NOAA Professional Paper NMFS 23 A guide to the corals of Alaska Robert P. Stone Stephen D. Cairns Dennis M. Opresko Gary C. Williams Michele M. Masuda December 2023 U.S. Department of Commerce NOAA Professional Gina M. Raimondo Secretary of Commerce Papers NMFS National Oceanic and Atmospheric Administration Richard W. Spinrad Scientific Editor Administrator José I. Castro National Marine Fisheries Service, NOAA National Marine 263 13th Avenue South Fisheries Service Saint Petersburg, Florida 33701 Janet Coit Assistant Administrator for Fisheries Associate Editor Cara Mayo National Marine Fisheries Service, NOAA Scientific Publications Office 7600 Sand Point Way NE Seattle, Washington 98115 Managing Editor Shelley Arenas National Marine Fisheries Service, NOAA Scientific Publications Office 7600 Sand Point Way NE Seattle, Washington 98115 The NOAA Professional Paper NMFS (ISSN 1931-4590) series is published by the Scientific Publications Office, National Marine Fisheries Service, NOAA, 7600 Sand Point Way NE, Seattle, WA 98115. The Secretary of Commerce has The NOAA Professional Paper NMFS series carries peer-reviewed, lengthy original determined that the publication of research reports, taxonomic keys, species synopses, flora and fauna studies, and data- this series is necessary in the transac- intensive reports on investigations in fishery science, engineering, and economics. tion of the public business required by Copies of the NOAA Professional Paper NMFS series are available free in limited law of this Department. Use of funds numbers to government agencies, both federal and state. They are also available in for printing of this series has been ap- exchange for other scientific and technical publications in the marine sciences. Profes- proved by the Director of the Office sional Papers are published online in PDF format at http://spo.nmfs.noaa.gov of Management and Budget. NOAA Professional Paper NMFS 23 A guide to the corals of Alaska Robert P. Stone (retired) (contact author)1 Stephen D. Cairns (retired)2 Dennis M. Opresko (retired)2 Gary C. Williams3 Michele M. Masuda1 1 Auke Bay Laboratories Alaska Fisheries Science Center National Marine Fisheries Service, NOAA 17109 Point Lena Loop Road Juneau, Alaska 99801 Present address for contact author: 3105 National Park Road Juneau, Alaska 99801 Email: bob.stone.coral@gmail.com 2 Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution P.O. Box 37012, MRC 163 Washington, D.C. 20013-7012 3 Department of Invertebrate Zoology and Geology California Academy of Sciences 875 Howard Street San Francisco, California 94103 December 2023 U.S. Department of Commerce Seattle, Washington Cover figure Top left: A school of yellowtail rockfish (Sebastes flavidus) feeding in a thicket of Primnoa pacifica in Dixon Entrance (eastern Gulf of Alaska) at a depth of 165 m. Top right: Coral habitat dominated by bubblegum corals (Paragorgia sp.) on the flank of the submarine volcano Amchixtam Chaxsxii (central Aleutian Islands) at a depth of 920 m. Bottom left: A Muriceides nigra colony highlights a coral garden on the flank of the Bobrof Volcano (central Aleutian Islands) at a depth of 160 m. Bottom right: Coral habitat dominated by hydrocorals (mostly Stylaster sp.) in northern Amchitka Pass (central Aleutian Islands) at a depth of 896 m. Suggested reference Stone, Robert P., Stephen D. Cairns, Dennis M. Opresko, Gary C. Williams, and Michele M. Masuda. 2023. A guide to the corals of Alaska. NOAA Professional Paper NMFS 23, 413 p. https://doi.org/10.7755/PP.23 Online dissemination This report is posted online in PDF format at https://spo.nmfs.noaa.gov (click on Professional Papers link). Copyright law Although the contents of the Professional Papers have not been copyrighted and may be reprinted entirely, reference to source is appreciated. Proprietary products The National Marine Fisheries Service (NMFS) does not approve, recommend, or endorse any proprietary product or proprietary material mentioned in this publi- cation. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or imply that NMFS approves, recommends, or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. Disclaimer The findings and conclusions in this report are those of the author(s) and do not necessarily represent the views of the National Marine Fisheries Service (NOAA). We dedicate this guide in memory of Bruce L. Wing 1938 – 2016 “Only as a biological oceanographer can one expect to work on whales, turtles, fish, zooplankton, phytoplankton, bacteria, water chemistry, ocean physics, [and deep-sea corals], sometimes all in one day.” CONTENTS Chapter 1 – Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 About this guide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Alaska – physiography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Eastern Gulf of Alaska . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Western Gulf of Alaska . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Gulf of Alaska Seamount Province . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Aleutian Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Bering Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Alaska Arctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Data sources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Alaska corals – inventory and systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Zoogeography of corals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Collection and preservation of specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Chapter 2 – Class Anthozoa, Subclass Hexacorallia, Order Antipatharia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Chapter 3 – Order Scleractinia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Chapter 4 – Order Zoantharia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 Chapter 5 – Subclass Octocorallia, Order Malacalcyonacea, Family Alcyoniidae . . . . . . . . . . . . . . . . . . . . . . . 88 Chapter 6 – Family Clavulariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104 Chapter 7 – Family Gorgoniidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111 Chapter 8 – Family Malacalcyonacea incertae sedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118 Chapter 9 – Family Paramuriceidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140 Chapter 10 – Order Scleralcyonacea, Family Chrysogorgiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150 Chapter 11 – Family Coralliidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167 Chapter 12 – Family Keratoisididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 200 Chapter 13 – Family Primnoidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233 Chapter 14 – Family Sarcodictyonidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292 Chapter 15 – Superfamily Pennatuloidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297 Chapter 16 – Class Hydrozoa, Order Anthoathecata, Family Stylasteridae . . . . . . . . . . . . . . . . . . . . . . . . . . . 341 Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 391 Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 392 Appendix I. Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401 Appendix II. Coral species reported from Alaska waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409 Appendix III. Additional supporting references on Alaska corals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413 v 1 Abstract—The Magnuson-Stevens CHAPTER 1 Fishery Conservation and Management Reauthorization Act of 2006 mandat- ed the research and management of the nation’s deep-sea coral resources through establishment of the National Oceanic and Atmospheric Administra- Introduction tion’s Deep Sea Coral Research and Technology Program. The challenge for Alaska was daunting, where expansive, world-class fisheries often coincided For more than a century Alaskan launched a new wave of explora- with extraordinarily rich coral habitats fishermen have inadvertently brought tion, including research using sub- for a high-latitude region. The first chal- corals to the sea surface tangled in mersibles and remotely operated ve- lenge was to inventory known locations nets and lines, and early on the cor- hicles to gather glimpses of Alaska’s of deep-sea corals. Many coral records and some museum collections existed als became symbolic of the rich coral habitats for the first time. Dis- from Alaska, but the taxonomy of cor- fauna and diverse communities in covery of vast coral resources in the als was little studied and field iden- deep marine waters there (Fig. 1-1). Aleutian Islands (Stone, 2006; Stone tification of corals was problematic. A few coral species from Alaska and Shotwell, 2007; Stone, 2014), Formal bycatch programs and research waters were described in the late Gulf of Alaska (GOA) Seamounts activities in recent decades provided many more specimens for taxonomic 1800s (Verrill, 1866; Dall, 1884), (Baco, 2007), and elsewhere in the study, but guides to species were largely but scientists paid little attention to United States (Lumsden et al., 2007) incomplete, inaccurate, and outdated the region’s coral resources until the greatly informed several important given the fast pace of species discovery U.S. Fisheries Steamer Albatross re- mandates of the Magnuson-Stevens in Alaska. We provide a comprehen- turned from Alaska with astonishing Fishery Conservation and Manage- sive, up-to-date guide, detailing 161 coral taxa identified from museum evidence of rich coral beds. The U.S. ment Reauthorization Act of 2006. collections, primary literature, and Fisheries Steamer Albatross expedi- The Reauthorization Act established video records. Each profile includes tions continued through 1906, and the National Oceanic and Atmo- a description, images for each taxon, collections made during that period spheric Administration’s Deep Sea taxonomic history, biology, ecology, prompted the first detailed work on Coral Research and Technology geographical distribution, and habitat, including depth distribution. Corals are Alaska octocorals (Nutting, 1912) Program with specific mandates to found in the six regions of Alaska but and hydrocorals (Fisher, 1938). With conduct research and manage deep- the coral fauna of the Aleutian Islands is specific regard to hydrocorals, Fisher sea coral resources. by far the most species rich. The state of (1938) noted “the North Pacific is One of the major objectives of taxonomy for some coral groups is ex- far richer in indigenous species than the Deep Sea Coral Research and cellent, while others require additional collections and more taxonomic work. the North Atlantic.” Opportunistic Technology Program was to inven- Construction of this guide resulted in and directed collections made dur- tory locations of deep-sea corals. In descriptions of several antipatharian ing expeditions in the latter 1900s Alaska, researchers mapped known species, published separately from provided for subsequent taxonomic locations of all coral records to this guide (Alternatipathes mirabilis, work on octocorals (Bayer, 1952a, identify “hot spots” or areas where Bathypathes alaskensis, B. ptiloides, B. tiburonae, and Parantipathes pluma) 1996; Cairns, 2011a), antipatharians corals appeared to be particularly and the scleractinian Flabellum (Flabel- (Opresko, 2005), stylasterids (Cairns diverse and/or abundant (see Stone lum) oclairi Cairns, sp. nov. described and Lindner, 2011), and a synthesis and Rooper, 2017). These locations herein. The guide provides informa- on scleractinians (Cairns, 1994). were prioritized for in situ explora- tion for targeting new collections and Despite early evidence that Alas- tion. Since bycatch in fisheries and identifying areas of high abundance and indicator species of vulnerable ka had surprisingly vast coral re- fisheries-independent surveys is a marine ecosystems. Stakeholders can sources for a high-latitude area of major source of information on the now more adequately assess Alaska’s the world, corals remained little location of coral fauna and a source coral resources and risks from natural more than curios until the U.S. of specimens for study, programs and anthropogenic stressors. Congress enacted amendments to were implemented to increase the the Magnuson-Stevens Fishery Con- awareness and ability of fisheries servation and Management Act in observers and scientists to collect 1996. Known as the Sustainable bycatch specimens for museum col- Fisheries Act, the legislation placed lections and taxonomic study (Stone emphasis on the impacts of fish- et al., 2015a). A few guides were eries on essential fish habitat and available for use in the field (Wing 2 Professional Paper NMFS 23 ion reference to “A guide to the deep-water sponges of the Aleutian Island Archipelago” (Stone et al., 2011) to assist in identifying the rich benthic fauna of Alaska. We provide detailed profiles for all coral taxa that we have identified from museum collections, the primary literature, and detailed analysis of high-confidence video records. Altogether, we present 15 coral chap- ters arranged systematically (Table 1): each of the three orders of Hexacorallia (Antipatharia, Scleractinia, and Zoantharia); each of the 10 families of Octocorallia Figure 1-1 (Alcyoniidae, Clavulariidae, Gorgoniidae, Malacalcyo- A canned sockeye salmon label (circa 1889) used by the nacea incertae sedis, Paramuriceidae, Chrysogorgiidae, Thlinket Packing Company located on Wrangell Island, Coralliidae, Keratoisididae, Primnoidae, and Sarcodi- Alaska during the earliest days of commercial fishing operations in the state. The label pays homage to the ctyonidae), given their relatively high species richness, fishermen’s familiarity with the thickets of red tree coral abundance, and ecological importance; the superfamily (Primnoa pacifica) on the nearby fishing grounds. Pennatuloidea; and the family Stylasteridae. Each coral chapter begins with a summary of taxo- nomic and systematic notes, biological and ecological information, and figure(s) illustrating basic morphologi- and Barnard, 2004; Clark1) and have been an impor- cal features. In Appendix I we provide a glossary of com- tant first step toward adequately monitoring coral by- mon terms subdivided by major taxonomic group (or- catch, but were quickly outdated given the fast pace der Antipatharia, order Scleractinia, order Zoantharia, of discovery in a largely unexplored region. The avail- subclass Octocorallia, superfamily Pennatuloidea, and able guides are largely incomplete, contain species that family Stylasteridae). We refer the reader to publications have never been confirmed to occur in Alaska waters, by Cairns and Kitahara (2012), Bayer et al. (1983), Wil- and use inaccurate taxonomic nomenclature. liams (1995), and Cairns (2011a) for additional keys, Here, we provide a comprehensive and up-to-date illustrations, and glossaries for morphological and ana- guide for the identification of Alaska corals. The main tomical terms applied to Scleractinia (azooxanthellate), purpose of this guide is to develop an awareness and ap- Octocorallia, Pennatuloidea, and Stylasteridae, respec- preciation of the importance of the coral fauna in Alas- tively. ka waters, where the diversity and abundance of corals We present scientific names using standard binomi- is extraordinary and bycatch in existing fisheries contin- nal nomenclature. The first name (always capitalized) ues to be a major concern for resource m anagers. This is the genus (e.g., Stylaster). The second name (never guide allows fisheries observers and scientists to identify capitalized) is the species (e.g., parageus). A few species corals so that more accurate data can be included in ex- have a third name (never capitalized) for a designated isting databases. The guide is also designed for use by subspecies (e.g., columbiensis), and a few others have scientists making observations of the fauna in situ with additional designations (never capitalized) for a variant submersibles, including remotely operated vehicles, au- (var.) or morph. Additionally, five species in this guide tonomous underwater vehicles, and towed camera sys- from the order Scleractinia have a designated subgenus tems (e.g., Goddard et al., 2016). These in situ observa- that is identified with parentheses and placed between tions are critical to identify areas of high abundance and the genus and species (e.g., Fungiacyathus (Bathyactis) the locations of indicator species of vulnerable marine marenzelleri). The authority or name(s) of the author(s) ecosystems such as coral gardens. We provide detailed of the species description (i.e., the person[s] who de- profiles for 161 coral taxa. scribed the species) and year of description follow the scientific name. Parentheses placed around the authority indicate there has been an accepted transfer to a differ- About this guide ent genus since the original description. Each profile begins with a description including 1) This guide brings together all known information for typical growth form(s), 2) branching patterns, 3) known every coral taxon in Alaska. It is intended as a compan- size range or maximum dimensions, 4) surface mor- phology, 5) axial skeletal and holdfast characteristics, 1Clark, R. N. 2006. Field guide to the benthic marine invertebrates 6) characteristics of the polyps and other surficial fea- of Alaska’s shelf and upper slope taken by NOAA/NMFS/AFSC/ RACE Division trawl surveys, 305 p. Unpubl. manuscript. Alaska tures, 7) fragility, and 8) color in life and with various Fish. Sci. Cent., Natl. Mar. Fish. Serv., 7600 Sand Point Way NE, preservation methods. For many taxa, examination of Seattle, WA 98115. these characteristics can provide a fairly accurate iden- Chapter 1 3 Table 1 Systematics for all coral taxa currently known to occur in Alaska waters. The taxonomic names are all modern, systematically arranged to family, and alphabetically arranged by genus and species within each family. Scientific names are presented using standard binominal nomenclature. The authority or name(s) of the author(s) of the species description (i.e., the person[s] who described the species) and year of description follow the scientific name. Parentheses placed around the authority indicate there has been an accepted transfer to genus assignment since the original description. We used the World Register of Marine Spe- cies as the taxonomic authority. PHYLUM CNIDARIA Subclass Octocorallia CLASS ANTHOZOA Order Malacalcyonacea Subclass Hexacorallia Family Alcyoniidae Order Antipatharia Alcyonium pacificum Yamada, 1950 Family Cladopathidae Alcyonium sp. Chrysopathes formosa Opresko, 2003 Gersemia fruticosa (Sars, 1860) Chrysopathes speciosa Opresko, 2003 Gersemia lambi Williams, 2013 Heteropathes pacifica (Opresko, 2005) Gersemia rubiformis (Ehrenberg, 1834) Trissopathes pseudotristicha Opresko, 2003 Gersemia sp. Family Schizopathidae Family Clavulariidae Alternatipathes mirabilis Opresko and Molodtsova, Clavularia armata Thomson, 1927 2021 Clavularia eburnea Kükenthal, 1906 Bathypathes alaskensis Opresko and Molodtsova, 2021 Clavularia rigida Broch, 1935 Bathypathes patula Brook, 1889 Clavularia sp. A Bathypathes ptiloides Opresko and Molodtsova, 2021 Family Gorgoniidae Bathypathes tiburonae Opresko and Molodtsova, 2021 Callistephanus pacificus Nutting, 1912 Bathypathes sp. A Callistephanus simplex (Nutting, 1909) Bathypathes sp. B Family Malacalcyonacea incertae sedis Dendrobathypathes boutillieri Opresko, 2005 Alaskagorgia aleutiana Sánchez and Cairns, 2004 Dendrobathypathes sp. Alaskagorgia splendicitrina Horvath and Stone, 2018 Lillipathes wingi Opresko, 2005 Calcigorgia beringi (Nutting, 1912) Lillipathes sp. A Calcigorgia gigantea Matsumoto et al., 2019 Parantipathes euantha (Pasternak, 1958) Calcigorgia japonica Dautova, 2007 Parantipathes pluma Opresko and Molodtsova, 2021 Calcigorgia matua Dautova, 2018 Parantipathes sp. A Calcigorgia spiculifera Broch, 1935 Order Scleractinia Cryogorgia koolsae Williams, 2005 Family Caryophylliidae Elenanthus cf. violaceus Caryophyllia (Caryophyllia) alaskensis Vaughan, 1941 Family Paramuriceidae Caryophyllia (Caryophyllia) arnoldi Vaughan, 1900 Acanthogorgia spissa Kükenthal, 1908 Crispatotrochus foxi (Durham and Barnard, 1952) Acanthogorgia sp. Desmophyllum dianthus (Esper, 1794) Muriceides cylindrica Nutting, 1912 Family Dendrophylliidae Muriceides nigra Nutting, 1912 Balanophyllia (Balanophyllia) elegans Verrill, 1864 Order Scleralcyonacea Family Flabellidae Family Chrysogorgiidae Flabellum (Flabellum) oclairi Cairns, sp. nov. Chrysogorgia sp. A Javania borealis Cairns, 1994 Chrysogorgia sp. B Javania cailleti (Duchassaing and Michelotti, 1864) Chrysogorgia sp. C Family Fungiacyathidae Pseudochrysogorgia sp. A Fungiacyathus (Bathyactis) marenzelleri (Vaughan, Pseudochrysogorgia sp. B 1906) Pseudochrysogorgia sp. C Family Micrabaciidae Radicipes stonei Cordeiro et al., 2017 Leptopenus discus Moseley, 1880 Family Coralliidae Order Zoantharia Hemicorallium sp. Family Epizoanthidae Paragorgia jamesi Herrera and Shank, 2016 Epizoanthus scotinus Wood, 1957 Paragorgia arborea var. pacifica Verrill, 1922 Family Parazoanthidae Paragorgia arborea var. pacifica morph nodosa Mesozoanthus sp. Paragorgia stephencairnsi Sánchez, 2005 Zibrowius cf. ammophilus Paragorgia sp. A gen. nov., sp. nov. Paragorgia sp. B Sibogagorgia cauliflora Herrera et al., 2010 Table continued 4 Professional Paper NMFS 23 Table 1 (continued) Subfamily Anthomastinae Family Anthoptilidae (cont.) Heteropolypus japonicus (Nutting, 1912) Anthoptilum sp. A Heteropolypus ritteri (Nutting, 1909) Anthoptilum sp. B Heteropolypus sp. Anthoptilum sp. C Pseudoanthomastus sp. Anthoptilum sp. D Family Keratoisididae Family Balticinidae Bathygorgia profunda Wright, 1885 Balticina californica (Moroff, 1902) Bathygorgia sp. A Balticina willemoesi (Kölliker, 1880) Isidella tentaculum Etnoyer, 2008 Balticina sp. A Isidella sp. A Family Kophobelemnidae Isidella sp. B Kophobelemnon sp. Keratoisis sp. A Family Pennatulidae Keratoisis sp. B Pennatula aculeata Danielssen, 1860 Keratoisis sp. C Pennatula sp. Keratoisis sp. D Ptilosarcus gurneyi (Gray, 1860) Keratoisis sp. E Family Protoptilidae Keratoisis sp. F Protoptilum sp. Keratoisis sp. G Family Stachyptilidae Orstomisis sp. Stachyptilum superbum Studer, 1894 Family Primnoidae Family Umbellulidae Arthrogorgia kinoshitai Bayer, 1952 Umbellula sp. Arthrogorgia otsukai Bayer, 1952 Family Veretillidae Arthrogorgia utinomii Bayer, 1996 Cavernularia vansyoci Williams, 2005 Callogorgia compressa (Verrill, 1865) Family Virgulariidae Callogorgia fraseri (Hickson, 1915) Virgularia bromleyi Kölliker, 1880 Calyptrophora laevispinosa Cairns, 2007 Virgularia cf. bromleyi Narella abyssalis Cairns and Baco, 2007 Virgularia cf. glacialis Narella alaskensis Cairns and Baco, 2007 CLASS HYDROZOA Narella arbuscula Cairns and Baco, 2007 Order Anthoathecata Narella bayeri Cairns and Baco, 2007 Family Stylasteridae Narella cristata Cairns and Baco, 2007 Crypthelia trophostega Fisher, 1938 Parastenella doederleini (Wright and Studer, 1889) Cyclohelia lamellata Cairns, 1991 Parastenella gymnogaster Cairns, 2007 Distichopora borealis Fisher, 1938 Parastenella ramosa (Studer, 1894) Errinopora dichotoma Lindner and Cairns, 2011 Plumarella aleutiana Cairns, 2011 Errinopora disticha Lindner and Cairns, 2011 Plumarella echinata Cairns, 2011 Errinopora fisheri Lindner and Cairns, 2011 Plumarella hapala Cairns, 2011 Errinopora nanneca Fisher, 1938 Plumarella nuttingi Cairns, 2011 Errinopora undulata Lindner and Cairns, 2011 Plumarella profunda Cairns, 2011 Errinopora zarhyncha Fisher, 1938 Plumarella robusta Cairns, 2011 Stylantheca papillosa (Dall, 1884) Plumarella spicata Nutting, 1912 Stylaster alaskanus Fisher, 1938 Plumarella superba (Nutting, 1912) Stylaster brochi (Fisher, 1938) Primnoa pacifica Kinoshita, 1907 Stylaster campylecus (Fisher, 1938) Primnoa pacifica var. willeyi (Hickson, 1915) Stylaster crassiseptum Cairns and Lindner, 2011 Primnoa wingi Cairns and Bayer, 2005 Stylaster elassotomus Fisher, 1938 Thouarella cristata Cairns, 2011 Stylaster leptostylus (Fisher, 1938) Thouarella trilineata Cairns, 2011 Stylaster parageus columbiensis Cairns and Lindner, Family Sarcodictyonidae 2011 Sarcodictyon incrustans (Broch, 1935) Stylaster parageus parageus (Fisher, 1938) Sarcodictyon sp. A Stylaster repandus Cairns and Lindner, 2011 Superfamily Pennatuloidea Stylaster stejnegeri (Fisher, 1938) Family Anthoptilidae Stylaster trachystomus (Fisher, 1938) Anthoptilum grandiflorum (Verrill, 1879) Stylaster venustus (Verrill, 1868) Anthoptilum murrayi Kölliker, 1880 Stylaster verrillii (Dall, 1884) Chapter 1 5 tification. Definitive identification of most species, how- region, the Pacific Plate moves roughly parallel to the ever, requires careful examination of the arrangement North American Plate along the Fairweather-Queen of microscopic skeletal structures, particularly the types, Charlotte Fault, forming an abrupt continental slope sizes, and locations of sclerites, and we provide that in- with an abbreviated shelf (von Huene, 1989). Promi- formation in each profile if available. nent geological features of the region include the Fair- For most taxa, we provide images of fresh or live weather Ground, a large offshore shoal on the outer specimens, preserved museum specimens, and specimens edge of the continental shelf in the northeast GOA, in situ. The former would be most useful to those identi- and the Shutter Ridge located 28 km west of Cape Om- fying specimens on the deck of fishing and survey vessels maney, Baranof Island, on the continental shelf. The while the latter would be useful to scientists attempting ridge comprises a series of rocky pinnacles along a sub- to identify and quantify coral fauna in situ with sub- duction zone where the Pacific Plate has docked with mersibles and remotely operated vehicles or from video Alaska (Greene2). footage collected in situ. We provide the known geo- graphic range and information on the physical habitat, Western Gulf of Alaska including depth, substrate, and associated fauna of each The western GOA region is delineated from longitude species both within Alaska waters and throughout its 146°W to longitude 163°W, near the east end of Uni- known range. The geographic range in Alaska for each mak Island, the first Island in the Aleutian Island Archi- taxon is depicted and finally, we include special remarks pelago (Fig. 1-2). The western GOA has a broad conti- with regard to taxonomic history, biology, and ecology. nental shelf extending seaward up to 200 km in some Appendix II provides a current and comprehen- areas and contains several deep troughs (NRC, 1990). sive taxonomic list of all coral taxa known to occur in The region includes smooth turbidity sediment scapes, Alaska. The list highlights the regions of occurrence and methane seeps, areas strongly influenced by glaciation, reported depth range for each taxon. Appendix III pro- and large basins with high rates of sedimentation (Reece vides a list of additional references not cited in the chap- et al., 2011). In this region the Pacific and North Ameri- ters that may provide the curious reader with valuable can plates slide, rather than slip, past each other and information on Alaska corals. form a convergent margin and subduction zone (von Huene, 1989). Alaska – physiography Gulf of Alaska Seamount Province Alaska forms a large portion of the northern margin of The GOASP is far offshore and encompasses a large the North Pacific Ocean and contains the majority of area from near the Aleutian Trench to the Juan de Fuca the U.S. marine coastline and continental shelf habitat. Ridge off Washington. The province contains over 100 The region has a highly varied submarine bathymetry, volcanic seamounts (>1000 m in height) arranged in owing to the numerous physical and geological process- two volcanic chains. The Kodiak–Bowie chain spans es at work in the three main physiographic provinces— 900 km from Kodiak Seamount in the western GOA to the GOA, the Bering Sea including the Aleutian Island Bowie Seamount off the Queen Charlotte Islands (Brit- Archipelago, and the Chukchi and Beaufort Seas in the ish Columbia) and contains 14 major and numerous Alaska Arctic. For the purposes of this guide, however, minor seamounts (Chaytor et al., 2007), most of which we delineate six distinct regions (Fig. 1-2) of Alaska were formed by movement of the Pacific Plate over the that have been used in previous reports on the distri- Bowie hot spot (Turner et al., 1973). The Cobb Sea- bution of marine resources (Stone and Shotwell, 2007; mount chain extends from Marchand Seamount in the Stone and Rooper, 2017) and more-or-less conform to northwestern GOA to Axial Seamount off the coast of resource management regions in the state. Those are 1) Oregon (Chaytor et al., 2007) where the Cobb hot spot the eastern GOA, 2) the western GOA, 3) the GOA Sea- is currently located on the Juan de Fuca Ridge (Desonie mount Province (GOASP), 4) the Aleutian Islands, 5) and Duncan, 1990). the Bering Sea, and 6) the Alaska Arctic. The 24 seamounts in Alaska waters span the entire GOA from Dickens Seamount in the southeastern sector Eastern Gulf of Alaska to Derickson Seamount in the western sector (Fig. 1-2). Sixteen of those seamounts summit within the range of The eastern GOA region is delineated by Dixon En- managed or targeted fish species and were designated trance on the state’s southern border with British Co- as Habitat Areas of Particular Concern in 2007 by the lumbia (Canada) and the line of longitude 146°W near the central part of Prince William Sound and includes 2Greene, G. 2014. Personal commun. Univ. Calif., 1850 Research the Inside Waters of Southeast Alaska (Fig. 1-2). In this Park Dr., Suite 300, Davis, CA 95618–6134. 6 Professional Paper NMFS 23 Figure 1-2 A map of the Alaska region showing the six broad geographical areas used in this guide: east- ern Gulf of Alaska (Region 1), western Gulf of Alaska (Region 2), Gulf of Alaska Seamount Province (Region 3), Aleutian Islands (Region 4), eastern Bering Sea (Region 5), and Alaska Arctic, including the Chukchi and Beaufort Seas (Region 6). These geographical areas were delineated for previous zoogeographical compilations on the distribution of marine resources (e.g., Stone and Rooper, 2017). North Pacific Fishery Management Council specifically insula to near the Kamchatka Peninsula in Russia and to protect deep-sea coral habitat from disturbance by is supported by the Aleutian Ridge, which largely sepa- fishing gear (Stone and Shotwell, 2007). The majority rates the deep North Pacific Ocean to the south and the of the seamounts have not yet been explored, but in this shallower Bering Sea to the north. The Aleutian Ridge guide we include coral records and video observations is a volcanic arc that contains more than 300 islands, made on nine of those seamounts. We also include re- including more than 20 active volcanoes. cords and observations from Pratt, Surveyor, and Mur- The region has frequent earthquakes and was formed ray Seamounts, which summit in international waters along zones of convergence between the North Ameri- but lie near the boundary with Alaska (Fig. 1-3). can Plate and other oceanic plates (Vallier et al., 1994). The island arc shelf is very narrow in the Aleutian Is- Aleutian Islands lands, drops precipitously on the Pacific side to depths The Aleutian Islands region is delineated by the line of greater than 6000 m in the Aleutian Trench, and con- longitude 163°W near the east end of Unimak Island to tains several large canyons, most notably, Adak and the international boundary with Russia west of Stale- Murray Canyons. A prominent feature of the Archi- mate Bank, and south to the boundary with interna- pelago is Bowers Ridge, which projects northward in a tional waters (Fig. 1-2). To the north, the region encom- 700-km arc from the central Aleutian Islands at Petrel passes all seafloor associated with the island arc slope Bank and along Bowers Bank to near the Ulm Plateau of the archipelago (Fig. 1-2). The Aleutian Island Ar- in the Aleutian Basin of the Bering Sea. Another notable chipelago extends over 1900 km from the Alaska Pen- geological feature of the region is Amchixtam Chaxsxii, Chapter 1 7 Figure 1-3 A map of the seamounts in the Gulf of Alaska Seamount Province where coral specimens and observations were collected: 1) Dickens Seamount, 2) Densen Seamount, 3) Welker Seamount, 4) Pratt Seamount, 5) Surveyor Seamount, 6) Quinn Seamount, 7) Giacomini Seamount, 8) Murray Seamount, 9) Patton Seamount, 10) Marchand Seamount, 11) Chirikof Seamount, and 12) Derickson Seamount. Historical (pre-1980) and recent research expeditions and Na- tional Marine Fisheries stock assessment surveys were a major source of coral specimens. Recent research expeditions were also an important source of video records. a nearly perfect conical submarine volcano discovered in ment deposited by the region’s major rivers (Johnson, northern Amchitka Pass in 2002 that is largely covered 2003). with the most lush and expansive coral gardens known in Alaska (Stone, 2006, 2014). Alaska Arctic The Alaska Arctic region is confined by the Bering Strait Bering Sea to the south, the boundary of territorial waters with The Bering Sea region is bounded by the Aleutian Is- Russia to the west, the boundary with international land Archipelago to the south, the Bering Strait to the waters to the north, and the line of longitude 141°W north, and the boundary of territorial waters with Rus- near the boundary with the Yukon Territory (Canada) sia to the west (Fig. 1-2). The Bering Sea is a shallow to the east (Fig. 1-2). The region contains large areas of sea with a massive continental shelf (1200 km long and the Chukchi and Beaufort Seas. The Chukchi Sea is a 500 km wide)—one of the largest continental shelves in relatively uniform, massive shallow shelf (only 20–60 the world (NRC, 1990). The continental shelf breaks at m deep) whereas the Beaufort Sea is a submarine ex- approximately 170 m depth. Seven major canyons incise tension of the North Slope coastal plain consisting of the continental slope, including the Zhemchug and Ber- a fairly broad (80–140 km wide) continental shelf and ing Canyons—the two largest submarine canyons in the a steep precipitous slope into the abyss of the Beaufort world (Johnson, 2003; Normark and Carlson, 2003). The Sea basin (Horowitz, 2002). Sediments on the continen- continental shelf and much of the slope contain limited tal shelf are predominantly soft and fine-grained and hard substrate but are rather heavily covered with sedi- are redistributed by longshore currents, wave action, 8 Professional Paper NMFS 23 entrainment in bottom-fast ice, ice gouging, ocean cur- Marine Fisheries Service stock assessment surveys, and rents, and internal waves (Horowitz, 2002). A promi- various fisheries, many coral records are available from nent geological feature of the Beaufort Sea is the Ste- past research expeditions made throughout Alaska. fansson Sound Boulder Patch, an area of isolated hard Many of these expeditions included video surveys of rock in an otherwise heavily sedimented region. the seafloor and the intentional collection of unknown specimens, particularly those that were common or abundant and presumably ecologically important, and Data sources conversely those that appeared to be quite uncommon and unusual. Detailed examination in the laboratory of All museum coral records from Alaska with appropriate video footage coupled with expertly identified voucher geospatial data were inventoried and inspected for data specimens provided a wealth of high-confidence identifi- quality. The majority of coral specimens collected in cations and ecological observations. Alaska are archived at the National Museum of Natural Previous research expeditions that contributed high- History (NMNH; Smithsonian Institution, Washington, confidence coral observations included those in 1) the D.C.) previously known as the U.S. Museum of Natural eastern GOA, Southeast Alaska, and Portland Canal History (USNM) and the California Academy of Sci- (Stone et al., 2013); 2) the eastern GOA continental ences (CAS; San Francisco, California). A few Alaska shelf and slope (Stone et al., 2015b); 3) the western records were obtained from the Royal British Columbia GOA continental shelf (Stone et al., 2005); 4) the central Museum (RBCM; Victoria, British Columbia, Canada), Aleutian Islands island arc shelf and slope (Stone, 2006, the Natural History Museum of London (NHMUK; 2014); and 5) the eastern Bering Sea shelf and slope London), and the Santa Barbara Museum of Natural (Miller et al., 2012). Additionally, to provide a wealth of History (SBMNH; Santa Barbara, California). We used new coral observations we carefully examined archived the World Register of Marine Species (WoRMS Edito- video footage from four expeditions on seamounts in rial Board, 2022) as the taxonomic authority and have the GOASP in 2002, 2004, and 2019. The vast video mostly adopted the recently revised systematics of the library maintained at the National Oceanic and Atmo- Octocorallia based largely on phylogenomics (McFad- spheric Administration Auke Bay Laboratories was also den et al., 2022). examined for high-confidence coral records, as were the The majority of Alaska museum specimens have been detailed observations made by the senior author during collected during National Marine Fisheries Service stock assessment surveys, using longlines (Auke Bay Labora- more than 2000 scuba dives throughout much of Alaska tories) and trawls (Resource Assessment and Conser- between 1987 and 2017. vation Engineering Division’s Groundfish Assessment Program), and typically the location and depth are re- corded as either the start or end point of the longline Alaska corals – inventory and systematics set or trawl haul. For both types of surveys, the start and end locations may vary by >10 km and the depth Our comprehensive inventory indicates there are cur- may span up to 400 m. To minimize the error associated rently 161 distinct coral taxa known in Alaska (Table with these records, we averaged the start and end values 1). The inventory includes two subclasses (Hexacorallia for both location and depth and present mean values. In and Octocorallia) of Anthozoa and a single order (An- no case did this method establish a new depth range for thoathecata) of Hydrozoa. The Hexacorallia includes any species. the orders Antipatharia, Scleractinia, and Zoantharia. Both the Auke Bay Laboratories and Resource As- The Antipatharia includes two families: Cladopathidae sessment and Conservation Engineering Division’s stock (four taxa) and Schizopathidae (14 taxa). The Sclerac- assessment programs maintain databases of coral by- tinia includes five families: Caryophylliidae (four taxa), catch records. However, because specimens are only re- Dendrophylliidae (one taxon), Flabellidae (three taxa tained upon special request and field identifications are including Flabellum (Flabellum) oclairi Cairns, sp. typically made only at higher taxonomic levels, we did nov.), Fungiacyathidae (one taxon), and Micrabaci- not consider them in our analysis. Curious fishermen idae (one taxon). The Zoantharia includes two families: and recreationists also occasionally retain coral speci- Epizoanthidae (one species) and Parazoanthidae (three mens and donate them to laboratories for identification. species). Many of those specimens have proved valuable since The Octocorallia includes two orders (Malacalcyo- they are often collected in areas not typically surveyed nacea and Scleralcyonacea) and is far more speciose by the other methods. than the Hexacorallia. The Malacalcyonacea includes In addition to historical collections like those from the five families: Alcyoniidae (six taxa), Clavulariidae (four U.S. Fisheries Steamer Albatross expeditions, National taxa), Gorgoniidae (two taxa), Malacalcyonacea incer- Chapter 1 9 tae sedis (nine taxa), and Paramuriceidae (four taxa). Clavulariidae, and Sarcodictyonidae (each family has The Scleralcyonacea includes five families: Chrysogorgi- only one taxon with incomplete taxonomy). Contrast- idae (seven taxa), Coralliidae (12 taxa), Keratoisididae ingly, the state of the taxonomy for Chrysogorgiidae, (13 taxa), Primnoidae (27 taxa), and Sarcodictyonidae Coralliidae, and Keratoisididae is poor, with the major- (two taxa). The Scleralcyonacea also includes the super- ity of taxa (85%) in need of additional work or col- family Pennatuloidea that is composed of nine families: lection. The taxonomy for the family Malacalcyonacea Anthoptilidae (six taxa), Balticinidae (three taxa), Ko- incertae sedis includes nine taxa in Alaska waters whose phobelemnidae (one taxon), Pennatulidae (three taxa), phylogentic positions are still unknown or uncertain (in- Protoptilidae (one taxon), Stachyptilidae (one taxon), certae sedis) above the genus level but is well established Umbellulidae (one taxon), Veretillidae (one taxon), and to species for eight of the nine taxa recently placed in Virgulariidae (three taxa). the group by McFadden et al. (2022). The state of tax- Corals in the class Hydrozoa are represented only onomy for Pennatuloidea is fair to poor. About 50% of by the order Anthoathecata with 23 taxa in the family the listed taxa have incomplete taxonomy that may re- Stylasteridae. quire higher-level taxonomic revision. Finally, the state Note that not all taxa identified in this guide have of taxonomy for Stylasteridae is excellent. been confirmed by examination of microscopic features and/or molecular analyses; those taxa are listed generi- cally until additional taxonomic work can be completed Zoogeography of corals (Table 1). Twenty taxa listed in Table 1 have been identified Corals have been documented in all six regions of to genus based on examination of microscopic features Alaska but the number of taxa and the representation and/or molecular analyses but have not yet been de- of corals from the six major coral groups (antipath- scribed to species level. Twenty-two taxa in Table 1 have arians, scleractinians, zoantharians, octocorals, pen- been confidently assigned to genus based on gross mor- natuloideans, and stylasterids) varies greatly between phological features and ecology, and are now queued regions. Three regions (eastern GOA, western GOA, for sclerite examination, molecular analyses, and for- and the Aleutian Islands) support corals from all ma- mal description if applicable. We have identified three jor coral groups, while stylasterids have not been re- taxa to genus based solely on the information available ported from the GOASP or the Bering Sea regions; and from videography and associated ecology. Finally, three the Alaska Arctic region is represented only by a single taxa are listed as “cf.,” meaning “confer or compare species of octocoral. The Aleutian Islands region is the to.” That is, specimens may be the species “compared most species rich with 104 reported taxa, followed by to” but appear sufficiently different enough that closer the GOASP (57 taxa), the eastern GOA (44 taxa), the examination is warranted. For example, we list “Vir- western GOA (31 taxa), the Bering Sea (21 taxa), and gularia cf. bromleyi” to mean the taxon is very similar the Alaska Arctic (one taxon). A summary for each of to “Virgularia bromleyi” but should be more carefully the six regions of Alaska follows. compared to known material for verification as it may Forty-four coral taxa from the six major coral groups represent an unknown or undescribed taxon. Altogeth- are reported from the eastern GOA region, including six er, 45 taxa are listed in Table 1 for which museum speci- antipatharians, four scleractinians, two zoantharians, mens are available for future taxonomic study. 15 octocorals, nine pennatuloideans, and eight stylaster- The state of taxonomy for Antipatharia is good with ids (Appendix II). Eleven taxa from the region are found only four taxa (Bathypathes sp. A, Bathypathes sp. B, nowhere else in Alaska, and of those, the eight taxa with Lillipathes sp. A, and Parantipathes sp. A) in need of complete taxonomy are found or suspected to occur fur- additional work, and the collection of Dendrobathy- ther south along the west coast of North America, indi- pathes sp. known only from video records is a high pri- cating a strong taxonomic affinity with the coral fauna ority. The taxonomy of all known Alaska Scleractinia of that region (Boutillier et al., 2019). is now complete with the description of Flabellum (Fla- Thirty-one coral taxa from the six major coral groups bellum) oclairi sp. nov. included in this guide. The state are reported from the western GOA region, including of taxonomy for Zoantharia is poor, but promising three antipatharians, three scleractinians, one zoanthar- leads using molecular genetics analyses are currently ian, 13 octocorals, seven pennatuloideans, and four underway. stylasterids (Appendix II). Four taxa known from the The state of the taxonomy for Octocorallia (exclud- region are found nowhere else in Alaska, and of those, ing Pennatuloidea) is fair (65% of the 86 taxa have the three taxa with complete taxonomy are found else- complete taxonomy) but varies greatly among families. where in the North Pacific Ocean. The state of taxonomy is complete for Primnoidae and Fifty-seven coral taxa from five major coral groups Gorgoniidae and nearly complete for Paramuriceidae, are reported from the GOASP region, including 12 an- 10 Professional Paper NMFS 23 tipatharians, three scleractinians, one zoantharian, 37 ing different from the 161 taxa presented in this guide. octocorals, and four pennatuloideans (Appendix II). This guide can be used to facilitate collection of the first Stylasterids have not been reported from the seamounts two priority categories by providing the known loca- region. Thirty-one (54%) of the taxa known from the tions of those taxa. Additionally, we recommend for region are found nowhere else in Alaska, indicating the collection those specimens that appear to be of known seamounts likely have a high degree of endemism. Of taxa but are evidently a geographic or bathymetric the 31 taxa, only 10 have complete taxonomy, which range extension for that species. Other high-priority is not surprising since the region has only recently been collections include those taxa for which museum col- the subject of exploration and specimen collection. Sev- lections are quantitatively or qualitatively inadequate to en of the 10 taxa with complete taxonomy appear to provide comprehensive species descriptions and to sup- be endemic to Alaska seamounts; that number is likely port taxonomy and systematics work using molecular to rise significantly as museum specimens are critically genetics. For example, some taxa are known only from examined and additional collections are made. one or few specimens; others from incomplete, frag- A total of 104 coral taxa from the six major coral mented specimens; and still others from poorly or inap- groups are reported from the Aleutian Islands region, propriately preserved specimens. including five antipatharians, six scleractinians, one The collection protocols outlined by Etnoyer et al. zoantharian, 58 octocorals, 13 pennatuloideans, and (2006) for deep-water corals are suitable for the collec- 19 stylasterids (Appendix II). Sixty-two (59%) of the tion of all Alaska corals. However, through the process taxa known from the region are found nowhere else in of constructing this guide, we have noted protocols Alaska, and of those, 50 taxa (81%) have complete tax- that have been performed inadequately or inconsis- onomy. Thirty-four of the 50 taxa (68%) with complete tently (and consequently have been problematic) so we taxonomy appear to be endemic to the region, indicat- provide the preferred protocols again here with slight ing that the archipelago has a high degree of endemism. modification. Of the remaining 16 non-endemic taxa, 11 are known Shortly after collection, specimens should be pho- only from Russian and/or Japanese waters, indicating tographed on a solid, clean, and dry background with a strong taxonomic affinity with the coral fauna of the (and without) an appropriate scale and specimen identi- Northwest Pacific Ocean. fier label (equivalent to “field number” in the USNM Only 21 coral taxa from five major coral groups database). This label will ultimately link the specimen are reported from the Bering Sea region, including two with metadata that importantly must include the follow- antipatharians, one scleractinian, one zoantharian, 11 ing station data: 1) field number, 2) date of collection, 3) octocorals, and six pennatuloideans (Appendix II). station number (station, haul, set or dive number, etc.), Stylasterids have not been reported from the Bering Sea. 4) vessel name, 5) name of collector, 6) collection meth- No species are endemic to the region as all known taxa od, 7) latitude and longitude, and 8) depth. occur in other regions of Alaska. When collecting specimens, imagery (still photos Only a single species of octocoral, the sea strawber- and/or video) should include the subject and surround- ry (Gersemia rubiformis) (Ehrenberg, 1834), is known ing habitat from multiple perspectives and with various from the Alaska Arctic region where it is abundant in lighting options. Imagery of the subject should also in- some areas (Table 1). Although this region of Alaska clude the nearby substrate, if possible, and the projection has been the focus of new research programs during the of scaling lasers on a flat surface (preferably on the sub- past decade, it is still by far the least explored region ject) so the area of view and size of objects are measured particularly with regard to coral habitat. There are sev- accurately. Broad-scale imagery can provide important eral museum specimens of indeterminate or question- habitat information regarding patch density and associ- able taxonomy that need attention so a few taxa may ated fauna. Close-up imagery can provide valuable infor- be added in the future to the inventory, but the region is mation on the presence of microfauna that may not be clearly depauperate in corals. retained with collected specimens and diagnostic features of the specimen, including polyps in both extended and retracted states. Documentation of polyp retraction can Collection and preservation of specimens easily be achieved by gently agitating the colony with the submersible’s manipulator. In each chapter summary, and in some species profiles, Sampling of a branch or small branch complex is we identify specific taxa as high priorities for future generally adequate for taxonomic identification, but collection. The highest priorities for collection are taxa if the specimen is suspected of being an unknown or with incomplete taxonomy, taxa identified solely on high-priority taxon (i.e., one that might be designated photographic evidence (of which there are three), and as a type), then the whole specimen including the hold- any specimens that are unfamiliar and suspected of be- fast should be collected if possible. If the whole speci- Chapter 1 11 men is difficult to detach from the substrate, then con- purposes may be fixed in a buffered 5% formaldehyde sider collecting the substrate (cobble or siltstone) with solution for 24 hours prior to storage in ethanol solu- the whole specimen attached. Save the substrate appro- tion. Specimens to be used for molecular genetic analy- priately; a geologist may be able to provide important ses should be frozen (preferably to −10°C or colder) or geological habitat information as a bonus. If specimens preserved by other methods (e.g., smeared on Whatman are small and plentiful, consider collecting several as FTA cards [GE Healthcare Lifesciences, Chicago, IL] species descriptions based on more than one speci- or placed in RNALater [Ambion, Inc., Austin, TX]) for men are most valuable. When sampling bamboo corals restriction-site associated DNA sequencing. For muse- (family Keratoisididae), attempt to collect branch com- um archiving, specimens should be stored in an 80–90% plexes consisting of both nodes and internodes since ethanol solution. Very large specimens may be dried but branching pattern is a valuable diagnostic characteris- a fragment of the specimen should be stored in ethanol tic for some taxa. solution. Great care should be taken to ensure that the actual Researchers responsible for the collected specimens process of specimen collection and securing specimens should ensure that the properly preserved specimens are in payload containers is carefully video-documented. transferred as soon as practical to the museum where Because multiple specimens are often stowed in the same they will be permanently archived. Each specimen containers, scientists often need to review imagery of the should be accompanied with all appropriate metadata collections to accurately sort and label specimens, ensur- and photographs. If the specimen has been partitioned, ing that they are linked to the proper station data. Speci- then the record should indicate the repository and cus- mens are occasionally not secured and properly stowed, todian of the sample, especially if it is the parent speci- only to be lost before arriving on deck. At a minimum, men. Finally, the principal researcher or assistant should the proper and secure storage of each specimen should verify that the specimen records entered into archived be documented with video imagery. databases are accurate and complete. Approximately All coral specimens may be frozen (preferably at tem- 5–10% of the thousands of data records we examined peratures below −10°C) but ideally stored in an 80–90% while constructing this guide contained inaccurate or in- ethanol solution. Specimens to be used for histological complete data. 12 CHAPTER 2 Class Anthozoa Subclass Hexacorallia Order Antipatharia Corals in the order Antipatharia are referred to as black or thorny corals, principally due to the color of their hard proteinaceous axial skeleton and the presence of thorns on the skeletal surface. Skeletons are covered with a thin, fragile coenenchyme bearing small non- retractable polyps with six non-retractile tentacles. Currently there are seven families in the order. Fami- lies are differentiated based on the internal and external morphology of the polyps (Fig. 2-1), and on the mor- phology of the spines (Fig. 2-2). Genera are recognized primarily by the morphology of the corallum, and spe- cies are separated by differences in the size of the polyps and spines and minor alterations in the growth pattern of the corallum (i.e., size and density of primary pin- nules; and number, size, and arrangement of subpin- nules, if present; Fig. 2-3). The current state of taxonomy for the group is good, but is likely to undergo revision as more DNA data be- Figure 2-1 come available to help define the limits of species vari- Characteristic features of polyps in the order Antipatharia. ability. For some genera, such as Bathypathes, the DNA results so far have revealed the possible existence of more species than that suggested by morphology alone. Figure 2-2 Characteristic morphology of skeletal spines in corals in the order Antipatharia. Most spe- cies found in Alaska waters are more similar to the illustration on the left. Chapter 2 13 Figure 2-3 Illustrations of pinnulation patterns typical of genera in the order Antipatharia found in Alaska waters. (A) Chrysopathes: view from a distal branch end showing a single set of pinnules, one from each of six axial rows, with two simple posterior pinnules, two lateral pinnules with secondary pinnules, and two anterior pinnules with secondary and tertiary pinnules. (B) Heteropathes: a lateral view of the front of the corallum showing two rows of simple bilateral pinnules and numerous small anterior pinnules with subpinnules. The pinnules are obscured by soft tissue. (C) Trissopathes: view from a distal branch end showing a single set of pinnules, one from each of four axial rows, with two simple lateral pinnules and two anterior pinnules, each with two subop- posite secondary pinnules. (D) Alternatipathes: a lateral view of the front of the corallum with simple alternating pinnules in two lateral rows. (E) Bathypathes: an oblique lateral view of the front of the corallum with simple pinnules arranged bilaterally and suboppositely in two rows. (F) Dendrobathypathes: a lateral view of the front of the corallum. Primary pinnules are bilateral and alternating in two rows. A few primary pinnules have a single secondary pinnule. (G) Lillipathes: a lateral view of the front of the corallum. The pinnules are simple, in four rows, and in bilateral alternating groups of two each. (H) Parantipathes: a lateral view of the front of the corallum. The pinnules are simple, in up to eight axial rows, and in irregular but generally bilateral alternating groups of three or four each. 14 Professional Paper NMFS 23 Figure 2-4 A map of the North Pacific Ocean showing the distribution of corals in the order Antipath- aria () in Alaska waters. Here we list eight previously known species and five and Parantipathes pluma Opresko and Molodtsova, new species that have recently been described due in 2021, found as far north as Zhemchug Canyon in the large part to an effort to make this guide as complete eastern Bering Sea, is one of the northernmost corals in and comprehensive as possible. Additionally we list five the North Pacific Ocean. taxa (Bathypathes sp. A, Bathypathes sp. B, Dendro- Antipatharians are widespread in Alaska waters, oc- bathypathes sp., Lillipathes sp. A, and Parantipathes sp. curring in all regions except the Arctic (Fig. 2-4), and A) that have not yet been formally described; although they are particularly abundant on seamounts in the Gulf these forms appear to be morphologically unique, ad- of Alaska Seamount Province. They are deep-water fau- ditional specimens and DNA analysis are needed to de- na in Alaska and occur at depths between 329 and 4950 termine if they represent different species or are only m. They typically grow on firm rock but are often found variants of known species. on siltstone, particularly on the continental slope in the The antipatharian fauna of the Northeast Pacific eastern Gulf of Alaska. Some species can grow to more Ocean has only been extensively described in the past than a meter high and wide and these larger colonies few decades (Opresko, 2003, 2005; Molodtsova and often harbor swarms of euphausiids and other fauna. Opresko, 2017; Opresko and Molodtsova, 2021). Based Colonies of most species are found either singly or in on those studies and more recently collected material, 18 low-density fields. They are a fairly common bycatch taxa from two families (Cladopathidae and Schizopathi- item in longline fisheries and stock assessment surveys dae) are currently known to occur in Alaska waters. and, given their very strong axes, are most typically dis- They are found in all major regions except the Arctic, lodged whole with the holdfast. Chapter 2 15 Family Cladopathidae 1. Chrysopathes formosa Opresko, 2003 Description (Adapted from Opresko, 2003) Colonies lower parts of the pinnules, but becoming more inclined (i.e., coralla) are branched primarily in a single plane upward on the distal sections. Spines are generally less with five or more orders of branches. Colonies are up to than 0.12 mm in height on the primary pinnules, slight- 53 cm in height, at least 32 cm in width, and 4 mm in ly taller on the secondaries, and shorter on the main basal stem diameter. The stem and branches are pinnu- branch and larger branches. late. Secondary pinnules are present on some primaries; Polyps have six primary mesenteries and no second- tertiary pinnules are absent. Over most of the corallum, ary mesenteries. Polyps are elongated transversely in the primary pinnules are arranged in six rows and in alter- direction of the axis; 1.8–3.0 mm in transverse diameter nating biserial groups of three pinnules; each group con- (as measured from the outer edge of the proximal lateral sists of one anterolateral, one lateral (or posterolateral), tentacles to the distal edge of the distal lateral tentacles); and one posterior pinnule. Near the distal end of the and are arranged in a single row on one side of the pin- branches there can be only four rows of primary pin- nules and subpinnules. nules. Lateral primary pinnules are up to 2 cm in length, Holdfasts are quite large but thin, asymmetrical, and simple or occasionally with a single secondary pinnule; fairly easily detached. Color of flesh in life is creamy anterolateral primary pinnules are generally less than white; gold to pale yellow when frozen or dried. 1.5 cm in length, simple or with one or two secondary pinnules arising near the base. Posterior primary pin- Remarks This species is very similar to Chrysopathes nules are usually not more than 0.5 cm in length and speciosa Opresko, 2003 with which it co-occurs, but dif- without subpinnules. Primary pinnules are spaced 3–4 fers in having fewer secondary pinnules. Secondaries are mm apart in each row, with 15–18 pinnules/cm. rare on the lateral primary pinnules and there are no ter- Pinnular spines are triangular to conical in lateral tiary pinnules. Spines are not as strongly inclined distally, view, acute, directed out almost horizontally on the the primary pinnules are spaced further apart, and the polyps are slightly smaller than those in C. speciosa. This species often har- bors ophiuroid and hippo- lytid shrimp associates. Distribution Locally com- mon. In Alaska – eastern Gulf of Alaska continental slope (Fig. 2-5). Elsewhere – known only from a few locations off the west coast of the Americas (Jasper Sea- mount off Baja California, Mexico to Ecuador). Habitat In Alaska – grows predominantly on siltstone but occasionally on bed- rock, cobbles, and pebbles in deeper areas of the con- tinental slope at depths be- tween 619 and 756 m and often in low density patches Figure 2-5 with the congener C. spe- ciosa. Elsewhere – depths A map of the eastern Gulf of Alaska showing the distribution of Chrysopathes formosa () in the Alaska waters. between 700 and 895 m. 16 Professional Paper NMFS 23 Photos A) A preserved (frozen then dried) whole The distance between the red laser marks is 10 cm. C. formosa colony (USNM 1484090) collected in the E) A patch of Chrysopathes spp. on a bedrock ledge eastern Gulf of Alaska at a depth of 756 m. B) A in the eastern Gulf of Alaska at a depth of 715 m. A close-up view of the same specimen in photo A. C) A giant grenadier (Albatrossia pectoralis) takes cover in close-up view of the holdfast of the same specimen in the patch of corals. The distance between the red laser photo A. D) A C. formosa colony (USNM 1288460) marks is 20 cm. in the eastern Gulf of Alaska at a depth of 619 m. Chapter 2 17 2. Chrysopathes speciosa Opresko, 2003 Description (Adapted from Opresko, 2003) Colonies to the direction of the primary pinnule, and projecting (i.e., coralla) are branched primarily in a single plane laterally or somewhat distally or basally. Tertiary pin- but sometimes bushy. Colonies are to at least 55 cm in nules are present on some secondaries. height and at least 45 cm in width, and up to 6 mm in Spines on pinnules are simple, smooth, conical, acute, basal stem diameter. Stem and branches are pinnulate. subequal, and are usually directed or curved distally, es- Primary pinnules are arranged equidistantly around the pecially towards the distal end of the pinnules. Spines axis in four rows near the tips of branches, increasing are generally less than 0.12 mm in height (maximum to six rows on lower sections of branchlets, and in al- 0.18 mm in height) on the primary pinnules and slightly ternating bilateral groups of two or three pinnules, one shorter on the secondaries. Spines on branches and the from each row; 18–27 pinnules/cm for all rows. Pri- stem are ≤0.05 mm. mary pinnules are usually not more than about 1 cm Polyps have six primary mesenteries and no second- in length, subequal or with laterals slightly longer than ary mesenteries. Polyps are elongated transversely in the anterior and posterior ones. Primary pinnules near tips direction of the axis; transverse diameter is 2.0–3.3 mm of branches are mostly simple or with one secondary (including tentacles) and arranged in a single row on pinnule on anterior primaries only; becoming increas- one side of the pinnules and subpinnules. ingly subpinnulate on the lower parts of the branches, Holdfasts are quite large but thin, asymmetrical, and with one or more secondary pinnules occurring on fairly easily detached. Color of flesh in life is creamy anterior and lateral primary pinnules, but only infre- white; gold to pale yellow in frozen or dried specimens. quently on posterior primary pinnules. There are up to four secondary pinnules on some primary pinnules. Ar- Remarks This species is very similar to Chrysopathes rangement and spacing of secondary pinnules are highly formosa with which it co-occurs but differs in having variable, either alternate, uniserial, subopposite, or ir- more numerous subpinnules on the anterolateral pri- regular. Secondary pinnules are inclined distally relative maries (Fig. 2-3A). Secondaries are not uncommon on the lateral primary pinnules and tertiary pinnules are present. Spines are more typically inclined distally, the primary pinnules are spaced closer together, and the polyps are slightly larg- er in transverse diameter than in C. formosa. Colo- nies often host ophiuroid and hippolytid shrimp as- sociates. Distribution Locally com- mon. In Alaska – eastern Gulf of Alaska continental slope (Fig. 2-6). Elsewhere – known only from a few locations off the west coast of North America from northern British Columbia to California. Habitat In Alaska – grows predominantly on siltstone Figure 2-6 but occasionally on bed- A map of the eastern Gulf of Alaska showing the distribution of Chrysopathes speciosa rock, cobbles, and pebbles () in Alaska waters. in deeper areas on the con- tinental slope at depths 18 Professional Paper NMFS 23 between 648 and 914 m and often in low-density patches Photos A) A preserved (frozen then dried) whole C. with the congener C. formosa. Elsewhere – depths be- speciosa specimen (USNM 1014116) collected in the tween 732 and 1168 m. eastern Gulf of Alaska at a depth of 812 m. B) A close- up view of the same specimen in photo A. C) A close-up view of the stalk of the same specimen in photo A. Chapter 2 19 3. Heteropathes pacifica (Opresko, 2005) Description (Adapted from Opresko, 2005) The cor- (from the distal edge of distal lateral tentacles to the prox- allum is monopodial with the stem extending to the top imal edge of the proximal lateral tentacles); arranged in of the colony, and pinnulate. Colonies are to at least 32 one row, with 2–3 polyps/cm. Polyps on anterior pin- cm in height and at least 19 cm in width. Pinnules are nules and subpinnules are highly modified, elongate and arranged along the stem in two lateral rows of long pin- tear-drop in shape, with the proximal section usually nules, one on each side, and also in one to two irregular wider than the distal portion. The mouth is offset to- anterior rows of very short pinnules (Fig. 2-3B). Lateral wards the distal end of the polyp. Tentacles are absent pinnules are simple (without subpinnules), ≥5.5 cm in or reduced to just one pair. length, arranged alternately, and inclined and curved The holdfast is relatively large but thin, asymmetrical, distally. Adjacent lateral pinnules are on the same side and not particularly strong. Color of the coenenchyme of the axis about 5 mm apart, resulting in 3 pinnules/ in life is orange (a darker orange than Alternatipathes cm. Adjacent lateral pinnules are on opposite sides of mirabilis Opresko and Molodtsova, 2021 and Bathy- the stem 2.0–2.5 mm apart. Anterior primary pinnules pathes ptiloides Opresko and Molodtsova, 2021 with are short, <1 cm, and subpinnulate with 1–4 secondary which it co-occurs) and lighter orange in ethanol. pinnules. Anterior pinnules are spaced 1.5–2.0 mm apart irregularly, with 8–10 occurring along 1 cm of the axis. Remarks This species was originally described as He- Secondary pinnules are equal in length to the anterior liopathes pacifica Opresko, 2005. primary pinnules. There are up to four secondary pin- nules on one anterior primary; the two lowermost ones are almost subopposite (about 0.2 mm apart) on lateral Distribution Rare. In Alaska – known only from De- sides of the primary and at right angles to the direction rickson Seamount in the western Gulf of Alaska (Fig. of stem. There are two distal secondaries (about 0.5 2-7). Elsewhere – similar morphotypes (Heteropathes cf. mm apart) on the upper and lower sides of the primary. pacifica) have been recorded from Gorda Ridge off Or- Secondary pinnules are curved distally to become nearly egon (USNM 1234550) and from New Zealand (USNM parallel to the anterior primary from which they origi- 1527070). nate. Secondary pinnules are sometimes subpinnulate, with tertiary pinnules occurring on the upper and lower Habitat In Alaska – grows on siltstone and fragment- sides. Anterior primary pinnules and subpinnules extend ed basalt at depths between 3563 and 4511 m but ob- out nearly perpendicular to the plane containing the stem served on video footage to depths of 4663 m (senior au- and lateral pinnules. thor, personal observ.). Elsewhere – morphotypes found Spines on the lateral pinnules are small (0.03–0.06 at depths between 1621 and 3030 m. mm in height from the center of the base to the tip), triangular, compressed, and nearly at right angles to the Photos A) The preserved (in ethanol) H. pacifica ho- axis. Four or five rows are visible in one lateral view. lotype (USNM 1070758; whole colony) collected on Spacing of spines is highly variable, but is typically Derickson Seamount in the western Gulf of Alaska at 0.32–0.37 mm, resulting in about 4 spines/mm. Spines a depth of 3563 m. B) A close-up dorsal view of the on the anterior primary pinnules are similar in size and same specimen in photo A. C) A close-up ventral view shape to those on the lateral primary pinnules. Spines on of the same specimen in photo A. D) The same colo- distal portions of some subpinnules are slightly larger ny in photo A at the time of collection in the western (up to 0.07 mm), more acute, and more distally directed Gulf of Alaska. E) A H. pacifica colony (indicated by than those on primary pinnules. the white circle) observed on Derickson Seamount at Polyps have six primary mesenteries and no second- a depth of 4663 m. F) An alternate view of the same ary mesenteries. Polyps on lateral pinnules are mostly colony in photo E. The distance between the red laser 4.5–5.2 mm (range 3.9–5.5 mm) in transverse diameter marks in photos D–F is 10 cm. 20 Professional Paper NMFS 23 3. Heteropathes pacifica (Opresko, 2005) (continued) Chapter 2 21 Figure 2-7 A map of the North Pacific Ocean showing the distribution of Heteropathes pacifica () and Trissopathes pseudotristicha (◆) in Alaska waters. 22 Professional Paper NMFS 23 4. Trissopathes pseudotristicha Opresko, 2003 Description (Adapted from Opresko, 2003) Coralla The color of the coenenchyme in life is bright to rusty are branched in a single plane to the fourth order or orange and creamy white in ethanol. more. Colonies are to at least 34 cm in height and at least 40 cm in width. The stem and branches are pinnu- Remarks This species is very similar to Trissopathes late and subpinnulate. Primary pinnules are arranged in tetracrada Opresko, 2003 but with a greater number of four rows: two anterior (or anterolateral) rows of simple secondary pinnules. Trissopathes tetracrada is known pinnules and two posterolateral rows of subpinnulate primarily from the Southwest Pacific (near New Zea- primary pinnules (Fig. 2-3C). Primary pinnules are also land, Tasmania, and south of Australia), with a single arranged alternately in bilateral groups containing one record from the North Atlantic (Cape Verde Islands). anterior and one posterolateral pinnule. Posterolateral primary pinnules are usually simple, mostly 1–2 cm in Distribution Uncommon. In Alaska – Gulf of Alaska length (up to 2.6 cm), and directed distally. Anterior pri- Seamount Province (Welker and Pratt Seamounts) and mary pinnules are 0.5–1.0 cm in length, directed nearly the central Aleutian Islands (Fig. 2-7). Elsewhere – wide at right angles to the direction of the branch, and usually ranging; known from a few locations off the west coast with a single subopposite pair of secondary pinnules oc- of North America south to Fieberling Guyot (west of curring near the base of the primary pinnule (Fig. 2-3C). the Channel Islands) and the Hawaiian Islands includ- Secondary pinnules are only rarely found on postero- ing Necker Ridge. lateral pinnules. Secondary pinnules are 0.5–1.5 cm in length. Secondary pinnules are not present on some an- Habitat In Alaska – grows on bedrock and fractured terior primaries, particularly those near the branch tips. bedrock including basalt in small patches (up to 10 in- Tertiary pinnules are usually absent. On thick branches dividuals together) at depths between 2635–2828 m but the primary pinnules may have the appearance of being video observations (Stone, 2014) indicate the species in more than four rows since the sclerenchyme of the branch overgrows the base of some secondary pinnules. ranges to as shallow as 2306 m. Elsewhere – eurybathic, Spines on pinnules are simple, smooth, conical, and depths between 227–2730 m. acute; 0.05–0.13 mm in height (from the center of the base to the apex), and often inclined distally, especially Photos A) A large portion of a preserved (in ethanol) near the distal ends of primary pinnules and on second- T. pseudotristicha colony (USNM 1070975) collected ary pinnules, in which case the abaxial edge is often northeast of Oahu Island, Hawaii, at a depth of 396 two to three times longer than the adaxial edge. The m. B) A close-up view of the same specimen in photo largest spines are found on the distal half of secondary A. C) A T. pseudotristicha colony collected on siltstone pinnules. Spines are arranged in axial rows with three in the central Aleutian Islands at a depth of 2829 m. or four rows visible in lateral view (excluding rows in The distance between the red laser marks is 10 cm. D) which the spines are only partially visible). The distance A close-up view of the same colony in photo C. E) A between adjacent spines in each row is variable, ranging cluster of T. pseudotristicha colonies on a rocky ridge from 0.25 to about 0.4 mm; on average there are about in Kailua Bay, Hawaii, at a depth of 432 m. The holo- 4 spines/mm in each row. type (USNM 98848) is the colony on the right. The dis- Polyps have six primary mesenteries and no second- tance between the red laser marks is about 10 cm. F) A ary mesenteries. Polyps are 2–4 mm in transverse diam- close-up view of the holotype immediately before collec- eter, arranged uniserially on the upper or lateral sides of tion. Photos E and F are courtesy of S. France, L. Mul- the pinnules and subpinnules, with 3–4 polyps/cm. lineaux, and the Hawaii Undersea Research Laboratory. Chapter 2 23 24 Professional Paper NMFS 23 Family Schizopathidae 5. Alternatipathes mirabilis Opresko and Molodtsova, 2021 Description (Adapted from Opresko and Molodtso- ered with coenenchyme, and relatively easy to detach. va, 2021) Colonies are monopodial (with the stem The color of the coenenchyme in life and in ethanol is reaching to the top of the corallum), unbranched, and very light orange. pinnulate. Colonies are to at least 50 cm in height and at least 20 cm in width. The lower unpinnulated stalk Remarks This species is similar to Alternatipathes al- can be considerably longer than the upper, strongly ternata (Brook, 1889), but usually has a longer unpin- triangular-shaped, pinnulated section. The pinnulated nulated stalk, longer pinnules, a wider distal angle of section of the holotype is quite flat and extends hori- the pinnules, and larger polyps. zontally in the water column in situ (see photos E and This species is superficially quite similar to Bathy- F). Pinnules are simple (without subpinnules), arranged pathes ptiloides with which it co-occurs but the pinnu- alternately in two lateral rows (Fig. 2-3D), and gener- lated section of the colony is relatively smaller and more ally decreasing in length from the lower part of the pin- obviously triangular-shaped. nulated section of the stem to the apex. Lowermost pin- nules are >15 cm in length in large colonies. Pinnular Distribution Rare. In Alaska – known from a single density is 11–12 pinnules/3 cm. specimen (USNM 1070972) collected from Derickson The spines on pinnules are smooth, triangular in pro- Seamount in the western Gulf of Alaska and several video file, moderately acute, laterally compressed, and mostly observations made nearby (Fig. 2-8). Elsewhere – not 0.04–0.05 mm in height on the polypar side of the axis. reported, but similar specimens identified only to genus Polyps have six primary mesenteries and four secondary have been collected off Washington (2453 m depth), Cal- mesenteries, are 5–7 mm in transverse diameter, with ifornia (west of the Channel Islands at 4100 m depth), 4–5 polyps/3 cm. Taney Seamount (3054 m depth), Gorda Ridge (2820 m The holdfast is small, inconspicuous, typically cov- depth), and near Hawaii (2638 m depth). Habitat Always occurs singly, attached to basalt and fragmented basalt at depths between 4600 and 4685 m. Photos A) The preserved (in ethanol) A. mirabilis holotype (USNM 1070972; whole colony) collected on Derickson Seamount at a depth of 4685 m. B) A close-up view of the branch- ing pattern of the same spec- imen in photo A. C) A close- up view of the polyps on the same specimen in photo A. D) A photo of the collec- tion of the holotype in situ. E) A suspected A. mirabilis colony observed on Derick- son Seamount at a depth of 4680 m. F) A close-up view Figure 2-8 of the same colony in photo A map of the North Pacific Ocean showing the distribution of Alternatipathes mirabilis () E. The distance between the in Alaska waters. red laser marks in photos D–F is 10 cm. Chapter 2 25 26 Professional Paper NMFS 23 6. Bathypathes alaskensis Opresko and Molodtsova, 2021 Description (Adapted from Opresko and Molodtso- Remarks Many of the Alaska specimens assigned to va, 2021) Colonies are monopodial and unbranched, this newly described species were originally identified but pinnulate. Colonies are up to about 43 cm in height as Bathypathes patula Brook, 1889. The new species with a basal stem diameter up to 2 mm. The stem is differs from B. patula in having 1) a stem that is not straight or curved but usually not distinctly sickle- as distinctly sickle-shaped, 2) relatively longer pin- shaped. Smaller colonies are usually upright or only nules, 3) larger and more numerous conical spines that slightly curved away from the polypar side of the coral- can be bilobed or trilobed, 4) more rows of spines, and lum, whereas larger colonies tend to be strongly curved 5) slightly smaller and more densely arranged polyps. away from the polypar side. The unpinnulated stalk is This species is also similar to B. galatheae Pasternak, typically 6.5–10.0 cm in length, while the pinnulated 1977, a species also found in the North Pacific Ocean section is often >30 cm in length. Pinnules are simple, but not in Alaska, and differs in having smaller spines arranged along the stem in two lateral or anterolateral (0.08–0.14 mm versus 0.18–0.26 mm) and more flex- rows, and grouped in subopposite pairs (see Fig. 2-3E ible pinnules. for a generic example). Pinnular density typically ranges 12–16 pinnules/5 cm. Pinnules are up to about 23 cm Distribution Common and locally abundant. In Alas- in length in the largest colonies (43 cm in height). The ka – principally in the eastern Gulf of Alaska includ- longest pinnules are usually found along the lower or ing the inside waters of Southeast Alaska, the Gulf of middle part of the pinnulated section of the stem. The Alaska Seamount Province (Dickens, Densen, Welker, interior angle formed by the two rows of pinnules is and Pratt Seamounts), and a single specimen from the highly variable; ranging from about 30° to 180° even central Aleutian Islands (Fig. 2-9). Elsewhere – known within the same colony. The distal angle of pinnules is from only two specimens: one collected on the conti- mostly 60–80° (range 45–90°). nental slope off Washington and another from Mon- Spines are smooth, simple, and conical with a rounded terey Canyon (northern California). apex. The largest polypar spines on pinnules are mostly 0.11–0.14 mm in height. Spines are often bilobed or tri- Habitat Found singly or in low-density patches. At- lobed at the apex, eventually forming double and triple taches principally to siltstone but occasionally on bed- spines. Five to seven rows of spines are visible in lateral rock and fractured bedrock, including basalt pillow view of the pinnules. lavas, at depths between 329 and 1837 m. Elsewhere Polyps are uniserially arranged; mostly 4–5 mm in – found at depths between 1215 and 1798 m. transverse diameter (range 3–6 mm) as measured from the distal edge of the distal lateral tentacles to the proxi- Photos A) A preserved (in ethanol) whole B. alasken- mal edge of the proximal lateral tentacles. Polyp density sis colony (USNM 1288462) collected in the eastern is 4–5 polyps/2 cm. Gulf of Alaska at a depth of 515 m. B) A close-up Colonies have large (up to 2 cm in diameter) circular view of a preserved (in ethanol) B. alaskensis specimen holdfasts that are often conspicuously greyish in color (USNM 1070916) collected on Densen Seamount at a and typically bear relatively large (>2 cm) sweeper tenta- depth of 1386 m. C) A B. alaskensis colony (USNM cles on the entire unpinnulated section of the stem (photo 1070919) in situ prior to collection on Pratt Seamount E). This is the only antipatharian in Alaska that often at a depth of 1837 m. D) A lateral view of the same col- bears sweeper tentacles (Bathypathes patula rarely bears ony in photo C. E) The same colony in photo C with a sweeper tentacles); the only other coral in Alaska with close-up view of the sweeper tentacles and grey-colored sweeper tentacles is the keratoisidid Isidella tentaculum holdfast. F) A B. alaskensis colony (USNM 1288461) Etnoyer, 2008. The typical color of the coenenchyme in collected in the eastern Gulf of Alaska at a depth of 515 life is dark orange but occasionally lighter orange; color m. The distance between the red laser marks in photos in ethanol is dull orange. This species attracts swarms of C–F is 10 cm. euphausiids. Chapter 2 27 28 Professional Paper NMFS 23 6. Bathypathes alaskensis Opresko and Molodtsova, 2021 (continued) Figure 2-9 A map of the North Pacific Ocean showing the distribution of Bathypathes alaskensis () in Alaska waters. Chapter 2 29 7. Bathypathes patula Brook, 1889 Description Colonies are monopodial and un- 90.4.9.19) collected in the North Pacific Abyssal Prov- branched, but pinnulate. Colonies are up to 30 cm or ince is a whole colony about 20 cm in height with rela- more in height and the stem usually has a sickle-shaped tively short pinnules (approximately 8 cm) and spines curvature. The unpinnulated stalk is of varying length. not more than 0.07 mm in height. This species is known Pinnules are simple (without subpinnules); arranged from only one specimen in Alaska and it has longer pin- along the length of the stem biserially in two lateral or nules and slightly taller spines than the type specimen. anterolateral rows, and grouped in subopposite pairs (see Fig. 2-3E for an example). Pinnules in each lateral Distribution Rare. In Alaska - known from only one row are spaced up to 12 mm apart. Pinnular density specimen (consisting of only fragments) collected on the along the stem is mostly 6–8 pinnules/3 cm (total for deep abyssal plain south of the Aleutian Trench in the both rows). The longest pinnules are found along the western Gulf of Alaska (Fig. 2-10). Elsewhere – report- lower or middle part of the pinnulated section of the edly cosmopolitan and wide ranging (North and South stem. Maximum length of pinnules is about 8 cm in Pacific, Antarctica, North Atlantic, and Mediterranean colonies with 10 pairs of pinnules, but longer in larger and Caribbean Seas) but designations are likely to be colonies. revised with further study of the genus. Spines on the stem and pinnules are simple, smooth, triangular, and laterally compressed. Spines are slightly Habitat Grows singly on bedrock and fractured larger on the polyp side of the axis. On pinnules 0.20– bedrock (cobble-size) at depths near 4950 m. This is 0.25 mm in diameter, the polypar spines are up to 0.07 a deeper water species than B. alaskensis. Elsewhere – mm in height and the abpolypar spines are 0.02–0.04 eurybathic; on fractured basalt and pillow lavas. The mm in height. On older colonies with thicker pinnules, type specimen was collected at a depth near 5200 m. polypar spines are up to about 0.1 mm in height and Other reported specimens from the Pacific Ocean range the abpolypar spines are 0.03–0.05 mm shorter than the in depth from 320 m (Hawaii) to 5020 m (Kurile-Kam- polypar spines. Typically, only three rows of spines are chatka Trench, Russia). visible in a lateral view of the pinnules. Polyps are usually 5–7 mm in transverse diameter as measured from the distal edge of the distal lateral ten- Photos A) A preserved (in ethanol) whole B. patula tacles to the proximal edge of the proximal lateral ten- colony (USNM 1453622) collected near American Sa- tacles; the polyp density along the length of a pinnule is moa in the central Pacific Ocean at a depth of 3683 m. approximately 2.5 polyps/2 cm. Live polyps are nearly B) The same specimen in photo A in situ just prior to translucent. collection. C) A close-up view of the same specimen in The holdfast is relatively inconspicuous but strong photo B. D) A B. patula colony (USNM 1467584) just and the unpinnulated section of the stem rarely bears prior to collection in the Musician Seamounts, Hawaii. small sweeper tentacles. The color of the coenenchyme E) A close-up view of the same colony in photo D. The in life is yellowish orange to pinkish orange and a dark- distance between the red laser marks in photos B and D er orange in ethanol. is 10 cm. Photos B–E are courtesy of the National Oce- anic and Atmospheric Administration’s Office of Ocean Remarks The type specimen of B. patula (NHMUK Exploration and Research. 30 Professional Paper NMFS 23 7. Bathypathes patula Brook, 1889 (continued) Chapter 2 31 Figure 2-10 A map of the North Pacific Ocean showing the distribution of Bathypathes patula (), B. pti- loides (◆), and B. tiburonae (▲) in Alaska waters. 32 Professional Paper NMFS 23 8. Bathypathes ptiloides Opresko and Molodtsova, 2021 Description (Adapted from Opresko and Molodtso- in having a longer unpinnulated section of the stem, larger va, 2021) Colonies are monopodial, unbranched, and polyps, relatively longer pinnules, and smaller spines. It pinnulate. Colonies are to at least 78 cm in height, differs from B. alaskensis in having larger polyps, but about 40 cm in width, with a basal diameter of the stem smaller spines. This species is superficially quite similar up to 2 mm. Coralla consist of a very long unpinnulated to Alternatipathes mirabilis with which it co-occurs but stalk, up to 66 cm in length, topped by a short pinnu- the pinnulated section of the colony is much longer and lated section only 10–20 cm in length. The lower sec- not as obviously triangular-shaped. tion of the stem is inclined towards the polyp side of the colony and then more strongly curved in the same direc- Distribution Uncommon but locally abundant. In tion, after which it curves in the opposite direction such Alaska – known from only two specimens; one collect- that the upper part of the pinnulated section is almost ed on Chirikof Seamount and the second collected on vertical. The pinnulated section of the colony is quite Derickson Seamount in the Gulf of Alaska Seamount convex and occasionally lies more or less horizontally in Province (Fig. 2-10). At both locations multiple video the water column. observations were made indicating that this species is Pinnules are simple, arranged along the stem in two locally common and occurs at patch densities up to 3 lateral or anterolateral rows, and in subopposite pairs colonies/m2. Elsewhere – not reported. (see Fig. 2-3E for a generic example). Pinnules are to at least 12 cm in length and spaced 6–13 mm apart in Habitat Forms small scattered patches (up to 25 colo- each lateral row. Pinnular density (total for both rows) nies) on bedrock and fractured bedrock including basalt is 6 pinnules/3 cm on the lowermost pinnulated portion pillow lavas at depths between 4491 and 4676 m. Often of the stem and 8 pinnules/3 cm distally. The pinnules occurs in association with the keratoisidid Bathygorgia are inclined distally such that the distal angle they form profunda Wright, 1885. with the stem is 70–80°. Interior angle formed by the two rows of pinnules is 120° or more. Photos A) Preserved (in ethanol) large sections of a Spines are simple, smooth, triangular in lateral view, B. ptiloides colony (USNM 1070974) collected on De- compressed, with a rounded apex and a base that ex- rickson Seamount at a depth of 2664 m. B) A close-up tends out in a very shallow slope in both the distal and view of the same specimen in photo A. C) A zoomed-in basal directions. Polypar spines are up to 0.04 mm in view of the same specimen in photo A. D) The same height and abpolypar spines are 0.035 mm tall. Six rows colony in photo A with the holdfast visible (indicated of spines are visible in one lateral view. by the white circle) just prior to collection. E) A B. pti- Polyps are uniserially arranged on one side of the cor- loides colony (USNM 1071408) with the holdfast vis- allum; 6–8 mm in transverse diameter, with 3 polyps/3 ible in the center of the white circle. The top part of cm. Polyps on the stem are 7 mm in transverse diameter. the colony was collected on Derickson Seamount at a The holdfast is small and inconspicuous but firmly depth of 4491 m. F) A close-up view of the collection of attached to the substrate. Color in situ is orange to light the top pinnulated section of the same colony in photo orange; creamy orange in ethanol. E. The stalk and holdfast are indicated by the white el- lipse. The distance between the red laser marks in pho- Remarks This species differs from Bathypathes patula tos D and E is 20 cm. Chapter 2 33 34 Professional Paper NMFS 23 9. Bathypathes tiburonae Opresko and Molodtsova, 2021 Description (Adapted from Opresko and Molodtso- pinnules (up to 0.09 mm compared to ≤0.07 mm in va, 2021) Colonies are monopodial, unbranched, and height), and larger polyps (up to 12 mm compared to pinnulate. Largest colonies measure up to 55 cm in 5–7 mm). Bathypathes tiburonae is similar to B. pti- height (total length of the stem), 30 cm in width, and loides in having a long unpinnulated stalk, but it has the basal diameter of the stem is up to 3 mm. Coralla larger pinnular spines (0.09 mm compared to 0.04 mm consist of an unpinnulated stalk up to 37 cm in length in height), and larger polyps (up to 12 mm compared to and topped by a pinnulated section only 18 cm in 8 mm in diameter). length. The stem is relatively straight and upright. Pin- nules are simple, arranged along the stem in two lateral Distribution Uncommon. In Alaska – known only or anterolateral rows and in subopposite pairs (see Fig. from the deep continental slope south of Davidson 2-3E for a generic example). Pinnules are to at least 24 Bank in the western Gulf of Alaska (Fig. 2-10). Else- cm in length and spaced up to 10–18 mm apart in each where – also known from Gorda Ridge off northern lateral row. Pinnular density (total for both rows) is 4 California (USNM 1116837) and off Oregon (USNM pinnules/3 cm on the lowermost pinnulated portion of 1234549). the stem and 6 pinnules/3 cm distally. The distal angle of the pinnules with the stem is about 80°. The interior Habitat Known from a single specimen collected on angle formed by the two rows of pinnules is about 120° a large siltstone bank at a depth of 3356 m and three nearest to the stem. specimens observed nearby; one on the same bank as Spines are simple, smooth, triangular in lateral view, the collected specimen (see photo C) and two specimens and compressed with a rounded apex. Polypar spines on a large boulder at a depth of 3280 m (see photo E). are up to 0.09 mm in height and abpolypar spines are Co-occurs with the keratoisidid Bathygorgia profun- up to 0.08 mm in height. Five or six rows of spines are da and the primnoid Narella bayeri Cairns and Baco, visible in one lateral view. 2007. Elsewhere – depths between 2516 and 3150 m. Polyps are uniserially arranged on one side of the cor- allum, mostly 9–10 mm (up to 12 mm) in transverse di- Photos A) A large portion of a preserved (in ethanol) ameter, and there are 3 polyps/4 cm or 4–5 polyps/5 cm. B. tiburonae colony (USNM 1116837; holotype) col- The polyps on the stem are up to 17 mm in transverse lected on southern Gorda Ridge off northern California diameter, with 2 polyps/4 cm. at a depth of 3121 m. B) A close-up view of the same The holdfast is relatively small and elliptical but quite specimen in photo A. Photos A and B are reprinted with strong and firmly attached to the substrate. The color permission from Zootaxa. C) A B. tiburonae colony of the coenenchyme is orange to light orange in life and (USNM 1459868) collected near Davidson Bank in the creamy orange in ethanol. western Gulf of Alaska at a depth of 3356 m. The hold- Remarks This species differs from B. alaskensis in fast is visible in the center of the white circle. Note the having a longer unpinnulated stalk (about 67% of the second colony present below the white circle. D) The total stem length compared to about 25% in colonies same colony in photo C during collection near David- 40–55 cm in height); smaller polypar spines (0.09 mm son Bank in the western Gulf of Alaska at a depth of in height compared to 0.11–0.14 mm in height), and 3356 m. The holdfast (not collected) is visible in the larger polyps (up to 12 mm in diameter compared to center of the white circle. E) Two B. tiburonae colo- only 3–6 mm in diameter). It differs from B. patula in nies (a large colony just right of the laser marks and a having a longer unpinnulated stalk (about 67% of the smaller one at the center of the white circle) observed total stem length compared to about 50% in B. patula near Davidson Bank at a depth of 3280 m. The distance of similar size), slightly larger polypar spines on the between the red laser marks in photos C–E is 20 cm. Chapter 2 35 36 Professional Paper NMFS 23 10. Bathypathes sp. A Description The collected colony is monopodial, un- in having relatively small spines as well as very small branched, and pinnulate with a maximum length of polyps. Although B. patula and B. ptiloides both have about 36 cm (estimated from in situ photos). The shape spines of a similar size, the polyps of these two species of the stem is unknown but it appears to be relatively are larger: 6–8 mm or more in transverse diameter. In straight, stout, and flexible. The unpinnulated stalk is addition, the pinnular density in this species, approxi- approximately 25% of the total colony length. Pinnules mately 14 total pinnules/3 cm (estimated from in situ are simple (without subpinnules) and arranged along photos), appears to be greater than in any other Alaska the stem biserially in two anterolateral rows. The lon- Bathypathes spp., including B. alaskensis which has gest pinnules are about 12 cm (estimated from in situ 8–12 pinnules/3 cm. However, an exact measurement photos). cannot be made because of the orientation of the colony Spines are simple, smooth, triangular to conical, and shown in the photos. Compared to Bathypathes alasken- 0.025–0.050 mm in height. Polyps are small, mostly sis, this species has pinnules that appear to be shorter 3.5–5.5 mm in transverse diameter, with a density of and stiffer and the coenenchyme is a lighter color. approximately 2 polyps/cm. This species occurs in habitats with tube-dwelling The holdfast is relatively large, circular, covered with polychaetes (family Sabellidae), hydroids, and the ker- a very thin coenenchyme, and firmly attached to the atoisidid coral Bathygorgia profunda, and it harbors substrate. Color of the coenenchyme in life is a uniform ophiuroids. light orange. Distribution Rare. In Alaska – known from only a Remarks Only pinnules and in situ photos of the sin- single specimen collected south of Davidson Bank in gle collected specimen were available for examination. the western Gulf of Alaska (Fig. 2-11). Elsewhere – un- This species differs from other known Bathypathes spp. known. Figure 2-11 A map of the North Pacific Ocean showing the distribution of Bathypathes sp. A () and Bathypathes sp. B (◆) in Alaska waters. Chapter 2 37 Habitat Occurs singly (although see photo A below) smaller colony (indicated by the white circle), likely the in areas of rough conglomerate (largely fragmented ba- same species, grows nearby. B) The same colony in pho- salt) at depths near 3275 m. to A with the holdfast visible in the white circle at the top of the photo. C) A broad view of the same colony in Photos A) A Bathypathes sp. A colony (USNM photo A. D) A close-up view of the pinnulated section 1071406) collected south of Davidson Bank in the of the same colony in photo A. The distance between western Gulf of Alaska at a depth of 3275 m. A second the red laser marks in photos A–D is 10 cm. 38 Professional Paper NMFS 23 11. Bathypathes sp. B Description The collected colony is monopodial, un- Remarks This species is unique from other nominal branched, and pinnulate. The colony is about 36 cm species of Bathypathes in that the unpinnulated stalk in height and the stem near the base has a diameter of is very short (only about 11 mm in length). Most spe- only about 3 mm. The stem is curved but not sigmoidal. cies of Bathypathes have unpinnulated stalks of 7 cm The unpinnulated stalk is very short (11 mm in length) or more. The very short unpinnulated stalk of the col- which is quite unusual for any species of Bathypathes. ony may be an abnormal condition resulting from an Note that in photos A and B (below) pinnules appear to injury followed by regrowth of the sclerenchyme and be growing directly from the holdfast but that is an ar- formation of the pinnules near the base. In terms of the tifact of the photographs. Pinnules are simple (without size and shape of the pinnular spines and length of the subpinnules) and arranged along the length of the stem pinnules, this species is similar to B. tiburonae, but in biserially in two anterolateral rows and in a suboppo- terms of the size and density of the polyps it is more site fashion. Pinnules are ≥24 cm in length with a basal similar to B. ptiloides with which it co-occurs. diameter of about 1 mm. Pinnules are 7–10 mm apart, resulting in a pinnular density (total for both sides) of Distribution Rare. In Alaska – known from only one 6 pinnules/3 cm on the lower part of the corallum and specimen (USNM 1070973) collected on Derickson Sea- 8–10 pinnules/3 cm on the upper part. mount in the Gulf of Alaska Seamount Province (Fig. The spines are simple, smooth, triangular, laterally 2-11). Elsewhere – unknown. compressed, and with a shallow slope on the distal and proximal edges. The polypar spines are 0.04–0.07 mm Habitat Occurs singly on fragmented bedrock (basalt) in height whereas the abpolypar spines are 0.02-0.03 at a depth of 4497 m. mm in height. Five to seven rows of spines are visible in lateral view, with 4 spines/mm in each row. Photos A) A preserved (in ethanol) whole Bathy- The polyps are 7–8 mm in transverse diameter, with pathes sp. B colony (USNM 1070973) collected on a density of 2.0–2.5 polyps/2 cm. The holdfast is small, D erickson Seamount at a depth of 4497 m. B) A close- cup-shaped, and firmly attached to the substrate. The up view of the same specimen in photo A. color of the coenenchyme in life is light orange and a lighter orange in ethanol. Chapter 2 39 12. Dendrobathypathes boutillieri Opresko, 2005 Description (Adapted from Opresko, 2005) Colo- relatively thick and blunt, and about 2 mm in length in nies are generally wider (≥150 cm in width) than taller preserved samples. (≥95 cm in height). Coralla are extensively branched and Larger colonies have a massive (up to 8×3 cm), asym- pinnulate; the stem, branches, and primary pinnules are metrical, very strong holdfast typically covered with generally in one plane. Primary pinnules are arranged coenenchyme and with a characteristic black stain on alternately along the stem and branches in two lateral the substrate surrounding the holdfast (see photo D on to anterolateral rows, and occasionally with one (rarely next page). Color of flesh in life is bright orange and two) simple secondary pinnules near the base (Fig. 2-3F). orange-brown in ethanol preserved specimens. Primary pinnules within each anterolateral row are spaced 5–8 mm apart resulting in 3–4 pinnules/cm (to- Remarks Larger colonies of D. boutillieri are often used tal for both rows). The interior angle formed by rows of as an elevated perch by king crabs (Paralomis spp.). primary pinnules is near 90°. Primary pinnules are in- Distribution Uncommon but widespread. In Alaska – clined distally, generally not more than 2 cm in length, eastern Gulf of Alaska, central Aleutian Islands (south- and with a basal diameter of about 0.5 mm (excluding west of Amlia Island to Adak Canyon), and Giacomini spines). Secondary pinnules are present on the anterior Seamount in the Gulf of Alaska Seamount Province (polypar) side of the primaries near the base and proj- (Fig. 2-12). Elsewhere – northern British Columbia. ect out from the polypar side of the corallum. Second- ary pinnules are usually not more than about 1 cm in Habitat Found singly on bedrock, siltstone, and boul- length with basal diameter of not more than 0.4 mm. ders. Collected specimens range in depth from 932 to Secondary pinnules are inclined towards the distal end 2161 m but video observations indicate the species of the primary pinnule from which they arise. Tertiary ranges to depths as shallow as 859 m in the central pinnules are not present. Aleutian Islands (Stone, 2014). Elsewhere – collected at Pinnular spines are simple (rarely forked), conical, depths between 1903 and 1908 m. with a rounded to somewhat acute apex, and slightly compressed; subequal to slightly larger on one side of Photos A) A dried whole D. boutillieri colony the axis (usually but not always corresponding to the (USNM 1094105) collected off British Columbia at a polypar side). Polypar spines are mostly 0.10–0.14 mm depth of 1460 m. B) A close-up view of the same speci- but up to 0.18 mm in height; abpolypar spines are gen- men in photo A. C) A close-up view of the holdfast on erally 0.02–0.04 mm shorter than polypar spines. Most the same specimen in photo A. D) A D. boutillieri col- spines project out at right angles to the axis, but some ony (USNM 1093060) sampled in the central Aleutian are angled slightly towards the base of the pinnule on Islands at a depth of 2019 m. Note the characteristic which they occur. Spines are not always arranged in reg- black stain around the holdfast indicated by the white ular longitudinal rows, four to six of which are usually circle. The octopus, Graneledone boreopacifica, takes visible in lateral view. Spines are spaced apart within refuge under the colony. The distance between the red each row at varying distances, from about 0.35 mm to laser marks is 20 cm. E) A D. boutillieri colony (USNM as much as 0.75 mm, but generally there are 3–4 spines/ 1093057) sampled in the central Aleutian Islands at a mm in each row. depth of 2161 m. A spiny paralomis (Paralomis multi- Polyps occur on one side of the corallum, correspond- spina) perches in the colony. F) A view of the top of a ing to the side on which the secondary pinnules are lo- D. boutillieri colony (USNM 1070762) sampled on Gi- cated. Polyps are mostly 3.5–5.0 mm in transverse diam- acomini Seamount at a depth of 1689 m. The distance eter (as measured from the distal edge of the distal lateral between the red laser marks in photos E and F is 10 cm. tentacles to the proximal edge of the proximal lateral G) A close-up view of the branches and polyps on the tentacles). Polyp density is 2–3 polyps/cm. Tentacles are same colony in photo F. 40 Professional Paper NMFS 23 12. Dendrobathypathes boutillieri Opresko, 2005 (continued) Chapter 2 41 Figure 2-12 A map of the North Pacific Ocean showing the distribution of Dendrobathypathes boutillieri () and Dendrobathypathes sp. (♦) in Alaska waters. 42 Professional Paper NMFS 23 13. Dendrobathypathes sp. Description Colonies are most similar to Dendro- Distribution Extremely rare. In Alaska – known from bathypathes boutillieri (especially specimen USNM only four specimens observed on Welker Seamount and 1070762 collected on Giacomini Seamount at 1689 m one specimen observed on Dickens Seamount in the depth) but with denser branching, longer primary pin- Gulf of Alaska Seamount Province (Fig. 2-12). Else- nules, and very few subpinnules. Colonies are bushy, where – unknown. flexible, not rigid, and have a very floppy or creeping appearance. Colonies are to about 15 cm in length, 10 Habitat Found singly or in clusters of up to four indi- cm in width, and 9 cm in diameter. Color of the coen- viduals on sloped habitat of bedrock and fractured bed- enchyme is pinkish-orange with only parts of the under- rock at depths of 1652–2634 m. lying corallum visible. Photos A) A Dendrobathypathes sp. colony photo- Remarks This taxon is known only from video foot- graphed on Welker Seamount at a depth of 2634 m. B) age of four specimens observed together (within 1 m of A close-up view of the same colony in photo A. C) A each other) on Welker Seamount and a fifth specimen Dendrobathypathes sp. colony photographed on Dick- observed on nearby Dickens Seamount. It is unlike any ens Seamount at a depth of 1652 m. The distance be- other antipatharian known in the region in that it is not tween the red laser marks in photos A and C is 10 cm. arborescent but has more of a creeping habitus. It un- doubtedly represents an unknown species and should be a high priority for collection. Chapter 2 43 14. Lillipathes wingi Opresko, 2005 Description (Adapted from Opresko, 2005) Coralla less near branch tips. The interior angle formed by two are unbranched or sparsely to moderately branched, anterior rows increases from about 90° at the tips of mostly in one plane, with up to six branches (see pho- branches to 120–150° basally. The distal angle of pin- to A on next page). Colonies are to at least 150 cm in nules is 60–70°. height and up to 75 cm in width. The stem usually ex- Spines are conical, simple, each with a rounded apex, tends to the top of the corallum. The stem and branches and are larger on the polyp side of the axis. Polypar are pinnulate. Pinnules are simple and without subpin- spines on pinnules are mostly 0.07–0.10 mm in height; nules; they are arranged in four rows along the length abpolypar spines are 0.03–0.05 mm shorter. Double of the stem and branches, two lateral and two anterior, spines are occasionally present, especially on the polypar and also in alternating biserial groupings of two pin- side of the axis. Four to six rows of spines are visible in nules each (see Fig. 2-3G for a generic example). One lateral view on pinnules. Spacing of spines within each or both anterior pinnules are occasionally missing. An- row is not uniform; the mutual distance is 0.18–0.37 terior pinnules are inserted slightly below (1–2 mm) ad- mm, on average 0.30 mm, resulting in 4–5 spines/mm jacent lateral pinnules. Pairs of pinnules on either side in a row. Polyps are 2.4–3.6 mm in transverse diameter. of the axis are 2.5–3.0 mm apart with 10–14 pinnules/ Holdfasts are relatively small and irregular in shape, cm (total for all rows). Pinnules on smaller branches are devoid of coenenchyme and firmly attached. The col- 1–2 cm in length. Pinnules on stem and larger branches or of the coenenchyme in life is bright orange, creamy are up to 5 cm in length with a basal diameter of about white in ethanol, and light orange when dried. 0.4 mm. Generally, the lateral pinnules are longer than anterior pinnules by 5–10 mm; however, anterior pin- Remarks In situ observations of large specimens indi- nules are occasionally longer than adjacent lateral pin- cate that they often harbor ophiuroids including Ophia- nules. The interior angle formed by the two rows of cantha sp. and swarms of euphausiids. lateral pinnules is typically close to 180°, but slightly Distribution Uncommon. In Alaska – this is predomi- nantly a continental slope species. Specimens have been collected in the east- ern Gulf of Alaska, a single seamount in the Gulf of Alaska Seamount Province (Murray Seamount), and the eastern Bering Sea (Fig. 2-13). Elsewhere – ranges south through British Co- lumbia to Astoria Canyon (off the southwest coast of Washington). Habitat Found on bed- rock, siltstone, and cobbles at depths between 600 and 909 m. Elsewhere – collect- ed at depths between 896 and 1903 m. Photos A) A preserved (in ethanol) whole L. wingi Figure 2-13 colony (USNM 1094142) A map of the North Pacific Ocean showing the distribution of Lillipathes wingi () in Alaska collected off northern Brit- waters. ish Columbia at a depth of 1057 m. B) A close-up view 44 Professional Paper NMFS 23 of the same specimen in photo A. C) A snailfish (Care- D) A branch (fresh) sampled from the colony in photo proctus sp.) seeks refuge in a large, multi-branched L. C. E) A fresh whole L. wingi colony (USNM 1482122) wingi colony (USNM 1249983) sampled in Zhemchug collected in the eastern Gulf of Alaska at a depth of Canyon in the eastern Bering Sea at a depth of 909 m. 805 m. Chapter 2 45 15. Lillipathes sp. A Description Colonies are large (up to 100 cm or more The sternostylid crab Sternostylus iaspis often uses in height and width), rather planar, and sometimes large colonies as platforms for suspension feeding. Some slightly concave. They are moderately branched (up to colonies host polychaetes (see photo C on next page) 12 side branches off the main stem) to the fourth or- and ophiuroids are often associated with larger colonies der or more; branches may overlap. Stem and branches too, presumably also using them for a feeding platform. have simple pinnules arranged bilaterally along the stem and branches in a variable number of rows ranging Distribution Common and locally abundant. In Alas- from two to six and also in alternating groups of up ka – this is predominantly a seamount species. Speci- to three pinnules each. Pinnules are mostly 2.0–3.5 cm mens have been collected on Dickens and Murray Sea- in length. Pinnular density (total for both rows) is 5–7 mounts in the Gulf of Alaska Seamount Province and pinnules/cm where there are only two rows and 10–11 a single specimen has been collected west of Prince of pinnules/cm where there are four rows. The distal angle Wales Island in the eastern Gulf of Alaska. Colonies of pinnules is about 60° or greater. have also been observed on video footage collected on Spines on pinnules are simple (without lobes or bifur- Welker, Pratt, Quinn, Giacomini, Patton, Marchand, cations), smooth, triangular, acute or slightly rounded and Chirikof Seamounts (Fig. 2-14). Colonies are par- at the apex, slightly compressed, and project out per- ticularly abundant on Murray Seamount where they pendicular to the axis. Polypar spines are up to 0.08 mm have been observed up to 5 colonies/m2. Elsewhere – in height on the thinner sections of the pinnules and up unknown. to 0.1 mm on the thicker sections. Abpolypar spines are usually about half the size of the polypar spines. Four or Habitat Occasionally found singly but more often five rows of spines are visible in lateral view. in patches of 10 or more colonies and typically in lin- Polyps on pinnules are 3–4 mm in transverse diam- ear rows (colonies may overlap) along bedrock ridges, eter and the interpolypar space is very narrow. Polyp including lava flows and breccias. Also found on frac- density is 2.5–3.0 polyps/cm. On the smallest branches tured bedrock, boulders, and cobbles. Total depth range the polyps are similar in size to those on the pinnules. is 600 to 2738 m. As the branches increase in size the polyps become fewer and fewer until they are not visible on the stem. Photos A) A Lillipathes sp. A specimen (USNM Holdfasts are relatively small and irregular in shape, 1070908; preserved in ethanol) collected on Dickens occasionally covered with coenenchyme, and firmly at- Seamount at a depth of 1652 m. B) A close-up view tached. The color of the coenenchyme in life is bright of the same specimen in photo A. C) A close-up view to light orange with reddish-orange branch axes; lighter of the same specimen in photo A with associated poly- orange to cream in ethanol. chaete (indicated by the white circle). D) A Lillipathes sp. A colony (USNM 1070908) collected on Dickens Remarks This species differs from L. wingi in having Seamount at a depth of 1652 m. E) A close-up view of a more densely branched corallum with the branches the same colony in photo D. The distance between the projecting at varying angles to the lower order branch- red laser marks in photos D and E is 10 cm. F) A Lil- es from which they arise. In addition, the pinnules are lipathes sp. A colony observed on Quinn Seamount at shorter (2.0–3.5 cm versus up to 5.0 cm in length) and a depth of 761 m. G) Lillipathes sp. A colonies on a the number of rows of pinnules is quite variable, ranging bedrock ridge on Murray Seamount at a depth of 1211 from two to six. Because six or more rows of pinnules is m. H) Lillipathes sp. A colonies on a bedrock ridge on characteristic of the genus Parantipathes, further study Murray Seamount at a depth of 1086 m. The distance is needed, including genetic sequencing, to determine between the red laser marks in photos F–H is 20 cm. whether the genetic boundaries between Lillipathes and Parantipathes need to be redefined. 46 Professional Paper NMFS 23 15. Lillipathes sp. A (continued) Chapter 2 47 Figure 2-14 A map of the North Pacific Ocean showing the distribution of Lillipathes sp. A () in Alaska waters. 48 Professional Paper NMFS 23 16. Parantipathes euantha (Pasternak, 1958) Description Coralla are monopodial, without branch- cm in length; those in anterior rows are not more than es or very sparsely branched. The stem and branches 2.8 cm in length (Molodtsova and Pasternak, 2005). are pinnulate. The corallum measures to more than 50 Some of the Alaska specimens have pinnules that are cm in height and to 14 cm in width. The lower part longer than those in the type specimen (7 cm versus 4 of the stem is without branches and may be sharply cm), and more densely arranged (up to 20 pinnules/cm curved in those colonies growing on the sides of ver- versus 12–14 pinnules/cm). tically sloped substrate. Pinnules are simple, without Spines on the pinnules are simple, smooth, rounded subpinnules, and arranged in up to six rows along the at the apex, compressed, and triangular in lateral view. length of the stem and branches (if present), and in They project perpendicular to the axis, are 0.07–0.10 alternating semi-spiral groups of up to three pinnules mm in height, arranged in longitudinal rows with five each on both sides of the axis (see Fig. 2-3H for a ge- to six visible in lateral view, and within each row the neric example). The lowermost pinnules on the stem spines are mostly 0.27–0.40 mm apart resulting in 3.0– may be in only two rows, increasing to four rows high- 3.5 spines/mm in each row (Molodtsova and Pasternak, er up, and then to six rows. In each group of semi- 2005). spiral pinnules, the anterior one is the most basal and Polyps on the pinnules are arranged uniserially on the the posterior one the most distal relative to the direc- front, upper or lower side, mostly 1.9–2.7 mm in trans- tion of the stem. Individual pinnules may be missing in verse diameter (maximum about 3 mm), and spaced some groups. Each semispiral group of three pinnules 0.34–1.23 mm apart resulting in 2.5–3.3 polyps/cm extends 2.25–5.00 mm along the axis, with 2.2–2.7 (Molodtsova and Pasternak, 2005). groups/cm and 12–14 pinnules/cm (total for all rows). Holdfasts are small, typically cup-shaped and incon- Pinnules may curve toward the back (abpolypar) side of spicuous, occasionally covered with coenenchyme and the corallum and may also extend distally (distal angle not so strongly attached to the substrate. Color of the 50–80°). Pinnules in posterior rows are mostly about 4 coenenchyme in life is medium to dark orange; light orange when preserved in ethanol. Remarks This species was originally described as Bathypathes euantha Pas- ternak, 1958. Parantipathes euantha primarily differs from P. pluma in having shorter pinnules (up to 7 cm versus up to 13 cm). Ophiu- roids and swarms of euphau- siids are often associated with this species. Distribution Uncommon. In Alaska – this is predomi- nantly a seamount species having been collected on Densen, Welker, and Pratt Seamounts and additionally observed on video footage collected on Quinn Sea- mount. The only specimen not collected on a seamount Figure 2-15 was collected in southern A map of the North Pacific Ocean showing the distribution of Parantipathes euantha (), P. Amchitka Pass in the central pluma (♦), and Parantipathes sp. A (▲) in Alaska waters. Aleutian Islands (Fig. 2-15). Elsewhere – not specifically Chapter 2 49 reported but Parantipathes morphotypes, likely P. euan- Photos A) A preserved (in ethanol) whole P. euantha tha or P. pluma, are reported off northern Vancouver colony (USNM 1070913) collected on Pratt Seamount Island (British Columbia), Juan de Fuca Ridge (Wash- at a depth of 1907 m. B) A close-up view of the same ington), and northern California. specimen in photo A. C) The same colony in photo A in situ just prior to collection. D) A P. euantha colony Habitat In Alaska – typically occur singly or in low- (USNM 1070631) just prior to collection on Densen density patches with Narella arbuscula Cairns and Baco, Seamount at a depth of 2345 m. E) A P. euantha colony 2007 on bedrock, including basalt pillow lavas and boul- observed on Quinn Seamount at a depth of 2159 m. ders, at depths between 862 and 2762 m. Elsewhere The base of the colony is indicated by the white circle. – Parantipathes range in depth from 859 to 1904 m in F) A close-up view of the same colony in photo E. The northern British Columbia, 2283 m on Juan de Fuca distance between the red laser marks in photos C, E, Ridge, and 2820 m off northern California. and F is 10 cm. 50 Professional Paper NMFS 23 17. Parantipathes pluma Opresko and Molodtsova, 2021 Description (Adapted from Opresko and Molodtso- bryozoans, actiniarians, and stoloniferous octocorals va, 2021) Coralla are monopodial and typically un- (Clavularia spp.) and are used as elevated perches by branched but occasionally branched (see photo E). The Verrill’s king crab (Paralomis verrilli). Other associated largest recorded colony is up to 220 cm in height with fauna include ophiuroids, euphausiids, and amphipods, 12 branches. Branches may develop within 10 cm of and large, multi-branched colonies often host snailfish the holdfast. Stem and branches are densely pinnulate. (Careproctus sp.). Lower 15–20% of the main stem lacks pinnules. Pin- nules are simple, without subpinnules, up to 13 cm in Distribution In Alaska – central Aleutian Islands length, inclined distally, arranged bilaterally in varying (southwest of Amlia Island to Amchixtam Chaxsxii) degrees of regularity in four to six rows along the length and Zhemchug Canyon in the eastern Bering Sea (Fig. of the stem (and branches when present), with two to 2-15). Elsewhere – unknown (but see the comment three rows on each side of the axis, and in alternating above for P. euantha). semi-spiral groups of two to three (very rarely four) pin- nules each (Fig. 2-3H). Pinnular density is quite vari- Habitat In Alaska – typically occurs on bedrock, able, but the total for all rows is commonly 11–13 pin- boulders, and cobbles, singly or in patches (up to 5 col- nules/cm (range 9–16 pinnules/cm). onies/10 m2). Collected specimens range in depth from Spines are smooth, usually simple, acute, and trian- 602 to 977 m but video observations (Stone, 2014) in- gular in lateral view with the base extending out along dicate that the species ranges to depths of 1562 m in the the axis; up to about 0.09 mm in height. Spines on thin- central Aleutian Islands. Elsewhere – unknown (but see ner parts of the pinnules are more rounded at the apex. the comment above for P. euantha). Polyps on pinnules are 2.8–3.2 mm in transverse diam- eter, arranged uniserially, with 2.5–3.0 polyps/cm on Photos A) The dried top half of a P. pluma colony the axis. (USNM 1482130) collected in Adak Canyon, cen- Holdfasts are large, typically circular, conspicuous in tral Aleutian Islands at a depth of 602 m. B) A close- larger colonies, often covered with coenenchyme, and up view of the same specimen in photo A. C) A dried strongly attached to the substrate. Color of the coen- whole P. pluma colony (USNM 1498742) collected enchyme in life is medium to dark orange and a duller in Zhemchug Canyon in the eastern Bering Sea at a orange when preserved in ethanol. depth of 977 m. D) A P. pluma colony photographed in the central Aleutian Islands at a depth of 1003 m. Remarks This species differs from P. euantha primar- E) A multi-branched P. pluma colony photographed in ily in having longer pinnules (up to 13 cm versus up to the central Aleutian Islands at a depth of 1003 m. F) A 7 cm). Holdfasts are very strong so the colony skeleton close-up view of the branching pattern of P. pluma. The often remains upright and in place after death. Conse- distance between the red laser marks in photos D and E quently, skeletons are often incrusted with hydroids, is 10 cm. Chapter 2 51 52 Professional Paper NMFS 23 18. Parantipathes sp. A Description Colonies are monopodial or sparsely Distribution Rare. In Alaska – known only from branched to the first order and measure to about 20 cm two collected specimens and seven video observations in height. Simple pinnules are arranged in a bottlebrush made on Giacomini Seamount in the Gulf of Alaska fashion along the stem and branches in six to eight rows. Seamount Province (Fig. 2-15). Colonies occur singly or Pinnules are up to about 5 cm in length and are also two within close proximity. Elsewhere – unknown. arranged in alternating semi-spiral groups of three to four pinnules each. Pinnular density (total for all rows) is Habitat Grows on bedrock (basalt) and fractured 20–22 pinnules/cm. The distal angle of the branches and bedrock (including boulders and cobbles) at depths be- pinnules is rather wide (80–90°). Spines are short, trian- tween 2715 and 2819 m. This is the deepest occurring gular, with a wide base, and up to 0.04 mm in height. species of Parantipathes in Alaska waters. Polyps are about 3 mm in transverse diameter, with 3 polyps/cm. Colonies have a small cup-shaped holdfast, not cov- Photos A) A preserved (in ethanol) whole Paran- ered with coenenchyme, that is relatively easy to detach tipathes sp. A colony (USNM 1070915) collected on from the substrate. Color of the coenenchyme in life and Giacomini Seamount at a depth of 2813 m. B) A pre- in ethanol is light orange. served (in ethanol) whole Parantipathes sp. A colony (USNM 1070914) collected on Giacomini Seamount at Remarks This species differs from Parantipathes euan- a depth of 2819 m. C) A close-up view of the same tha and P. pluma in having smaller spines, and in having specimen in photo B. D) A Parantipathes sp. A col- pinnules in up to eight rows and in alternating groups ony (USNM 1070914; indicated by the white circle) of four, whereas in the other two species there are usu- just before collection at a depth of 2813 m. Next to ally up to six rows and alternating groupings of three the specimen is a colony of N. arbuscula and a large- (although in P. pluma there can very rarely be a group clawed crab (Macroregonia macrochira). E) A Paran- of four pinnules). The length of pinnules is similar to that tipathes sp. A colony (USNM 1070915) just before in P. euantha but much shorter than that in P. pluma. collection at a depth of 2819 m with a N. arbuscula Colonies often grow immediately next to the prim- colony. The distance between the red laser marks in noid Narella arbuscula and host ophiuroid associates. photos D and E is 10 cm. Chapter 2 53 54 CHAPTER 3 Order Scleractinia Corals in the order Scleractinia are known as “stony and pali within the calice. An illustrated key and glos- corals,” but they are also sometimes referred to as “true sary of morphological terms to the 120 genera of extant corals.” Because almost all species known from polar azooxanthellate scleractinians are provided by Cairns latitudes (including all in Alaska) and those from deep and Kitahara (2012). water are non-colonial and cup-shaped, they are collo- The polyp morphology of scleractinians is nearly quially referred to by coral biologists as “solitary cor- identical to that of the actiniarians, or true sea anemo- als” or “cup corals.” All are ahermatypic, that is, they nes, except they lack ciliated lobes on the mesenterial do not construct reefs like their colonial, tropical coun- filaments. Polyps are fully retractile. Oral tentacles are terparts. Ten species of scleractinians from five families conical and arranged hexamerally, reflecting the sym- are known from Alaska waters. The current state of metry of the septa; one tentacle is associated with each taxonomy for the group is excellent. All specimens col- septum. The skeleton (corallum) is typically conical or lected in Alaska should be properly preserved for taxo- discoidal and composed entirely of aragonite except for nomic examination and zoogeographic study. Caryophyllia (Caryophyllia) alaskensis Vaughan, 1941 Conventional terminology is used to describe the and C. (C.) arnoldi Vaughan, 1900, which contain small scleractinian corallum, including basic morphological amounts of low-magnesium calcite (senior author and S. features (Fig. 3-1) and septal insertion patterns (Fig. D. Cairns, unpubl. data). 3-2). Diagnostic features most important for taxonomic resolution include 1) the characteristics of the corallum, theca, and columella, and 2) the arrangement of septa 1 half- S3 S4 S1 1 system S4 system S2 S2 CALICE S4 S3 S3 EXSERT SEPTUM S4 SEPTAL FACE S1 S1 AXIAL SEPTAL PALUS Fossa EDGE FASCICULAR COLUMELLA THECA S1 S1 S4 S5 S3 S S54 S5 S S S2 4 S S 3 S5PEDICEL 1 4 S5 S5 BASE Figure 3-2 A composite cross-sectional diagram of a calice, illustrat- Figure 3-1 ing various septal insertion patterns: upper right system A cutaway diagram of a species of Caryophyllia illus- with three cycles of septa, upper left system with four trating the basic morphological features of an attached, cycles, and lower two systems with various stages of solitary scleractinian (reproduced after Cairns, 1981 and development of the Pourtalès Plan (after Cairns, 1994). Cairns, 1994). The numbers refer to the cycle to which septa belong. Chapter 3 55 Figure 3-3 A map of the North Pacific Ocean showing the distribution of corals in the order Sclerac- tinia () in Alaska waters. Scleractinians are found in all regions of Alaska ex- sediment. This mode is likely an adaptation to the pau- cept the Arctic (Fig. 3-3), and records for most taxa city of hard substrate at the great depths (2370–6328 m) represent the northernmost in the world, perhaps most they occupy in Alaska. notably for the otherwise cosmopolitan cockscomb cup Most taxa are quite small, generally only a few cen- coral (Desmophyllum dianthus) (Esper, 1794). They timeters in diameter even when polyps are fully extend- are an extremely eurybathic fauna in Alaska, occupying ed. Exceptionally large specimens of Javania borealis depths between 17 and 6328 m. The eight taxa found Cairns, 1994 may reach 10 cm in height, with polyps in waters shallower than 2000 m are firmly attached or fully extended. Most taxa are found either singly or fixed and typically grow on rock but are often found on in low-density patches but at least one species, C. (C.) siltstone, particularly on the continental shelf and upper alaskensis, occasionally forms high density patches (278 slope of the eastern Gulf of Alaska. Two taxa found in corals/m2; Stone, 2014). These patches may provide im- very deep water, Fungiacyathus (Bathyactis) marenzel- portant refuge and foraging habitat for juvenile fish and leri (Vaughan, 1906) and Leptopenus discus Moseley, crabs. They are an uncommon bycatch item in longline 1880, are free-living; that is they are not attached to fisheries and stock assessment surveys and are most typ- solid substrate but rather are settled into unconsolidated ically dislodged whole at the holdfast. 56 Professional Paper NMFS 23 Family Caryophylliidae 1. Caryophyllia (Caryophyllia) alaskensis Vaughan, 1941 Description The skeleton is solitary, ceratoid, and al- bia and the Gulf of Alaska (Cairns, 1994). It is one of ways attached by a slender pedicel. The calice is circu- the more common scleractinians collected in Alaska wa- lar to elliptical; the calicular edge is finely serrate. Most ters. The 66 Recent species of Caryophyllia are keyed in specimens are less than 20 mm in height and 21 mm Kitahara et al. (2010). in greater calicular diameter, but exceptional specimens measure up to 27 mm in height and 34 mm in calicular Distribution Widespread and locally common. In diameter. The theca is smooth (porcelaneous) and milky Alaska – eastern Gulf of Alaska including the inside white; the polyp is typically orange. waters of Southeast Alaska, the Aleutian Islands (north Septa are hexamerally arranged in at least four com- of Unalaska Island to east of Attu Island) including Pe- plete cycles (48 septa, 12 pali), but larger specimens may trel Bank and Bowers Ridge, and the eastern Bering Sea have up to five complete cycles of septa (96 septa, 24 continental slope (Fig. 3-4). Elsewhere – southeast to pali), with all variations between. The septal formula the Strait of Georgia, British Columbia, and southwest is S1–2>S3>S4>S5. Septa S1–2 are only slightly exsert to the Commander Islands (Russia) and Sea of Japan (less than 1 mm); pali have very sinuous axial edges. (Republic of South Korea). The fossa is shallow, containing a fascicular columella of 5–15 slender twisted laths. Habitat In Alaska – typically found in patches, some- Remarks Caryophyllia (Caryophyllia) alaskensis is times quite dense (278 corals/m2; Stone, 2014), on hard very similar to C. (C.) arnoldi (see next species descrip- rock in areas of moderate current and at depths be- tion), but C. (C.) alaskensis has a more northern distri- tween 90 and 1397 m. Elsewhere – depths between 103 bution, the two species overlapping off British Colum- and 223 m. Figure 3-4 A map of the North Pacific Ocean showing the distribution of Caryophyllia alaskensis () in Alaska waters. Chapter 3 57 Photos A) A fresh C. (C.) alaskensis specimen tral Aleutian Islands, at a depth of 119 m. D) A calicu- (USNM 1116848) collected in Zhemchug Canyon, east- lar view of a dried C. (C.) alaskensis specimen (USNM ern Bering Sea, at a depth of 909 m. B) A view from 99373) collected near the Semichi Islands, western the top of the same specimen in photo A. C) A C. (C.) Aleutian Islands, at a depth of 95 m. alaskensis specimen collected near Bobrof Island, cen- 58 Professional Paper NMFS 23 2. Caryophyllia (Caryophyllia) arnoldi Vaughan, 1900 Description The skeleton is solitary, robust, ceratoid waters of Southeast Alaska, the western Gulf of Alaska to trochoid, and always firmly attached by a thick pedi- (Prince William Sound), the Gulf of Alaska Seamount cel. The calice is circular to elliptical; the calicular edge Province (Giacomini Seamount), and northwest of Un- is finely serrate. The largest known specimen is 17 mm alaska Island to Amchixtam Chaxsxii in the Aleutian Is- in height and 16 mm in greater calicular diameter. The lands (Fig. 3-5). Elsewhere – distributed south to Queen theca is white, bearing granular costae, whereas the pol- Charlotte Islands (British Columbia), Strait of Juan de yp is typically light orange to orange. Fuca (Washington), Cordell Bank to off San Diego (Cal- Septa are hexamerally arranged in four complete ifornia) and the Galapagos Islands (Ecuador). cycles (48 septa, 12 pali) with a septal formula of S1– 2>S3>S4. Septa S1–2 are moderately exsert (over 1 Habitat In Alaska – typically found in patches on mm); pali have extremely sinuous axial edges. The fossa hard rock, including cobbles, in areas of moderate is shallow, containing a fascicular columella of 5–25 current and at depths between 21 and 1702 m. Else- twisted laths, occasionally fused into a massive structure. where – depths between 75 and 657 m. Co-occurs in dense mats with the zoantharian Mesozoanthus sp. in Remarks As reported by Cairns (1994), Caryophyl- Prince William Sound. lia (Caryophyllia) arnoldi is similar to C. (C.) alasken- sis but differs in having a more robust, denser skeleton with a thicker pedicel; having granular costae (not Photos A) A fresh C. (C.) arnoldi specimen (USNM smooth theca); never having more than 48 septa; having 1407047) collected in the West Arm of Glacier Bay, more exsert septa; and having wider S4, almost as wide Southeast Alaska at a depth of 292 m. B) A side view of as their S3. the same specimen in photo A. C) The same specimen in photo A, dried. D) Two C. (C.) arnoldi, tentacles fully Distribution Widespread and locally common. In extended, photographed on bedrock in the central Aleu- Alaska – eastern Gulf of Alaska including the inside tian Islands at a depth of 862 m. Figure 3-5 A map of the North Pacific Ocean showing the distribution of Caryophyllia arnoldi () in Alaska waters. Chapter 3 59 60 Professional Paper NMFS 23 3. Crispatotrochus foxi (Durham and Barnard, 1952) Description The skeleton is solitary, trochoid, and al- Distribution Rare. In Alaska – known from only four ways firmly attached by a robust pedicel. The calice is specimens collected at three locations between Umnak circular to slightly irregular in shape. Most specimens Island and northern Amchitka Pass in the Aleutian Is- are less than 19 mm in height and 15 mm in greater lands (Fig. 3-6). One of the four specimens could be calicular diameter. Granular costae are slightly ridged confirmed only to genus but is very likely Crispatotro- in the upper half to third of skeleton. The corallum is chus foxi (Fig. 3-6). Elsewhere – known only from the white; the polyp is orange to pale orange. type locality off San Miguel Island, Channel Islands, Septa are hexamerally arranged in five complete cycles southern California. (96 septa) with a septal formula of S1–2>S3>>S4>S5. Septa S1–2 are only slightly exsert and have straight ax- Habitat In Alaska – typically found on cobbles at ial edges. Pali are absent. The fossa is moderately deep, depths between 135 and 199 m except one specimen containing a small fascicular columella consisting of 4–7 (the one confirmed only to genus) at a depth of 702 m. broad, twisted laths that are usually laterally fused to Found in association with the scleractinians Caryophyl- one another. lia (Caryophyllia) alaskensis and Javania spp. Elsewhere – 82 m depth. Remarks This species can be distinguished from all other Alaska scleractinians by having fascicular colu- Photos A) A calicular view of a dried C. foxi speci- mella and lacking pali; it is otherwise similar to Caryo- men (USNM 19210) collected in the eastern Aleutian phyllia except for its lack of pali. It is rarely collected, Islands at a depth of 199 m. B) Fresh C. foxi specimens so if encountered should be properly preserved and sent (USNM 1482090) collected in the eastern Aleutian Is- to a museum, such as the National Museum of Natural lands at a depth of 135 m. History (Smithsonian Institution), for verification and archival. Figure 3-6 A map of the North Pacific Ocean showing the distribution of Crispatotrochus foxi () in Alaska waters. One specimen (♦) could be confirmed only to the genus. Chapter 3 61 62 Professional Paper NMFS 23 4. Desmophyllum dianthus (Esper, 1794) Description The skeleton is solitary, ceratoid, often most common and widespread living scleractinians, with a flared calice (trumpet-shaped), and always at- and one of only a dozen species worldwide that is truly tached by a robust pedicel. The calice is circular, slight- cosmopolitan in distribution. Even though solitary, it is ly elliptical, or even scalloped (if large). The Alaska considered to be a framework-forming species (Roberts specimen measures up to 5 cm in height and 4×6 cm et al., 2009). in calicular diameter, although larger specimens are known from New Zealand fjords. The theca is granular Distribution Rare. In Alaska – known from only a sin- and the C1–3 are usually slightly ridged in the upper gle specimen (representing the northernmost in the world) half of the skeleton. The corallum is white, light brown collected north of Dixon Entrance, eastern Gulf of Alaska or grey; the polyps are light orange. near the border with British Columbia (Fig. 3-7). Else- Septa are hexamerally arranged in up to six cycles (192 where – widespread, cosmopolitan (found in all world septa), according to the formula S1–2>S3>>S4>S5>S6; oceans). septa S6 are complete only in the largest of specimens. Septa S1–2 are often highly exsert, up to 10 mm, and Habitat In Alaska – the single specimen was col- have straight axial edges. Pali are absent. The fossa is lected at a depth of 398 m and was attached to the deep and slender, usually lacks a columella, but may heavily calcified base of a large red tree coral (Prim- have a rudimentary papillose element that is usually hid- noa pacifica Kinoshita, 1907). Elsewhere – extremely den from view in an intact corallum. eurybathic. Considered a deep-sea emerged species in southern Chilean fjords (to 8 m depth; Försterra and Remarks Morphologically, Desmophyllum dianthus Häussermann, 2003) and Milford Sound, New Zea- is unique and perhaps the simplest of the scleractinian land (to 25 m depth; NMNH 94068 and 94072), but corals in having the combination of being solitary (not found to depths as great as 2200 m on the Charles- colonial), lacking pali, columella, and any other unique ton Bump off the southeastern Atlantic Coast of the features. Its simplicity may explain why it is one of the U.S. Figure 3-7 A map of the eastern Gulf of Alaska showing the distribution of Desmophyllum dianthus () in Alaska waters. Chapter 3 63 Photos A) A large dried fragment of D. dianthus naris. B) A lateral view of two dried D. dianthus speci- (USNM 1482091) collected in the eastern Gulf of mens (USNM 36545) collected off the coast of Chile Alaska at a depth of 398 m. The specimen is partially at a depth of 821 m. C) A calicular view of the same encrusted with the demosponge Poecillastra tenuilami- specimen in photo B. 64 Professional Paper NMFS 23 Family Dendrophylliidae 5. Balanophyllia (Balanophyllia) elegans Verrill, 1864 Description The skeleton is solitary, cylindrical to columella that fuses to the axial edges of many of the trochoid in shape (often short and squat), with a thick septa. attached pedicel and polycyclic base. Large specimens are up to 18 mm in height and 17 mm in greater calicu- Remarks Among the Scleractinia known from Alaska lar diameter. The theca is vermiculate (spongy, porous) waters, Balanophyllia (Balanophyllia) elegans is unique and rough to the touch (called a synapticulotheca), but in being the only dendrophylliid in the region, and thus on the lower third to half of the corallum, the synaptic- the only species to have a porous skeleton and septa ulotheca is covered by a smooth thin epitheca, on which arranged in a Pourtalès Plan. Although quite common encrusting organisms are often attached. The skeleton is off the west coast of the continental United States and white, but shallow-water (<30 m) specimens are often Canada, it is quite rare in Alaska waters but found in encrusted with purple coralline algae (Lithothamnion small patches where present. spp.; see photo D); polyps are bright orange or reddish- orange. Distribution Rare. In Alaska – known from only two The septa are distinctively arranged in a stellate specimens (representing the northernmost in the world) pattern (the Pourtalès Plan) in four to five cycles; the collected at two locations in the eastern Gulf of Alaska last cycle is never complete, resulting in 48–88 septa. (Fig. 3-8) but also known from photographs collected Axial edges of S1–2 are smooth, but those of higher during shallow-water resource surveys (see photos cycle septa are laciniate. Pali are absent. The fossa is D–F). Elsewhere – Vancouver Island, British Columbia, of moderate depth, containing an elongate, spongy to southern California. Figure 3-8 A map of the North Pacific Ocean showing the distribution of Balanophyllia (Balanophyllia) elegans () in Alaska waters. Chapter 3 65 Habitat In Alaska – specimens have been collected at ka at a depth of 22 m. B) A calicular view of a dried B. 22 m depth but observed as shallow as 15 m (Harris3). (B.) elegans specimen (USNM 1482079) collected in the Grows on bedrock and boulders in areas of moderate eastern Gulf of Alaska at a depth of 22 m. C) A cali- to strong current. Elsewhere – grows on bedrock, boul- cular view of a dried B. (B.) elegans specimen (USNM ders, cobbles, and pebbles from tide pools to depths of 92623) collected north of the Farallon Islands, Califor- 177 m. Durham (1947) lists the depth range to 296 m nia at a depth of 53 m. D) A small patch of Balanophyl- but provides no documentation for the record. lia (B.) elegans photographed near Cape Ulitka, Noyes Island in the eastern Gulf of Alaska at a depth of about Photos A) Five dried B. (B.) elegans specimens 15 m. E) A close-up view of several corals in the same (USNM 1482079) collected in the eastern Gulf of Alas- patch (with tentacles fully extended). F) A close-up view of a coral in the same patch (with tentacles retracting). 3Harris, D. K. 2018. Personal observ. Alaska Dep. Fish Game, Juneau, Photos D–F are courtesy of D. K. Harris, Alaska De- AK 99801. partment of Fish and Game. 66 Professional Paper NMFS 23 Family Flabellidae 6. Flabellum (Flabellum) oclairi Cairns, sp. nov. Synonymy Remarks Cairns (1994, 1995) suggested that this spe- cies was very similar to the ceratoid form of the Ant- Flabellum (F.) sp. A Cairns, 1994:28–29, plates 10a–b. arctic species Flabellum (F.) impensum Squires, 1962, even though that would have postulated a broadly Flabellum impensum Cairns, 1995:100. disjunct distribution pattern. Flabellum (F.) impensum is reported from only the Southern Hemisphere, prin- Flabellum sp. Stone and Shotwell, 2007:105 (listed).— cipally in Antarctic seas, but also occurs in the South Stone and Cairns, 2017:3 (listed). Atlantic (Scotia Sea, Drake Passage, and off the coast of Argentina) and South Pacific (New Zealand) at depths Material examined Holotype: Station 69RD3, Na- of 46–2260 m (Cairns, 1982, 1994, 1995). Now that tional Museum of Canada (Canadian Museum of Na- additional specimens are available for examination, it ture), Invertebrate Collection, Ottawa, 1982-1492. appears that the Alaska specimens are indeed very simi- Paratypes: ex AB02-0123B, scuba collection, central lar but morphologically distinct from F. (F.) impensum, Aleutian Islands, 51°52.20´N, 179°49.85´W, 27 m and thus represent an undescribed species. It differs in depth, 17 July 2002, four specimens, USNM 1011163; 1) having a more circular calice—GCD:LCD about 1.2 RV Vesteraalen, haul station 5, eastern Aleutian Is- versus up to 2.8 in F. (F.) impensum, 2) achieving a full lands, 52°40.76´N, 169°06.61´W, 110 m depth, 21 complement of S5 at an earlier size (17 mm GCD ver- May 2001, six specimens, USNM 1010473; RV Alas- sus 19 mm GCD), and 3) having straight axial edges of kan Leader, haul station 35, eastern Aleutian Islands, the larger septa—those of F. (F.) impensum are sinu- 53°04.55´N, 170°13.88´W, 407 m depth, 4 June 2002, ous. Also, F. (F.) impensum achieves a much larger size one specimen, USNM 1011169; RV Dominator, (128 mm GCD) and eventually inserts a full sixth cycle haul station 4, eastern Aleutian Islands, 52°47.01´N, of septa (Cairns, 1982). Although these differences are 168°41.98´W, 109 m depth, 23 May 1996, six speci- minor, they do appear to be consistent. mens, USNM 98509; RV Dominator, haul station 218, western Aleutian Islands, 52°05.32´N, 172°42.99´E, Distribution Uncommon. In Alaska – throughout the 113 m depth, 4 August 1997, eight specimens, USNM Aleutian Islands from Unalaska Island to south of Attu Island (Fig. 3-9). Elsewhere – not reported. 99369. Habitat Grows on bedrock and other hard substrates Type locality Station 69RD3: 51°25.50´N, 179°15.20´E at depths between 27 and 507 m, although most re- (Kirilof Bay, northern Amchitka Island), 30 m depth, 9 cords are shallower than 100 m. Occasionally 3–4 cor- October 1969. als may be fused together along their thecal edges (see photo C). Description The skeleton is solitary, ceratoid to tro- choid, and always attached by a thick pedicel (3.6–4.1 Etymology Named in honor of Dr. Charles (Chuck) mm in diameter) that broadens to a slightly larger scal- O’Clair, renowned marine biologist and naturalist of loped basal plate up to 5.5 mm in diameter; the basal Alaska and the Pacific Northwest, who so carefully plate contains 12 protosepta. The calice is slightly ellip- saved the holotype he collected near Amchitka Island in tical, with a greater calicular diameter (GCD) to lesser the Aleutian Islands in 1969. calicular diameter (LCD) of about 1.2. The largest spec- imen is 38.5 mm in height and 26×22 mm in calicular Photos A) A calicular view of the F. (F.) oclairi sp. diameter. The theca is milky white and porcelaneous; nov. holotype (NMCIC 1982-1492; dried) collected the polyps are bright orange. near Amchitka Island in the western Aleutian Islands The septa are hexamerally arranged in five com- at a depth of 30 m. B) A lateral view of the F. (F.) plete cycles in larger specimens with a formula of S1– oclairi sp. nov. holotype. C) A calicular view of F. (F.) 2>S3>>S4>S5 (S5 rudimentary). The fourth cycle of 48 oclairi sp. nov. paratypes (USNM 1011163; dried) col- septa is attained at a GCD of about 11 mm and the fifth lected near Semisopochnoi Island, central Aleutian Is- at a GCD of about 17 mm. The septa are not exsert, thin lands, at a depth of 27 m. D) Flabellum (F.) oclairi and quite fragile, having straight axial edges. Pali are sp. nov. (at left in photo C) photographed immediately absent; the fossa is deep; a columella is absent. after collection. Chapter 3 67 Figure 3-9 A map of the Aleutian Islands showing the distribution of Flabellum oclairi Cairns, sp. nov. () in Alaska waters. 68 Professional Paper NMFS 23 7. Javania borealis Cairns, 1994 Description (Adapted from Cairns, 1994) The skel- Distribution In Alaska – Aleutian Islands (north of eton is solitary, trochoid, and attached by a stereome- Unalaska Island to northeast of Attu Island) includ- reinforced pedicel up to 11 mm in diameter. The calice ing Petrel Bank (Fig. 3-10). Elsewhere – the only other is elliptical and slightly serrate. The largest specimen is known record is from the Sea of Japan (west of Hok- 35 mm in height and 36×28 mm in calicular diameter kaido). (greater calicular diameter to lesser calicular diameter of 1.3). The theca is smooth, white, and porcelaneous; Habitat In Alaska – grows on bedrock and other hard the polyps are typically white or pale orange at shallow substrate at depths between 17 and 1266 m. Elsewhere depths but orange in deeper waters. – at depth of 347 m. Septa are hexamerally arranged in five complete cycles (96 septa) with a formula of S1–2>S3>S4>>S5. Photos A) A calicular view of a dried J. borealis speci- Septa are not exsert, thin and quite fragile, and have men (USNM 1011164) collected northwest of Adak slightly sinuous axial edges. Pali are absent; the fossa is Island, central Aleutian Islands, at a depth of 150 m. deep; a columella is absent. B) A fresh J. borealis specimen (USNM 1086634) col- lected west of Bobrof Island, central Aleutian Islands, Remarks Javania borealis is distinguished from the at a depth of 17 m. C) A lateral view of the same speci- only other Alaska species, J. cailleti (Duchassaing and men in photo A. D) A patch of J. borealis in the central Michelotti, 1864), by having five (not four) cycles of sep- Aleutian Islands at a depth of 1266 m. E) A close-up ta. Otherwise, it closely resembles Flabellum (Flabellum) view of a patch of J. borealis in the central Aleutian Is- oclairi sp. nov. in many features, but differs in having a lands at a depth of 1258 m. The distance between the stereome-reinforced pedicel (characteristic of the genus) red laser marks in photos D and E is 10 cm. and sinuous axial septal edges. The ten Recent species of the genus Javania are keyed by Cairns (2004). Figure 3-10 A map of the North Pacific Ocean showing the distribution of the Javania borealis () in Alaska waters. Chapter 3 69 70 Professional Paper NMFS 23 8. Javania cailleti (Duchassaing and Michelotti, 1864) Description The skeleton is solitary, ceratoid, and the world) collected from two locations in the central firmly attached by a robust stereome-reinforced pedicel Aleutian Islands (including Amchixtam Chaxsxii) and a up to 14 mm in diameter; the pedicel may be curved. single location in the Gulf of Alaska Seamount Province Specimens measure up to 46 mm in height; the calice (Giacomini Seamount), but also observed on video foot- is elliptical and up to 31 mm in greater diameter; the age collected on Welker, Quinn, Giacomini, and Mur- upper corallum is distally flared. The theca is smooth, ray Seamounts (Fig. 3-11). Elsewhere – widespread and white, and porcelaneous; the polyps are typically light common (Cairns, 1994). Pacific Ocean records include orange or light pink. a single record from the Strait of Juan de Fuca (Brit- Septa are hexamerally arranged in four complete cy- ish Columbia), Hawaii, Hokkaido (Japan), and New cles (48 septa) with a formula of S1–2>S3>S4; the S1–2 Zealand. Also reported from the Southeast coast of the is usually highly exsert. The axial edges of larger septa United States, Gulf of Mexico and Caribbean Sea, off both coasts of South America, and the Azores. are straight and their lower edges fuse into a rudimen- tary solid columella deep in the fossa. Habitat In Alaska – grows singly and occasionally in small clusters on fractured bedrock, basalt, and cobbles Remarks This species is compared to J. borealis in the at depths between 150 and 1799 m. But observed much previous species account. It is one of four species in the deeper on video footage collected on Welker (1783–2635 genus to have only four cycles of septa (Cairns, 2004). m), Quinn (2100–2373 m), Giacomini (2431–2476 m), and Murray (1787 m) Seamounts. Elsewhere – grows on Distribution Uncommon. In Alaska – known from hard rock at depths between 78 and 1809 m. only three specimens (representing the northernmost in Figure 3-11 A map of the North Pacific Ocean showing the distribution of Javania cailleti () in Alaska waters. Chapter 3 71 Photos A) A lateral view of dried J. cailleti specimens a depth of 1799 m. D) A lateral view of the same speci- (USNM 62021) collected off northeastern Little Abaco men in photo C. E) Javania cailleti photographed on Island, Bahamas, North Atlantic Ocean, at a depth of Giacomini Seamount at a depth of 1804 m. F) Javania 288 m. B) A calicular view of the same specimens in cailleti photographed on Quinn Seamount at a depth of photo A. C) A preserved (in ethanol) J. cailleti specimen 2350 m. The distance between the red laser marks in (USNM 1075808) collected on Giacomini Seamount at photos E and F is 10 cm. 72 Professional Paper NMFS 23 Family Fungiacyathidae 9. Fungiacyathus (Bathyactis) marenzelleri (Vaughan, 1906) Description The corallum is solitary, discoidal, free- off central California at a depth of 4100 m (Flint et al., living (not attached), and has a flat base covered with 2007). finely serrate, ridged costae. Coralla measure up to 27 mm in diameter. The skeleton is white to light brown; Distribution Uncommon. In Alaska – known only the polyps are pale white to light pink. from the deep continental slope region of the central Septa are hexamerally arranged in four cycles; only and western Gulf of Alaska including Derickson Sea- the S1–2 is independent, reaching the columella. Pairs mount. But also observed on video footage collected of S3 fuse with adjacent S2 and pairs of S4 fuse to ad- on Quinn Seamount at depths between 2370 and 2371 jacent S3, forming triangular canopies at the point of m (Fig. 3-12). Elsewhere – widespread and common fusion. Upper edges of the septa bear numerous elon- (Cairns, 1994); Pacific Ocean (Japan and the Kurile Is- gate trabecular spines; the septal faces are ridged. The lands, California and Baja Mexico, and New Zealand) columella is circular, consisting of a mass of trabecular and the Atlantic Ocean from Greenland to Antarctica. spines, similar to those on axial septal edges. Habitat In Alaska – free-living (not attached) in areas Remarks Fungiacyathus (Bathyactis) marenzelleri is of soft sediment (fine sand and silt) with low current in one of two species known from Alaska waters that has very deep water (2370–6328 m). The latter record is the an unattached corallum, the other being Leptopenus dis- deepest known for any scleractinian species worldwide. cus. It is distinguished from Leptopenus by having typical Observed on video footage in patches up to 4 corals/ (non-bifurcating) higher cycle septa, and septa that cor- m2. Elsewhere – found in areas of soft sediment in deep respond to their costae (in Leptopenus they alternate). water but eurybathic (300–4882 m). The shallowest re- The reproductive ecology of this species has been studied cords are from Antarctica. from specimens collected in the Northeast Pacific Ocean Figure 3-12 A map of the North Pacific Ocean showing the distribution of Fungiacyathus marenzelleri () in Alaska waters. Chapter 3 73 Photos A) A calicular view of two dried F. (B.) the same specimen in photo B during collection. E) A marenzelleri specimens (USNM 98974) collected west patch of F. (B.) marenzelleri in situ. Specimen USNM of the Channel Islands (California) at a depth of 4100 1081142 is visible within the black circle. F) Fungiacy- m. B) A calicular view of a dried F. (B.) marenzelleri athus (B.) marenzelleri photographed on Quinn Sea- specimen (USNM 1081142) collected on Derickson mount at 2371 m. The distance between the red laser Seamount at a depth of 4704 m. C) A ventral view of marks in photos D–F is 10 cm. the same specimen in photo B. D) A photograph of 74 Professional Paper NMFS 23 Family Micrabaciidae 10. Leptopenus discus Moseley, 1880 Description The corallum is solitary, discoidal, and gether only by their tissue. This species was originally free-living (not attached); the largest known specimen described as L. irinae Keller, 1977, which is now con- is 25 mm in diameter. The base is flat, and the costae sidered a junior synonym of L. discus. alternate in position with the septa with a row of large pores occurring between each adjacent costa and re- Distribution In Alaska – known only from two speci- sulting in an extremely fragile skeleton. The skeleton is mens collected on the deep continental slope in the white; the color of the polyps in life is unknown. western Gulf of Alaska (Fig. 3-13). Elsewhere – wide- Septa S1–2 extend from the columella to the calice spread but rarely collected, especially intact. Known edge, each septum bearing a series of spines on its upper from off Washington and southern California in the edge. Septa S3 also originate at the columella but appear North Pacific Ocean and near Cuba in the North Atlan- to bifurcate repeatedly one to three times, resulting in tic Ocean. relatively short septa at the calicular margin. Altogether there are 48–72 septa in an adult corallum. The colu- Habitat In Alaska – collected at depths between 4820 mella is a spinose central mound. and 5000 m. Elsewhere – collected at depths between 2842 and 4655 m. Remarks All micrabaciids, including Leptopenus, are distinctive in having septa that alternate with costae and Photos A) A dorsal view of a dried L. discus specimen a unique septal arrangement in which the S3 appear to (USNM 93938) collected on Fieberling Guyot, west of bifurcate repeatedly. They are all quite fragile, usually the Channel Islands (California) at a depth of 4100 m. damaged during collection, and their skeleton is held to- B) A ventral view of the same specimen in photo A. Chapter 3 75 Figure 3-13 A map of the North Pacific Ocean showing the distribution of Leptopenus discus () in Alas- ka waters. 76 CHAPTER 4 Order Zoantharia Corals in the order Zoantharia (Class Anthozoa, Sub- hexactinellid sponges and the other (Zibrowius cf. class Hexacorallia) are generically referred to as “zo- ammophilus) on octocorals and occasionally antipath- anthids” but the correct term is simply “zoanthar- arians. We note that this group of corals is in des- ians.” They are often mistaken for actiniarians (true perate need of taxonomic attention. All encountered sea anemones) so are largely ignored by marine biolo- specimens should be carefully collected, cataloged, and gists and seafloor ecologists with the goal of studying preserved in ethanol (minimum 70%) or deep frozen more traditional corals (i.e., those with a noticeable for both morphological and molecular phylogenetic hard skeleton made of calcium carbonate). They are study. also often overlooked given their small size and rather Zoantharians are colonial corals in which the polyps inconspicuous habitus and there is even debate as to arise from a basal tissue, the coenenchyme, which can whether they are actually corals. We include them in form a mat or stolon containing gastrodermal canals. this guide because they appear to be more common Each polyp has two rings of tentacles and they are never than previously thought and are ecologically important pinnate. Colonies very rarely have an intrinsic skeleton in habitats where they occur. They have been poorly but most species incorporate sand, sponge spicules, and documented in Alaska waters until recently. Here we other debris particles into the thick body wall or me- recognize four distinct taxa from two families (Epizo- sogloea. The pharynx is flattened and contains a single anthidae and Parazoanthidae) in Alaska waters, two siphonoglyph. Mesenteries are numerous but typically of which are epizoic; one (gen. nov., sp. nov.) on with weak musculature. B A C Figure 4-1 Illustrations of typical corals in the order Zoantharia showing diagnostic morphological features. (A) A planular-section of a polyp. (B) A cross-section of a zooid showing the septal arrangement. (C) The view of a horizontal-section through a colony. Adapted from Ryland and Lancaster (2003). 5 cm 2 cm 2 cm Chapter 4 77 Diagnostic morphological features (Fig. 4-1) of the S. lucifica the gonads are located on the macrocnemes zoantharian fauna have traditionally included 1) the just below the actinopharynx and are ripe during the anatomy and position of the sphincter muscle, 2) the summer months. The zoanthella-type larvae are retained relationship between the number of septa and polyp col- within the body wall until they are well developed and umn diameter, and 3) the complement and morphom- are commonly observed in the lumens of the tentacles etry of the cnidom (Ryland and Lancaster, 2003). Re- (Cutress and Pequegnat, 1960). These characteristics in- cently, DNA information based on multiple molecular dicate that the larvae do not travel far from the parent markers and ecological parameters have played a major and likely account for the highly contagious distribution role in determining phylogenetic relationships among displayed by these corals. zoantharians (Sinniger et al., 2010, 2013). None of the Alaska species are zooxanthellate nor do Most species of zoantharians are reported to be gono- they appear to emit bioluminescence when disturbed in choric (either male or female) and the reproductive ecol- situ. They are uncommon but can be abundant where ogy for at least one Pacific Ocean species, Savalia lucifica they do occur. They are found in the Gulf of Alaska, the (Cutress and Pequegnat, 1960) (originally described as Gulf of Alaska Seamount Province, the eastern Aleutian Parazoanthus lucificum Cutress and Pequegnat, 1960), Islands, and the eastern Bering Sea (Fig. 4-2) and are has been well studied (Cutress and Pequegnat, 1960). In eurybathic fauna ranging in depth from 87 to 2511 m. Figure 4-2 A map of the North Pacific Ocean showing the distribution of corals in the order Zoan- tharia () in Alaska waters. 78 Professional Paper NMFS 23 Family Epizoanthidae 1. Epizoanthus scotinus Wood, 1957 Description The original description of this species and was a globular mass measuring about 6.3×4.4 cm. by Wood (1957) is poorly documented. Additionally, The colony consists of 14 tubular polyps of variable but Epizoanthus is one of the most difficult genera to iden- quite large size (up to 24 mm in height and 10 mm in tify based on external appearance with the exception of width, contracted). Polyps are variably crowded and species with very unique morphologies or ecology. Cur- with a relatively thick coenenchyme that is obviously rently, the most reliable identifications rely on simple speckled with black foreign material, likely sand par- DNA analyses or histological examination. However, in ticles. The color of the colony preserved in ethanol is a the case of E. scotinus, individual polyps are up to 5 cm uniform golden-light brown. in length but colonies forming large irregular mats may cover several square meters or more. Polyps are moder- Remarks This species is not known to be epizoic but ately crowded and each possesses approximately 30 rel- rather grows directly on bedrock or large boulders and atively short (up to 22 mm in length), stout but pointed typically does not incorporate particles into the ecto- tentacles. Color in life is a golden-orange with a notably derm or mesogloea. lighter oral disc rim and tentacles. The coloration of the larger polyps and the better-developed coenenchyme are Distribution Rare. In Alaska – known from only a good characteristics to distinguish this species from the single specimen collected near Unalaska Island in the potentially sympatric Mesozoanthus lilkweminensis Re- eastern Aleutian Islands (Fig. 4-3). Elsewhere – un- imer and Sinniger, 2010. common but locally abundant; widespread through- The only known Alaska specimen (photos A and B) out the eastern North Pacific Ocean (British Columbia was attached to an aggregate of fractured bivalve shell to Cordell Bank, California). According to Lamb and Figure 4-3 A map of the Aleutian Islands showing the distribution of Epizoanthus scotinus () in Alas- ka waters. Chapter 4 79 Hanby (2005) this species is known from Siberia to Photos A) A large portion of a preserved (in ethanol) southern California. E. scotinus colony (CAS 168222) collected in Akutan Pass, eastern Aleutian Islands, at a depth of 116 m. B) Habitat In Alaska – attaches to cobbles, pebbles, and An alternate view of the same specimen in photo A. C) bivalve shells in areas with moderate to strong currents A large E. scotinus colony photographed in Weynton at depths near 116 m. Elsewhere – often found in near- Passage, central British Columbia, at a depth of about shore and coastal habitats on vertical or undercut bed- 25 m. D) A close-up view of the polyps on the same rock with moderate currents. Colonies are found in the colony in photo C. Photos C and D are courtesy of Neil low intertidal zone (Lamb and Hanby, 2005) to depths McDaniel. of 149 m (off Oregon). 80 Professional Paper NMFS 23 Family Parazoanthidae 2. Mesozoanthus sp. Description This species is known in Alaska from reaster swifti), and the spiny red sea star (Hippasteria only a few small specimens collected in Pribilof Can- phrygiana) in the eastern GOA. yon (eastern Bering Sea) and observations of similar specimens at 12 locations in the eastern and central Distribution Uncommon. In Alaska – the GOA from Gulf of Alaska. Colonies occur as clusters of 6–50 pol- Dixon Entrance to south of the Kenai Peninsula (includ- yps that form nearly continuous mats covering 5 m2 ing Prince William Sound) and Pribilof Canyon in the or more. Polyps are relatively large, up to 32 mm in eastern Bering Sea (Fig. 4-4). Elsewhere – not reported. length and 12 mm in diameter when contracted, and often increase slightly in width from the base to the Habitat Occurs principally on bedrock including ver- distal section of the polyp. Polyps have 24–27 long tical and overhanging walls but also on cobbles and (up to 14 mm in length), slender tentacles that are the pebbles at depths between 123 and 625 m. Co-occurs in same color as the polyp. Polyps are orangish-brown dense mats with the scleractinian Caryophyllia arnoldi in color, sometimes with obvious ribbing when con- in Prince William Sound. tracted, and often speckled with black sand particles. As with other zoantharians, the oral disc is a lighter Photos A) A Mesozoanthus sp. specimen collected color than the rest of the polyp. in Pribilof Canyon, eastern Bering Sea, at a depth of 208 m. B) An alternate view of the same specimen in Remarks There are only two accepted species in the photo A. C) A Mesozoanthus sp. specimen collected in genus: Mesozoanthus fossii Sinniger and Haussermann, Pribilof Canyon at a depth of 204 m. The colony is at- 2009 and Mesozoanthus lilkweminensis (WoRMS Edi- tached to pebbles that were incorporated into the base torial Board, 2022). The former species is from shal- of the hexactinellid sponge. D) Small Mesozoanthus sp. low water in Chilean fjords and the latter is from the colonies (including the colony in photo A) attached to coastal waters of British Columbia at depths between pebbles in Pribilof Canyon at a depth of 208 m. E) Nu- 37 and 43 m. Based on geographical and bathymetrical merous Mesozoanthus sp. colonies attached to bedrock differences between the two described species and the and forming a nearly continuous mat on Portlock Bank one listed here, we suspect that the Alaska specimens in the western GOA at a depth of 197 m. An immature may be one or more undescribed species. All specimens redbanded rockfish (Sebastes babcocki) perches on the suspected of being this taxon should be collected and corals. F) Mesozoanthus sp. colonies associated with de- properly preserved for expert identification. mosponges (Latrunculia austini) in the Edgecumbe Pin- We initially thought this species was epizoic since we nacles Marine Reserve in the eastern GOA at a depth of observed it growing on live demosponge Poecillastra 161 m. G) A close-up view of the same colonies in pho- tenuilaminaris (see photo I) in the eastern GOA and on to F. H) Large mats of Mesozoanthus sp. colonies in the live hexactinellid sponge (Aulosaccus sp.) in the eastern Edgecumbe Pinnacles Marine Reserve at a depth of 165 Bering Sea (see photo C). Upon closer inspection, how- m. A lingcod (Ophiodon elongatus) perches on the cor- ever, we determined that in both cases the zoantharians als. I) The same colonies in photo H showing zoanthar- were actually growing on rock fragments or pebbles ians growing on fragmented rock that has fallen from that were in turn attached directly to the sponges. above onto the demosponge Poecillastra tenuilaminaris. This species is preyed upon by the dorid nudibranch The distance between the red laser marks in photos D (Peltodoris lentiginosa), the gunpowder star (Gephy- and F–I is 10 cm. Chapter 4 81 82 Professional Paper NMFS 23 2. Mesozoanthus sp. (continued) Figure 4-4 A map of the North Pacific Ocean showing the distribution of Mesozoanthus sp. () in Alaska waters. Chapter 4 83 3. Zibrowius cf. ammophilus Preliminarily molecular DNA analysis based on the mi- but occasionally zoantharians covered only a portion tochondrial cytochrome oxidase c subunit 1 and 16S of a living host (see photo F on next page). We suspect markers and the 8S and internal transcribed spacer nu- that the process that occurs for this zoantharian settling clear markers indicated that specimens recently collect- on and then overtaking its host is very similar to the ed on seamounts in the Gulf of Alaska Seamount Prov- process described for Savalia lucifica and Muricea cali- ince are likely a novel species, very closely related to fornica Aurivillius, 1931 (see Cutress and Pequegnat, Zibrowius ammophilus Sinniger et al., 2013. Additional 1960), where larvae settle directly on living tissue rath- analyses are underway and, if it is a new species, a de- er than on bare areas of skeleton and then overgrow scription will be prepared (Sinniger Harri4). The genus and kill the host colony in the process. All the host cor- Zibrowius was established to accommodate a unique als we observed have strong holdfasts and persistent group of zoantharians on Cross Seamount near Hawaii skeletons after death providing excellent habitat for the (Sinniger et al., 2013). There are three known species in zoantharians, but we observed very few coral skeletons the genus, all from seamounts and all characterized by a without the zoantharians. This evidence indicates that well-developed coenenchyme that completely covers the the zoantharians may be more parasitic than epizoic, host and has sand incrustations in the ectoderm. This is a suggestion made previously for Hawaiian seamount the first Zibrowius species documented from the north- Parazoanthidae (Sinniger et al., 2013). ern North Pacific Ocean. Swarms of euphausiids have been observed in the im- mediate vicinity of larger colonies, and colonies harbor Description This is strictly an epizoic species that can similar associated fauna (ophiuroids and the sternostyl- form large encrustations covering host coral colonies id crab Sternostylus iaspis) that the host colony would as large as 70 cm in height by 94 cm in width. Polyps have in life. are relatively small (up to 9 mm in length and 5 mm in diameter), variably crowded, and with a very thin coe- Distribution Uncommon but locally abundant. In nenchyme. Each polyp has approximately 20 pointed Alaska – known only from seamounts in the Gulf of tentacles that are relatively shorter and stouter than Alaska Seamount Province (Welker, Pratt, Quinn, Gia- those of other Alaska species. The scapus (or column) comini, Murray, Patton, and Marchand Seamounts; Fig. is somewhat transparent and lighter in color than the 4-5). Elsewhere – not reported. capitulum in such a way that the mesenteries are often visible inside the polyp wall. Color of colonies in life Habitat In Alaska – encrusts octocoral, antipatharian, is uniform and a bright rusty orange to golden-orange; and hexactinellid sponge skeletons at depths between this variation in coloration is likely due to differential 745 and 2511 m. Rarely grows on rock alone and likely artificial lighting. only when the stolon has overgrown the host colony or Remarks The species completely overgrows or en- skeleton. crusts octocoral colonies, mostly the primnoids Para- stenella ramosa (Studer, 1894) and Parastenella gym- Photos A) A whole preserved (in ethanol) Z. cf. am- nogaster Cairns, 2007, occasionally the gorgoniids Cal- mophilus colony (NMNH 1661883) collected on Quinn listephanus pacificus Nutting, 1912 and Callistephanus Seamount at a depth of 895 m. The colony completely simplex (Nutting, 1909) and keratoisidids (Isidella ten- encrusted a whole P. ramosa skeleton. B) A preserved taculum and Keratoisis sp. D), and rarely the antipath- (in ethanol) branch of Z. cf. ammophilus (NMNH arian Lillipathes sp. A and fossil hexactinellid (likely 1661884) collected on Giacomini Seamount at a depth Chonelasma oreia). Small colonies (less than five pol- of 745 m. The colony completely encrusted a P. ramosa yps) have been observed on the octocoral Paragorgia skeleton. C) A close-up view of the same specimen in sp. B. photo B. D) A zoomed-in view of the same specimen in This species has a contagious distribution apparently photo C, showing the mesenteries visible inside the pol- settling from one colony to neighboring colonies giv- yp wall. E) The same colony in photo A prior to collec- ing the appearance of what we call “infestations.” Most tion. F) A Z. cf. ammophilus colony (upper right) newly host colonies we observed were completely covered settled on a P. ramosa colony on Pratt Seamount at a with zoantharians (see photos E and G on next page) depth of 916 m. G) Two P. gymnogaster colonies on Marchand Seamount at a depth of 2316 m; a healthy 4Sinniger Harri, F. 2021. Personal commun. Tropical Biosphere Re- colony (at left) and a colony completely covered with search Center, Univ. Ryukyus, 3422 Sesoko, Motobu 905-0227, Oki- zoantharians (at right). The distance between the red la- nawa, Japan. ser marks in photos E and G is 10 cm. 84 Professional Paper NMFS 23 3. Zibrowius cf. ammophilus (continued) Chapter 4 85 Figure 4-5 A map of the North Pacific Ocean showing the distribution of Zibrowius cf. ammophilus () in Alaska waters. 86 Professional Paper NMFS 23 4. gen. nov., sp. nov. Preliminarily molecular DNA analysis based on the mi- eter of the scapulus disc in live polyps), slender, pointed tochondrial cytochrome oxidase c subunit 1 and 16S tentacles that are a slightly lighter color than the polyp. markers and the 8S and internal transcribed spacer Color in life and in ethanol is a light golden brown. Un- nuclear markers indicated that specimens collected on like other Alaska zoantharians, the oral disc of this spe- hexactinellid sponge reefs in Southeast Alaska on the cies is a darker color than the rest of the polyp. border with British Columbia are a novel genus and species in the family Parazoanthidae. Additional analy- Remarks An unknown species, apparently a Parazo- ses are underway and a new genus and species descrip- anthus, was collected on Pioneer Seamount off northern tion are in preparation (Sinniger Harri4). There are 16 California. The depth of that collection is unknown but known genera in the family Parazoanthidae (WoRMS it was likely quite deep since the specimens had com- Editorial Board, 2022) and the external appearance of pletely overgrown an antipatharian skeleton. the new specimens most closely resemble Parazoanthus but the molecular data clearly distinguish this genus Distribution Uncommon. In Alaska – known only from all other genera in the family Parazoanthidae. from specimens collected in Portland Canal, Southeast Alaska, on the border with British Columbia (Stone et Description Colonies consist of clusters of up to 32 al., 2013; Fig. 4-6). Elsewhere – Portland Canal, north- polyps and a very thin coenenchyme. Columns are most ern British Columbia (same reefs as reported here). often uniformly cylindrical and up to 26 mm in length and 14 mm in diameter in contracted specimens. Par- Habitat In Alaska – grows exclusively on the exposed ticles are not incrusted in the mesogloea of the column skeletal surfaces of the hexactinellid sponge reefs (87– or elsewhere. The oral disc is flat but broadly fluted. 107 m depth) formed principally by Aphrocallistes vas- Each polyp has 22–24 long (about two-thirds the diam- tus and Heterochone calyx along with oxide crusts. Figure 4-6 A map of the eastern Gulf of Alaska showing the distribution of gen. nov., sp. nov. () in Alaska waters. Chapter 4 87 Photos A) Preserved (in ethanol) portions of gen. of the North Reef in Portland Canal, Southeast Alaska, nov., sp. nov. colonies (USNM 1482149), photo- at a depth of 103 m. E) Gen. nov., sp. nov. colonies graphed from the top, collected in Portland Canal, growing on hexactinellid skeletons on the top of the Southeast Alaska, at a depth of 87 m. B) A ventral North Reef in Portland Canal, Southeast Alaska, at a view of the same specimens in photo A showing at- depth of 90 m. F) A close-up view of the same colonies tachment to hexactinellid skeletons. C) A close-up view in photo E. The distance between the red laser marks of the top specimen in photo A. D) Gen. nov., sp. nov. in photos D–F is 10 cm. colonies growing on hexactinellid skeletons on the top 88 CHAPTER 5 Subclass Octocorallia The Octocorallia are a diverse subclass of colonial an- guide for all octocorals, with a separate subsection for thozoans characterized by the subdivision of the polyp Pennatuloideans. by eight mesenteries, each division giving rise to a tenta- cle adorned with lateral pinnules, and tissues containing sclerites (Fig. 5-1; Watling et al., 2011). The two mono- Order Malacalcyonacea phyletic clades of the Phylum Cnidaria, Medusozoa and Anthozoa (Daly et al., 2007), were recently elevated The World Register of Marine Species currently recog- to the rank of sub-phylum by McFadden et al. (2022) nizes 46 families in the order Malacalcyonacea, only as part of their revised systematics of the Octocoral- four of which are represented in Alaska waters—Al- lia based largely on phylogenomics. Additionally, two cyoniidae, Clavulariidae, Gorgoniidae, and Paramuri- subclasses of Anthozoa, Hexacorallia and Octocorallia, ceidae. The order also includes corals in 46 genera and were elevated to the rank of class. Note that WoRMS two unaccepted families whose phylogenetic positions has not yet accepted the two rank suggestions made are still unknown or uncertain and are tentatively listed above by McFadden et al. (2022). Two new orders (Malacalcyonacea and Scleral- cyonacea) were established to accommo- axis date taxa from the six former suborders in coenenchymecentral cord the order Alcyonacea, five of which were solenia represented in Alaska waters—Alcyoniina, axis cortex Calcaxonia, Holaxonia, Scleraxonia, and gastrodermal canals Stolonifera. Corals from both orders are mesenteries common and abundant in most regions of Alaska, particularly in the Aleutian Islands gonads where they are relatively diverse compared gastric cavity to other high-latitude ecosystems world- peristome wide. The Malacalcyonacea include the major- pharynx ity of taxa in the former suborders Holaxo- nia and Alcyoniina and many taxa formerly classified as Stolonifera and Scleraxonia mesenterial filaments (McFadden et al., 2022). The Scleralcyona- loculi cea include all corals in the former order pinnules points crowntentacles Pennatulacea (now superfamily Pennatuloi- dea), most members of the former Calcaxo- nia, and a morphologically heterogeneous group of taxa from all five former subor- ders. Both orders contain taxa with a wide neck zone range of growth morphologies, including those with and without skeletal axes (Mc- anthocodia anthostele Fadden et al., 2022). axial sheath Definitions of the various growth forms and commonly used technical terms can be found in the “Illustrated trilingual glossary of morphological and anatomical terms Figure 5-1 applied to Octocorallia” by Bayer et al. A diagram showing the general anatomy of typical octocorals. This figure (1983). In Appendix I we also provide an is adapted from plate 2 in Bayer et al. (1983). abbreviated glossary of terms used in this Chapter 5 89 as incertae sedis (McFadden et al., 2022). The World Register of Marine Species lists these taxa as the fam- ily Malacalcyonacea incer- tae sedis. The vast majority of taxa in the Malacalcyo- nacea have either a largely proteinaceous or no skel- etal axis (McFadden et al., 2022). Family Alcyoniidae (Adapted from Fabricius and Alderslade, 2001 and McFadden et al., 2022) The family Alcyoniidae are soft corals in which most or all of the polyps are united in a common, fleshy mass without a supporting axis of horny and/or calcareous material. The coenenchyme Figure 5-2 is occasionally subdivided A map of the North Pacific Ocean showing the distribution of corals in the family Alcyoni- into an outer cortex and idae () in Alaska waters. inner medula. Colonies are membranous, lobate to digi- tate, arborescent with a single stalk, or bramble-like North Pacific Oceans, including most regions of Alaska, without a main stalk. Polyps are monomorphic and so the likelihood that the designation has been misap- usually fully retractile into the coenenchyme or promi- plied is high. Accordingly, Williams (2013) recommends nent calyces. Polyps are distributed over the colony but that molecular studies be done to help determine the are sometimes more densely packed on branch tips or taxonomic status of this group. A pilot molecular study terminal lobes. Sclerites are spindles, sticks, rods, clubs, was conducted in 2016 in Alaska to help determine the and radiates, and often brightly colored. taxonomic status of this group (Everett5).The large mu- In Alaska, the Alcyoniidae include corals in the gen- seum collections and ready access to large numbers of era Alcyonium and Gersemia (until recently designated fresh specimens in Alaska presents an ideal opportunity in the family Nephtheidae) and are found throughout to undertake a much-needed re-examination of the fam- much of the Gulf of Alaska, including a few seamounts ily in the North Pacific Ocean. in the Gulf of Alaska Seamount Province, the Aleutian We list here six taxa found in Alaska waters, only Islands where they are most common and abundant, the four of which have definitive taxonomy: Alcyonium eastern Bering Sea, and throughout the Arctic region pacificum Yamada, 1950; Gersemia fruticosa (Sars, (Fig. 5-2). As a group, they are extremely eurybathic 1860); G. Lambi Williams, 2013; and G. rubiformis. (1–2564 m depth), all attach to hard substrate, and they The group Alcyonium spp. includes specimens other may be found singly or in dense patches. than A. pacificum that may represent one or more spe- The family Alcyoniidae is in need of revision. All Alcy- cies, either not yet described or one of two other species, onium specimens collected in Alaska should be retained Scleronephthya spiculosa (Kükenthal, 1906) and A. ro- for taxonomic study. Williams (2013) recommends that bustum Utinomi, 1976, known from the North Pacific a taxonomic revision and determination of the validity region. of Gersemia rubiformis be undertaken and specifically notes that North Pacific Ocean specimens may actually be Alcyonium rather than Gersemia. Gersemia rubi- 5 formis reportedly has an extremely broad geographi- Everett, M. 2019. Personal commun. National Marine Fisheries Service, Northwest Fisheries Science Center, 2725 Montlake Bou- cal range over much of the North Atlantic, Arctic, and levard East, Seattle, WA 98112. 90 Professional Paper NMFS 23 1. Alcyonium pacificum Yamada, 1950 Description (Adapted from McFadden and Hoch- rudyi (Verseveldt and van Ofwegen, 1992), formerly berg, 2003) Colonies are mushroom-shaped with a Alcyonium rudyi Verseveldt and van Ofwegen, 1992. distinct capitulum and stalk. The capitulum is up to 4 Discophyton rudyi has been documented from south- cm in diameter and 1.5–2.0 cm in height, and slight- ern British Columbia to southern California and from ly depressed in the center. A short, irregularly shaped the low intertidal zone to <10 m depth (McFadden and stalk narrows basally to a cup-shaped attachment disc. Hochberg, 2003; Verseveldt and van Ofwegen, 1992). Polyps are large, up to 1 cm in length, and are evenly distributed over the upper surface and margins of the Distribution Uncommon. In Alaska – the Aleutian Is- capitulum only. The underside of the capitulum and lands from Adak Island to Attu Island (Fig. 5-3). Else- stalk lacks polyps. Tentacles have 15–19 pinnules along where – Northwest Pacific Ocean. The type locality is each edge. Polyps are fully retractile, leaving slightly re- Akkeshi Bay, Hokkaido, Japan and this species has also cessed apertures surrounded by eight triangular lobes been reported from shallow water near Matua Island (see photo F) formed by sclerites located in the most (Kuril Islands, Russia) in the Sea of Okhotsk (Sanamyan proximal region of the anthocodia. The distal region of and Sanamyan, 2020). the anthocodia lacks armature entirely. Sclerites are white and distributed throughout the Habitat In Alaska – typically found in small but dense capitulum surface and in the coenenchyme of the ca- patches on bedrock at depths between 6 and 12 m. Else- pitulum and stalk. Sclerites are radiates that are often ir- where – the type specimens were collected in the low regularly shaped and 0.09–0.11 mm in length. The color intertidal zone. of the colonies in life is bright orange; creamy white in ethanol. Photos A) A preserved (in ethanol) whole A. pacifi- cum colony (SBMNH 145135), with polyps fully re- Remarks This species closely resembles Discophyton tracted, collected near Attu Island, western Aleutian Figure 5-3 A map of the Aleutian Islands showing the distribution of Alcyonium pacificum () in Alaska waters. Chapter 5 91 Islands, at a depth of 6 to 12 m. B) A preserved (in ethanol) whole A. pacificum colony (SBMNH 348114) ethanol) whole A. pacificum colony (SBMNH 145135), collected near Adak Island, central Aleutian Islands, at with polyps partially retracted, collected at the same lo- a depth of 10 m. E) A close-up view of the same speci- cation and depth as the specimen in photo A. C) The men in photo B. F) A close-up view of the same speci- underside of the specimen in photo B showing the rela- men in photo D. Photos E and F are courtesy of Daniel tively short, irregularly shaped stalk. D) A preserved (in Geiger (Santa Barbara Museum of Natural History). 92 Professional Paper NMFS 23 2. Alcyonium sp. Description Colonies are mushroom-shaped with a rubiformis specimens and urges that a taxonomic revi- polypiferous capitulum that is covered densely and uni- sion be undertaken with special regard to the species. formly with autozooids. The stalk is robust and may Preserved specimens of Alcyonium sp. can easily be be relatively short or longer than the diameter of the mistaken for the pennatuloidean Cavernularia vansyoci capitulum. The stalk is up to 7 cm in height and the Williams, 2005 without careful examination to confirm capitulum is up to 7 cm in diameter. On rare occasion the absence of siphonozooids between the autozooids two colonies growing in close proximity to each other and to verify that the lower part of the colony is a hold- will fuse together along the stalk (see photo E). fast and not a peduncle. Color of the entire colony is a uniform bright to pale The nudibranch Tritonia festiva has been document- orange; lighter orange to yellow in ethanol. ed to prey on Discophyton rudyi along the west coast of North America (Verseveldt and van Ofwegen, 1992, Remarks The taxonomy of this group is unresolved so personal observ. by J. Goddard on p. 173). We have all specimens should be carefully re-examined. For the made similar observations of T. festiva preying on Al- purpose of this guide, we separate this group from Al- cyonium sp. in the central Aleutian Islands (see photo cyonium pacificum but the current status is taxon inqui- H). Shallow-water specimens are often covered partially renda. We are confident with the designation of Alcyo- with a brown diatomaceous film and heavily secrete mu- nium, and specimens grouped here are clearly neither cous upon collection. Heteropolypus, Pseudoanthomastus, nor Anthomastus. This group of specimens differs from all other Alcyoni- Distribution Uncommon. In Alaska – the Aleutian Is- idae known to occur in Alaska and undoubtedly repre- lands (Islands of Four Mountains to near Buldir Island) sents either one or more undescribed species or one(s) including Petrel Bank (Fig. 5-4). Elsewhere – unknown. not previously reported from the region. Alcyonium is a very large genus with 58 valid spe- Habitat In Alaska – in shallow water (5–24 m depth), cies currently recognized worldwide (WoRMS Editorial typically found in small but dense (>10 colonies/m2) Board, 2022). Most species are found in the Indo-Pacific patches on bedrock. In deeper water (103–600 m region and the Atlantic Ocean including the Mediterra- depth), typically single or in low-density patches on nean Sea. Only three species are known from the North bedrock, boulders, cobbles, and pebbles including frac- Pacific Ocean: 1) A. pacificum (discussed on page 90), tured basalt. 2) Scleronephthya spiculosa reported from the Sea of Okhotsk at 117 m, and 3) A. robustum reported from Photos A) A fresh whole Alcyonium sp. colony (CAS Japanese waters and inhabiting rocky areas at depths # pending) collected near Bobrof Island in the central 120–150 m. Aleutian Islands at a depth of 25 m. B) The capitulum Several taxa may be represented in this group. Speci- of a fresh Alcyonium sp. specimen (CAS # pending) mens partition into two distinct depth ranges (5–24 m collected near Semisopochnoi Island, western Aleutian and 103 m to about 600 m) and specimens with very Islands, at a depth of 22 m. C) The dried capitulums short and noticeably long stalks. No records exist from of two Alcyonium sp. specimens (USNM 1482135) col- depths between 24 and 103 m. We suggest that the lected near Bobrof Island at a depth of 18 m. D) A pre- short-stalked, shallow-water taxon (photos B, C, G, served (in ethanol) whole Alcyonium sp. colony (USNM and H) may be A. pacificum. The long-stalked shallow- 100742) collected near the Islands of Four Moun- water taxon (photo A) and long-stalked deeper-water tains, eastern Aleutian Islands, at a depth of 150 m. E) taxon (photos D, E, and F) likely represent additional Two conjoined whole Alcyonium sp. colonies (USNM taxa. 100802) preserved in ethanol, collected near Kiska Is- Two specimens (CAS 029138) from the Arctic Ocean land, western Aleutian Islands, at a depth of 103 m. F) (near Point Barrow) listed in Williams (2013) more An Alcyonium sp. colony photographed north of Little closely conform to Gersemia rubiformis as presented Tanaga Island, central Aleutian Islands, at a depth of in this guide so we list them there. However, Williams 25 m. G) An Alcyonium sp. colony photographed near (2013) argues that these specimens likely represent an Semisopochnoi Island at a depth of 26 m. H) A fresh unidentified species of Alcyonium, clearly not the one(s) whole Alcyonium sp. colony (USNM 1482136) collect- presented here. Williams (2013) furthermore questions ed on Petrel Bank at a depth of 160 m. The nudibranch the taxonomic validity of northeastern Pacific Ocean G. (T. festiva) was preying on the coral. Chapter 5 93 94 Professional Paper NMFS 23 2. Alcyonium sp. (continued) Figure 5-4 A map of the Aleutian Islands showing the distribution of Alcyonium sp. () in Alaska waters. Chapter 5 95 3. Gersemia fruticosa (Sars, 1860) Description (Adapted from Verrill, 1922) Mature though the geographical ranges for the two species are colonies consist of an obvious naked stalk branching ir- separated by more than 1800 km, we recommend that regularly near the base with numerous branches further the Alaska specimens be compared to specimens of G. dividing and subdividing into branchlets. Colonies are juliepackardae. up to 40 cm in height and 25 cm in width when fully The cyclical rhythm of colony expansion and con- expanded. The terminal branchlets are blunt or capi- traction has been studied in this species (Slephkova and tate but mostly clavate in contracted specimens. Each Seravin, 1983; Seravin and Gudkov, 1990), and it oc- branchlet may terminate in a few or many slender, elon- casionally harbors large numbers of ophiuroid commen- gated polyps, arising from slightly raised eight-lobed ca- sals (see photo C on next page). lyces. The polyps are so stiffened with small sclerites that they are nearly incapable of complete retraction within Distribution Uncommon. In Alaska – western Gulf the calyces. The tentacles are long, somewhat swollen of Alaska (Shumagin Bank) and the Aleutian Islands to proximally, but tapered to slender tips; their larger pin- Amchitka Pass (Fig. 5-5). Elsewhere – generally consid- nae are long and slender. The anthocodia terminates in ered a North Atlantic Ocean species where it was origi- eight small obtuse lobes or scallops corresponding to the nally described (Maritime Canada to Florida and Baffin tentacle bases. All sclerites are white in ethanol. Bay, Arctic Ocean) but also reported from the Kara Sea Colonies appear light orange or pinkish in situ and (Arctic Ocean), the Sea of Japan (Broch, 1932), and off often the stalk is a lighter shade or even whitish. Speci- the U.S. Pacific Coast. mens are darker orange or pink on deck, and orange to brown in ethanol. Habitat In Alaska – always found singly but occasion- ally has daughter colonies near their base (see photo D Remarks This taxon is represented by a single con- on next page). Found on hard rock (boulders, cobbles, firmed specimen (USNM 60475, identified by F. M. and pebbles) in areas of unconsolidated sediment at Bayer, 1982), 21 additional specimens collected in deep depths between 1005 and 2564 m. Elsewhere – eury- water (1199–2248 m) and identified as Gersemia cf. bathic but principally a deep-water species: U.S. Pacific fruticosa, and 13 colonies observed in situ in the central Coast (to 1946 m), Sea of Japan (70–100 m), Maritime Aleutian Islands (Stone, 2014). Canada (91–361 m), mid-Atlantic U.S. (to 3506 m), Verrill (1922) reports that this species is similar to G. Florida (to 770 m), and shallowest in the Kara Sea (Arc- rubiformis in modes of branching and general morphol- tic Ocean) at 32–43 m. ogy but has less coenenchyme between the calyces and more sclerites in the anthocodial region and proximal Photos A) A preserved (in ethanol) whole G. fruticosa part of the polyps. Additionally, the two species have colony (USNM 60475) collected on Shumagin Bank, vastly different depth distributions and our observations western Gulf of Alaska, at a depth of 1143 m. B) A close- indicate that G. fruticosa has a more stalked rather than up view of the same specimen in photo A. C) A preserved clustered morphometry. (in ethanol) whole G. fruticosa colony (CAS 234586) Some of the specimens included here in this group collected in northern Amchitka Pass at a depth of 2248 (e.g., the one pictured in photos E and F on next page) m. The base of the colony is on the left. D) The same are quite similar in appearance to G. juliepackardae colony in photo C, only partly expanded, prior to col- Williams and Lundsten, 2009, known from the conti- lection. A daughter colony is present immediately to the nental slope off northern Washington, central Oregon, right of the collected specimen (black circle). E) A G. fru- and northern California including Monterey Canyon ticosa colony (CAS 234577) in situ northwest of Kanaga and several seamounts. This species attaches to hard Island, central Aleutian Islands, at a depth of 1199 m. substrate including hexactinellid skeleton at depths be- The colony is fully expanded. F) A close-up view of the tween 520 and 2034 m. The Alaska specimens listed same colony in photo E. The distance between the red here occupy similar habitats including depth range. Al- laser marks in photos D–F is 10 cm. 96 Professional Paper NMFS 23 3. Gersemia fruticosa (Sars, 1860) (continued) Chapter 5 97 Figure 5-5 A map of the North Pacific Ocean showing the distribution of Gersemia fruticosa () in Alaska waters. 98 Professional Paper NMFS 23 4. Gersemia lambi Williams, 2013 Description (Adapted from Williams, 2013) Colo- Distribution Uncommon but reportedly abundant lo- nies are composed of dense concentrations of auto- cally (Fig. 5-6). In Alaska – known only from obser- zooids distributed in isolated clusters on several short vations made in situ between Cape Ommaney at the lobes that emanate from a very short stalk and asym- southern tip of Baranof Island and Cross Sound and Icy metrical holdfast. Each cluster usually contains 15–30 Strait north of Chichagof Island, eastern Gulf of Alaska polyps. Polyps are erect and curve upward from their (McDaniel6). Elsewhere – known from three locations bases. Polyps are monomorphic, non-retractile, tubular, in central to northern British Columbia, including near and relatively large (4–7 mm in height and 1.5–2 mm in Langara Island in southern Dixon Entrance, near the width). The width of the polyps is greatest at the distal border with Southeast Alaska. extremity. Most coenenchymal sclerites of the polyps are sharp- Habitat In Alaska – occurs in dense patches from the ly tuberculated radiates (0.03–0.12 mm in length), and shallow subtidal zone to 30 m on bedrock and in areas most of those in the stalk are variably ornamented radi- of strong current and wave surge. Elsewhere – shallow ates and modified radiates (0.03–0.12 mm in length). subtidal zone (9–20 m depth) on bedrock and in areas Polyp wall sclerites are abundant, uniformly and densely of strong current and wave surge. distributed, and most are variably shaped radiates and Photos A) A lateral view of a preserved (in ethanol) rods with a few irregularly shaped elongate forms and whole Gersemia lambi colony (CAS 171940; paratype) crosses (0.03–0.13 mm in length). Tentacular sclerites collected near Langara Island, Haida Gwaii, in Dixon are densely and uniformly distributed and mostly radi- Entrance, northern British Columbia, at a depth of 12 ates and rods, although a few club-shaped or torch-like forms are also present. Colonies are pinkish-orange in life and creamy white 6McDaniel, N. 2019. Personal commun. McDaniel Marine Sur- to pale orange when preserved in ethanol. veys, 354 West 35th Ave., Vancouver, BC, Canada V6N 2N5. Figure 5-6 A map of the eastern Gulf of Alaska showing the distribution of Gersemia lambi () in Alaska waters. Chapter 5 99 m. B) A view from the top of the preserved (in etha- Islands, Haida Gwaii, central British Columbia, at a nol) G. lambi holotype (CAS 171939; whole colony) depth of 6 m. E) Gersemia lambi colonies with polyps collected near Langara Island, Haida Gwaii, in Dixon fully contracted and surrounded by a bed of what ap- Entrance, northern British Columbia, at a depth of pears to be G. rubiformis with much smaller polyps 12 m. C) Gersemia lambi colonies with polyps fully fully extended. The colonies were photographed near extended, photographed near the type collection sites. the Kerouard Islands, Haida Gwaii, central British Co- The photo is courtesy of Marc Chamberlain. D) Ger- lumbia, at a depth of 6 m. Photos D and E are cour- semia lambi colonies with polyps fully contracted. tesy of Neil McDaniel. All photos are reproductions of The colonies were photographed near the Kerouard those provided in Williams (2013). 100 Professional Paper NMFS 23 5. Gersemia rubiformis (Ehrenberg, 1834) Description (Adapted from Verrill, 1922) Dried (con- elongated, rough, fusiform sclerites arranged en chev- tracted) specimens consist of rounded or ovate clusters ron. The extended mature polyps are much smaller than of rather hard, short, thick branches or lobes, convex those of G. lambi, about 2.0–2.5 mm in length and 0.7 externally, and attached to the main stalk by short mm in width; tentacles are usually without sclerites. stems, smaller than the enlarged ends. The main stem Color of colonies in life and in ethanol is uniform may be very short or somewhat elongated and without shades of orange and red. branches or calyces near the base, which usually spreads out in a thin expansion for attachment. The lower part Remarks This species is gonochoric and broods of the stem and holdfast often lack the surface layer of its larvae (Verrill, 1922; Nørrevang, 1973). The effects red sclerites that occurs elsewhere in the colony. The of simulated fishery disturbance on the reproduction surface of the branches bears polyps and is covered with and regenerative capability of this species have been a thin but firm layer of compact minute rough sclerites studied in the laboratory (Henry et al., 2003). Gerse- (red to pale red in color) which give the surface a finely mia rubiformis includes 51 specimens definitively iden- granular appearance and the corallum a red, pinkish, or tified and 15 additional specimens identified as G. cf. light to bright orange color. Sclerites include spindles rubiformis. (irregular, short, lobed, or warty), double spindles, el- lipsoids, and a few elongated rough forms. Contracted Distribution Locally common and widespread. This specimens resemble raspberries or strawberries and bear species is the northernmost coral in the Pacific Ocean. the common name “sea raspberries” or “sea strawber- In Alaska – generally a northern taxon, particularly ries.” They typically range to about 6 cm in diameter. common and abundant on the broad continental shelf The polyps in preserved specimens are usually com- of the eastern Bering Sea, Chukchi Sea, and Beaufort pletely contracted, but not always revealing a conical Sea. Also present in the Gulf of Alaska and the Aleu- anthocodia with eight feeble double rows of minute and tian Islands from Unimak Pass to southern Amchitka Pass (Fig. 5-7). Elsewhere – nearly circumpolar with a geographical range simi- lar to G. fruticosa: off the Kamchatka Peninsula (Rus- sia) including the Sea of Okhotsk, the Pacific Coast of North America (Brit- ish Columbia to northern California), Arctic Ocean (Kara Sea to Baffin Bay and Greenland), and the Atlan- tic Coast of North America (Maritime Canada to North Carolina). Habitat In Alaska – oc- curs in dense patches on bedrock in shallow water but singly or in small clus- ters on cobbles and pebbles in areas of unconsolidated sediment in the deeper part of its range. Total depth range is 3–90 m. Elsewhere – reported on cobbles, peb- Figure 5-7 bles, and shell at depths be- A map of the North Pacific Ocean showing the distribution of Gersemia rubiformis () in tween 1 and 457 m. Alaska waters. Chapter 5 101 Photos A) A preserved (in ethanol) whole G. rubifor- depth of approximately 40 m. D) A G. rubiformis col- mis colony (USNM 1482143) collected near Adak Is- ony photographed near Cape Ulitka, Noyes Island, in land, central Aleutian Islands, at a depth of 18 m. B) the eastern Gulf of Alaska at a depth of about 15 m. E) A preserved (in ethanol) whole G. rubiformis colony A G. rubiformis colony photographed at the same loca- (USNM 1482144) collected near St. Paul Island in the tion and depth as the specimen in photo D. Photos D eastern Bering Sea at a depth of 8 m. C) Gersemia ru- and E are courtesy of D. K. Harris (Alaska Department biformis specimens collected in the Chukchi Sea at a of Fish and Game). 102 Professional Paper NMFS 23 6. Gersemia sp. Description Colonies are dentritic to slightly bushy Distribution Rare. In Alaska – Pratt and Giacomini with branches emanating above the base and on all Seamounts in the Gulf of Alaska Seamount Province sides. Branches may terminate in up to five large, (Fig. 5-8). Elsewhere – unknown. brightly colored polyps (approximately 60 polyps to- tal). The largest contracted specimen measures 18 mm Habitat In Alaska – occurs singly or in clusters up to in height and 16 mm in width, but the same speci- seven colonies on bedrock and fractured bedrock and at men measured in situ fully expanded is up to 10 cm in depths between 897 and 1095 m. height and 7 cm in width. The color of the main colony and base in situ is creamy white and the distal end of Photos A) A preserved (in ethanol) whole Gersemia branches and polyps is orange-red to pink. In ethanol, sp. colony (USNM 1075807) collected on Pratt Sea- most of the colony is reddish-orange with a creamy bas- mount at a depth of 1092 m. B) The opposite side view al plate. of the same specimen in photo A. C) Mostly contracted Gersemia sp. colonies in situ on Pratt Seamount at 1092 Remarks The species is known from only two collect- m depth. Specimens in photos A and B are visible in the ed specimens and several video observations of similar center of the photo. D) A Gersemia sp. colony (fully specimens made on the same and a nearby seamount. expanded) observed on Pratt Seamount at a depth of At the collection site, seven specimens were clustered 1095 m. The distance between the red laser marks in together, each separated by 2–10 cm; the two largest photos C and D is 10 cm. E) A close-up view of the specimens were collected. same colony in photo D. F) A zoomed-in view of the same colony in photo D. Figure 5-8 A map of the North Pacific Ocean showing the distribution of Gersemia sp. () in Alaska waters. Chapter 5 103 104 CHAPTER 6 Family Clavulariidae (Adapted from Verrill, 1922 and McFadden et al., are rare or uncommon and have a limited distribution in 2022) Clavulariids are stoloniferous octocorals with- the western Gulf of Alaska, Pratt Seamount in the Gulf out a skeletal axis and with polyps connected basally of Alaska Seamount Province, the Aleutian Islands, and by ribbon-like stolons. Polyps are monomorphic; an- the eastern Bering Sea as far north as Zhemchug Canyon thocodiae are retractile into clavate calyces that may (Fig. 6-1). They are eurybathic fauna occupying depths be extraordinarily tall. Pinnules may be free or fused, between 82 and 3277 m. Aside from growing directly forming broad blade-like expansions of the tentacle. on hard rock (bedrock, boulders, cobbles, pebbles, and The prominent calyces are packed with sclerites (most- siltstone), they often encrust a wide array of biotic sub- ly warty spindles), and the polyps and anthocodiae are strates including the skeletons of octocorals (keratoisi- quite spiculose (Verrill, 1922). Colonies are generally dids, gorgoniids, and paramuriceidids), antipatharians, small and consist of cylindrical or bluntly conical pol- stylasterids, hexactinellid sponge skeletons, mollusk yps, usually joined only at their bases by reticulating shells, and bryozoans. stolons which may further coalesce into thin membra- All clavulariid specimens collected in stock assess- nous expansions. ment and research surveys should be retained for tax- Of the three known genera of Clavulariidae, only onomic studies, including advanced molecular DNA Clavularia (four species) is found in Alaska. Most taxa techniques. Figure 6-1 A map of the North Pacific Ocean showing the distribution of corals in the family Clavulariidae () in Alaska waters. Chapter 6 105 1. Clavularia armata Thomson, 1927 Description (Adapted from Thomson, 1927) The and the other had completely overgrown a Bathygorgia most peculiar characteristic of this species is the ex- profunda colony that was attached to fractured basalt. treme spiculation of the entire colony, including the sto- Also, two additional colonies were observed together on lon and calyces. This is due to the projection of the tips unknown hexactinellid skeletons growing on another of the spindles, which, in general, are directed upwards boulder nearby. The largest collected specimen has a and outwards from the calyx. Abundant sclerites at the relatively thick coenenchyme densely populated with upper part of the calyx are arranged in eight groups, large polyps (18 mm in length) that taper from the base each of which exhibits roughly an en chevron arrange- (approximately 6 mm) to the top (4 mm). The tentacles ment, but progresses to distinct protrusions. Sclerites are quite long (up to 12 mm in length) and during col- are almost entirely spindles (0.35–0.40 mm in length by lection of the specimen the polyps were very slow to 0.04–0.06 mm in diameter) and a few distinct crosses retract. The color of the coenenchyme is a uniform, light (0.23 mm in length). The largest spindles (0.43–0.46 orange in life; the polyps are a lighter orange, almost mm in length by 0.07–0.09 mm in diameter) bear a few translucent along their length. warts. The type specimen has polyps that are about 4 mm in length and only partially retracted into the ca- Distribution Rare. In Alaska – known only from lyx. This specimen was growing on the bare skeletal Pratt Seamount in the Gulf of Alaska Seamount Prov- section of a live bamboo coral (Keratoisis sp.) and was ince (Fig. 6-2). Elsewhere – the type specimens were col- described as slightly bluish in color, presumably due to lected near the Azores (Portugal) in the North Atlantic the presence of dark mineral particles inside the polyps. Ocean. This species is known in Alaska from only two col- lected specimens; one had mostly overgrown the skeletal Habitat In Alaska – grows on hexactinellid and bam- stalk of a hexactinellid sponge (Crateromorpha sp.) that boo coral skeletons that are attached to boulders at was attached to a boulder (photos A–E on next page) depths between 2667 and 2730 m. Elsewhere – grows Figure 6-2 A map of the North Pacific Ocean showing the distribution of Clavularia armata (), C. eburnea (♦), and C. rigida (▲) in Alaska waters. 106 Professional Paper NMFS 23 on the skeleton of bamboo corals (Keratoisis sp.) in ar- completely overgrown the attached stalk (relic) of a eas of sand and limestone at depths between 1153 and hexactinellid sponge (Crateromorpha sp.). B) A close- 1600 m. up view of the same colony in photo A. C) The same colony in photo A in situ. D) The same colony in photo Photos A) A preserved (in ethanol) fragmented but A during collection. The holdfast of the sponge is indi- whole Clavularia armata colony (USNM 1076621) col- cated by the white circle. E) A close-up view of the same lected on Pratt Seamount in the Gulf of Alaska Sea- colony with fully extended polyps. The distance be- mount Province at a depth of 2689 m. The colony has tween the red laser marks in photos C and D is 10 cm. Chapter 6 107 2. Clavularia eburnea Kükenthal, 1906 Description (Adapted from the original description tian Islands and identified by Nutting (1912). Nutting by Kükenthal, 1906) Numerous polyps originate from (1912) noted that the specimen was attached to an al- a membranous base that thickens in places at acute an- cyonarian stem and had small polyps (7 mm in length), gles. Polyps are large (up to 12 mm in length and 3 mm but otherwise conformed to the description for the type in width), at least partly retractile, and each has a calyx specimen. The specimen has clearly and completely half the height of the total polyp. The calyx has a sharp overgrown an octocoral skeleton, possibly Calcigorgia wall with eight distinct longitudinal furrows; the upper, japonica Dautova, 2007, and without very careful ex- retractile section of the polyp is more delicately walled, amination would likely have been mistaken as the latter cup-shaped, and supports stout tentacles up to 3 mm species. The specimen appears to have a very thick coe- in length with 10–11 thick pinnules on each side, the nenchyme and the color in ethanol is very pale orange. middle of which are the longest. The pinnules contain rod-shaped, wide-thorned sclerites that are 0.09 mm in Distribution Rare. In Alaska – known only from a length; they are slightly larger in the tentacle axis and single specimen collected southeast of Agattu Island in arranged in double rows that converge downwards. The the western Aleutian Islands (Fig. 6-2). Elsewhere – the upper polyp section contains longitudinal spindles up type specimen was collected in the Sea of Japan. to 0.25 mm in length, as well as clubs and more irregu- lar bodies, all irregularly adorned with large, rounded Habitat In Alaska – grows on attached octocoral skel- thorns. The calyx wall contains many spiked lobes eton at depths near 715 m. Elsewhere – the type speci- 0.13 mm in length that are strongly constricted in the men was collected at a depth between 600 and 1200 m. middle. The basal section of the polyp and the coenen- chyme contain smaller (0.07 mm in length), irregular- Photos A) A preserved (in ethanol) whole Clavularia shaped sclerites. The color of the specimen was a uni- eburnea colony (USNM 30106) collected southeast of form ivory white. Agattu Island, western Aleutian Islands at a depth of The only record of this species in Alaska is a speci- 715 m. The specimen has completely overgrown an oc- men (USNM 30106; photos A and B below) collected by tocoral (likely Calcigorgia sp.). B) A close-up view of the U.S. Fisheries Steamer Albatross in the western Aleu- the same colony in photo A. 108 Professional Paper NMFS 23 3. Clavularia rigida Broch, 1935 Description (Adapted from the original description ily armored with small thorns. The tentacles are quite by Broch, 1935) Colonies form a membranous crust- small (0.15–0.34 mm in length). like stolon that gives rise to relatively large polyps (up The only record of this species in Alaska is a speci- to about 8 mm in length) that are completely retractile men (USNM 1081183; photos A and B below) collect- into cylindrical or weakly club-shaped calyces. Caly- ed on the deep abyssal plain near the Aleutian Trench ces reach 2.3 mm in diameter near the slightly wider and identified by S. D. Cairns in 2005. The specimen top and are quite rigid due to a dense concentration has completely overgrown the stem of an antipathar- of sclerites. The polyps are packed close together. The ian skeleton, probably Bathypathes tiburonae, which calyx wall has eight weakly protruding longitudinal are fairly common in that area and at that depth. The bulges, which are reinforced in younger polyps with specimen appears to have a very thin coenenchyme with eight double rows of densely lying, upwardly converg- irregularly situated polyps, and the color in ethanol is a ing sclerites. In older individuals, the double rows are very pale orange. better described as eight dense lines of sclerites that are arranged almost completely parallel. The closed calyx Remarks Close examination of the preserved speci- resembles an eight-pointed star-shaped feature above men revealed that there was actually a second very the retracted head. small (only three polyps) colony growing on the hold- A crown of well-developed sclerites adorns the pol- fast. yp head, which is sharply bordered and separated from the calyx by a wide area without sclerites. A basal wreath of horizontally lying sclerites, from which rise Distribution Rare. In Alaska – known only from a eight short double rows, forms the crown. The ten- single specimen collected on the deep island arc slope tacle stem contains only densely packed sclerites. The south of Davidson Bank, eastern Aleutian Islands (Fig. stolon plate contains densely packed sclerites, which 6-2). Elsewhere – the type specimen was collected from are irregularly curved, quite slender (0.5–0.7 mm in the Sea of Okhotsk, Russia. length), spindle- or rod-shaped, and heavily thorned. The densely packed sclerites of the calyx are relatively Habitat In Alaska – the specimen was growing on the large (1.40–1.56 mm in length); the smaller and me- unpinnulated lower (7 cm) stalk of a black coral, likely dium-sized sclerites are slender and heavily thorned, Bathypathes tiburonae, that occurs in the same area and sometimes slightly curved spindles. The larger ones are depth range. The black coral was growing on siltstone basic spindles that are also heavily thorned, but other- at a depth of 3277 m. Elsewhere – the type specimen wise irregular variably sized wart-like or thorny spurs, was collected at a depth of 1076 m. often with forked ends. The sclerites of the crown of the polyp head are 0.34–0.55 mm in length and are Photos A) A preserved (in ethanol) whole Clavularia otherwise similar to those of the stolon plate, except rigida colony (USNM 1081183) collected south of Uni- more weakly curved. The tentacle sclerites are short, mak Island, eastern Aleutian Islands, at a depth of 3277 broad, oval- to rod-shaped with rounded ends; heav- m. B) A close-up view of the same specimen in photo A. Chapter 6 109 4. Clavularia sp. A Description Colonies form irregularly shaped mats up sue of other marine life but there is some evidence (see to 15 cm in length and width. Polyps are randomly dis- photos A and B on next page) that they may also be tributed throughout the stolon in distinct clusters and epizoic, similar to zoantharians that settle and grow on are up to 8 mm in height when fully expanded. Calyces live animals and kill them in the process. are almost perfectly cylindrical, short, and stout (up to The coenenchyme of this species is composed almost 4 mm in height and 4 mm in diameter). Polyps appear entirely of high-magnesium calcite (8.3 mol% MgCO3; to be at least partly non-retractile, and the collarets in senior author and S. D. Cairns, unpubl. data). Associ- fresh specimens are about the same length and diameter ated microfauna include pleustid amphipods (Watling as the calyx. The tentacles and pinnules are short and and Stone7). stout. The sclerites have not yet been described for this taxon. The coenenchyme is relatively thick and the col- Distribution Locally common. In Alaska – western or in life is a uniform orange with slightly darker pol- Gulf of Alaska (south of Sanak Island), Aleutian Islands yps; the color fades somewhat in dried specimens and (near Unalaska Island to southeast of Agattu Island), turns a golden brown in ethanol. and Pribilof Canyon to north of Zhemchug Canyon in the eastern Bering Sea (Fig. 6-3). This species is par- Remarks Our observations of this species growing in ticularly abundant on Amchixtam Chaxsxii in northern heavily disturbed habitats in the central Aleutian Islands (see photo E on next page) indicate that it might be a 7 pioneer species with a relatively fast growth rate. It is Watling, L., and R. P. Stone. 2007. Appendix E. Macro-fauna as-sociated with deep corals in the Aleutian Islands. In Deep sea also an opportunistic species, as are other Alaska sto- coral distribution and habitat in the Aleutian Archipelago (J. loniferans, in that they settle and grow on both abiotic Heifetz, D. Woodby, J. Reynolds, and R. P. Stone, eds.), p. 295– and biotic substrates. Most of our observations indicate 304. North Pacific Res. Board Final Rep. 304. [Available from https://projects.nprb.org/#metadata/ebf15fb7-dc07-406a-bb74- that these corals grow on the skeletons or non-living tis- 8dd3c978a1e0/project] Figure 6-3 A map of the North Pacific Ocean showing the distribution of Clavularia sp. A () in Alaska waters. 110 Professional Paper NMFS 23 Amchitka Pass (central Aleutian Islands). Elsewhere – Aleutian Islands at a depth of 744 m. B) A fresh whole unknown. Clavularia sp. A colony (USNM 1116851) growing on the bare skeleton of a Callistephanus pacificus colony Habitat In Alaska – colonies grow on hard rock (bed- collected in Zhemchug Canyon in the eastern Bering Sea rock, boulders, cobbles, pebbles, and siltstone) and often at a depth of 494 m. C) A close-up view of the same encrust a suite of biotic substrates including the skeletons colony in photo B. D) A close-up view of the section of octocorals (gorgoniids and keratoisidids), stylasterids, of the same colony in photo B growing directly on the hexactinellid skeletons, bivalve shell (scallop), and bryo- cobble to which the host coral was attached. E) A Cla- zoans at depths of 82 to 990 m. The corals appear to vularia sp. A colony (indicated by the white circle) ob- grow partly on living tissue, at least in the case of the served on a slope heavily disturbed by fishing activities stylasterids (see photo A). on Amchixtam Chaxsxii at a depth of 165 m. The dis- tance between the red laser marks is 10 cm. F) Several Photos A) A fresh whole Clavularia sp. A colony Clavularia sp. A colonies observed northwest of Tanaga (CAS 235595) growing on a Distichopora borealis Island in the central Aleutian Islands at a depth of 160 colony collected on Amchixtam Chaxsxii in the central m. 111 CHAPTER 7 Family Gorgoniidae (Adapted from McFadden et al., 2022) Gorgoniids are diates and small spindles; all are often brightly colored. octocorals with a proteinaceous skeletal axis that may Of the 13 known genera of Gorgoniidae, only Cal- contain non-scleritic carbonate hydroxylapatite. The listephanus (two species) is found in Alaska waters. axis is hollow with a narrow cross-chambered core Both species are common and have a broad distribution and a dense cortex with little loculation. Colonies are along the Gulf of Alaska continental shelf, seamounts erect, variably branched, planar or bushy, and often an- in the Gulf of Alaska Seamount Province, the Aleutian chored by a strong holdfast that may be calcified. Polyps Islands, and the eastern Bering Sea as far north as Per- are monomorphic, retractile into coenenchyme or low venets Canyon (Fig. 7-1). They are deep-water and eu- mounds, and distributed over all surfaces or arranged bi- rybathic fauna occupying depths between 96 and 2778 serially on the branches. Polyp sclerites are flattened rods m and grow directly on hard rock (bedrock, fractured with variable margins and coenenchyme sclerites are ra- bedrock, boulders, cobbles, pebbles, and siltstone). Figure 7-1 A map of the North Pacific Ocean showing the distribution of corals in the family Gorgoniidae () in Alaska waters. 112 Professional Paper NMFS 23 1. Callistephanus pacificus Nutting, 1912 Description (Adapted from Nutting, 1912 and Hor- is crimson red in color with pink or dark orange high- vath, 2019) Colonies are flabellate, almost always in lights. The hydroid was observed in southern Amchitka one plane, and moderately to densely branched. The Pass at depths between 274 and 352 m where the sea- largest known specimen (USNM 1481935) is 28 cm in floor was heavily disturbed by bottom-contact fishing height and 40 cm in width but larger specimens have gear, littered with sponge and coral debris, and mostly been observed in situ. In general, branching is highly ir- inhabited by fast-growing species of hydroids. Cal- regular but may be opposite or alternate and to the 4th listephanus pacificus were not observed at this site but order and 5th order in the largest colonies. Secondary they do occur elsewhere in Amchitka Pass, usually in branches may be quite short, giving the appearance of slightly deeper water. anastomosis; most branches curve upwards. Secondary This species hosts an extremely rich suite of associ- branching starts very close to a relatively stout and cup- ated fauna, including amphipods (caprellids, pleustids, shaped holdfast; sometimes two colonies apparently stenothoidids, and the ischyrocerid Bonnierella sp.), share the same holdfast. The axis is woody, strong, and isopods (antarcturids and aegids), and pycnogonids difficult to tear; it is dark brown but lighter near the (Watling and Stone7). Hydroids and small actiniarians branch tips. Colonies are crimson red to brilliant orange often grow on bare sections of the axial skeleton, and in life and the polyps are often highlighted with orange; several species of ophiuroids (including Ophiopholis the coenenchyme is a paler cranberry red when dried or japonica and Ophiura leptoctenia) are common associ- preserved in ethanol. ates. Colonies on seamounts in the Gulf of Alaska Sea- The calyces are prominent, conical and rounded, mount Province occasionally host the large sternostylid mostly lateral on the branches, alternate and opposite crab Sternostylus iaspus and the large ophiuroid Astero- in position, and about 3–4 mm apart. Typical calyces nyx. The coenenchyme and sclerites are principally com- measure 1.2 mm in height and 2.2 mm in diameter. posed of high-magnesium calcite (8.6 mol% MgCO3) The anthocodiae are well developed, fully retractile but with minute amounts of aragonite (senior author and S. slightly exsert in preserved specimens, with only the D. Cairns, unpubl. data). crown resting on the calyx margin. Colonies are able to retract their polyps very quickly when disturbed during Distribution Widespread (Fig. 7-2) and locally com- collection. mon. In Alaska – eastern and western Gulf of Alaska Polyp sclerites are bright red or orange rods and pale (west of Prince of Wales Island to Prince William pinkish-red warty spindles and capstans. The tentacles Sound), the Gulf of Alaska Seamount Province (Densen, are armed with numerous spindles arranged en chevron Welker, Pratt, Surveyor, Marchand, and Chirikof Sea- basally and in longitudinal rows distally. The coenen- mounts), the Aleutian Islands (south of Unimak Island chyme sclerites are unilaterally developed superficial to near Agattu Island), and the eastern Bering Sea slope capstans and spindles (0.08–0.17 mm in length) with (Pribilof Canyon to north of Pervenets Canyon). No- flatter warty spindles in the layer beneath. tably, the holotype (USNM 30024) collected east of Semichi Island by the U.S. Fisheries Steamer Albatross Remarks This species was originally described by in 1906 is one of only two records from the western Nutting (1912) as Callistephanus pacificus but until just Aleutian Islands. Elsewhere – northern British Colum- recently was known as Swiftia pacifica (Nutting, 1912). bia (west of the Queen Charlotte Islands) to central Or- This species has a very broad geographic and bathymet- egon (off Cape Meares). ric distribution and morphological characteristics that apparently overlap with congeners (Horvath, 2019). Habitat In Alaska – attaches to hard rock including Accordingly, Breedy et al. (2015) suggested that a thor- bedrock, fractured bedrock, siltstone, boulders, cobbles, ough review of Callistephanus be undertaken to clarify and large pebbles on sloped habitats and at depths be- taxonomic issues, including the presence of sibling or tween 96 and 2778 m. There are, however, only three cryptic species. records shallower than 485 m (96 m, 210 m, and 278 We have observed a hydroid that is superficially simi- m). Found in small patches with maximum densities of lar to Callistephanus including C. pacificus in the cen- 8 colonies/m2, often in association with C. simplex, and tral Aleutian Islands. This hydroid is bushy with what additionally Paragorgia sp. A and Isidella tentaculum we call a “creeping” appearance, not upright, delicately on seamount habitats. Elsewhere – found at depths be- branched and wiry, with up to eight scraggly branches tween 251 m (off Washington) to 2904 m (British Co- (8–10 cm in length) that originate near the base, and lumbia). Chapter 7 113 Photos A) A fresh whole C. pacificus colony (USNM attached caprellid amphipod collected south of Tanaga 1481937) collected west of Chichagof Island, eastern Island, central Aleutian Islands, at a depth of 824 m. GOA, at a depth of 816 m. B) A fresh whole C. paci- E) Callistephanus pacificus colonies photographed on ficus colony (USNM 1481941) collected in Zhemchug Welker Seamount at a depth of 1120 m. F) A C. pacifi- Canyon, eastern Bering Sea, at a depth of 520 m. C) A cus colony (USNM 1075771) collected on Densen Sea- close-up view of the same specimen in photo B. D) A mount at a depth of 2402 m. The distance between the fresh C. pacificus specimen (USNM 1481933) with an red laser marks in photos E and F is 10 cm. 114 Professional Paper NMFS 23 1. Callistephanus pacificus Nutting, 1912 (continued) Figure 7-2 A map of Alaska showing the distribution of Callistephanus pacificus () in Alaska waters. Chapter 7 115 2. Callistephanus simplex (Nutting, 1909) Description (Adapted from Nutting, 1909) Colonies The reproductive biology (Feehan and Waller, 2015) are sometimes scraggly, whip-like, quite flexible, and and connectivity of populations along the U.S. Pacific unbranched or only slightly branched; colonies have Coast have been studied using restriction-site associated up to 12 branch tips but with 5th order branching (see DNA sequencing (Everett et al., 2016). photo A on next page). The main axis and branches are round, slender, and of relatively uniform thickness Distribution Locally common. In Alaska – the eastern throughout (3.6–5.2 mm). Colonies are up to at least Gulf of Alaska continental slope and Dickens, Densen, 50 cm in height. The holdfast is generally discoid, stout, Welker, Surveyor, Pratt, Quinn, Giacomini, and Pat- and robust, and often covered with coenenchyme. The ten Seamounts in the Gulf of Alaska Seamount Prov- coenenchyme is relatively thin for the family. ince (Fig. 7-3). A single specimen was collected west of Calyces are uniformly distributed, not crowded, of- Tanaga Island in the central Aleutian Islands. Elsewhere ten spaced apart by as much as 2 mm, tubular, small – Washington to the Channel Islands, California. The (approximately 1 mm in height), and usually higher type specimens (USNM 25431) were collected off Santa than broad. The polyp walls contain red spindle-shaped Cruz Island (817–819 m). sclerites, as do the areas on and near the tentacle bases, where they are arranged more or less en chevron. Other- Habitat In Alaska – found in small patches (up to 5 wise the sclerites are longitudinally arranged. colonies/m2), occasionally singly, on bedrock, siltstone, The sclerites are mainly of two types: 1) small double boulders, cobbles, and pebbles at depths between 500 spindles, rosettes, stars, and small clubs located most- and 1352 m. Often co-occurs with C. pacificus. Else- ly in the superficial layer of the coenenchyme, and 2) where – found at depths between 549 and 1463 m. larger spindles that are slender, pointed, often slightly curved, covered with regularly distributed verrucae, and Photos A) A fresh whole (except the holdfast) C. much more abundant than the small double spindles. simplex colony (GOA06-100B-01) collected west of The clubs are much less numerous than the other sclerite Chichagof Island, eastern Gulf of Alaska, at a depth forms. of 816 m. B) A dried whole C. simplex colony (USNM Color of the coenenchyme in life is crimson red to 1481924) collected southeast of the Fairweather bright orange, and the polyps are a lighter shade than Ground, eastern Gulf of Alaska, at a depth of 769 m. the coenenchyme. Color in dried specimens is similar, C) A close-up view of the same specimen in photo B. but polyps are a darker shade than the coenenchyme. D) A close-up view of a dried C. simplex specimen (USNM 1481923) collected southeast of the Fairweath- Remarks Until recently this species was known as er Ground at a depth of 651 m. E) A C. simplex col- Swiftia simplex (Nutting, 1909). Seamount colonies of- ony (USNM 1075782) collected on Welker Seamount ten host ophiuroids including the large Asteronyx sp. at a depth of 793 m. F) A C. simplex colony (USNM and the large sternostylid crab (Sternostylus iaspus) that 1075781) collected at the same location as the speci- uses the colony as an elevated feeding platform, and men in photo E with a C. pacificus colony (USNM damaged or dying specimens are often covered with 1075781). G) A C. simplex colony (USNM 1075798) hydroids. The coenenchyme and sclerites are princi- collected on Pratt Seamount at a depth of 1179 m. H) pally composed of high-magnesium calcite (8.9 mol% A close-up view of the same colony in photo G. The MgCO3) with minute amounts of aragonite and calcite distance between the red laser marks in photos E–G is (senior author and S. D. Cairns, unpubl. data). 10 cm. 116 Professional Paper NMFS 23 2. Callistephanus simplex (Nutting, 1909) (continued) Chapter 7 117 Figure 7-3 A map of Alaska showing the distribution of Callistephanus simplex () in Alaska waters. 118 CHAPTER 8 Family Malacalcyonacea incertae sedis As part of their recent revised systematics of the Octo- prising if some of them prove to be unique family-level corallia, McFadden et al. (2022) placed 46 genera and clades. two unaccepted families of corals into a group incertae Alaska corals included in this new family include two sedis. They determined that the phylogenetic positions species of Alaskagorgia (previously placed in the family of these corals were unknown or uncertain. The World Plexauridae, suborder Holaxonia), five species of Cal- Register of Marine Species currently lists these taxa in cigorgia (previously placed in the family Acanthogorgi- the family Malacalcyonacea incertae sedis. The majority idae, suborder Holaxonia), the only species of Cryogor- of these taxa are rare and few specimens are available gia (previously placed in the family Plexauridae), and for molecular analyses. The mtMutS phylogeny for all the only species of Elenanthus (previously placed in the of the Alaska genera placed in this group (Alaskagorgia, family Clavulariidae). These corals have a limited and Calcigorgia, Cryogorgia, and Elenanthus) do not fit into peculiar distribution in Alaska (Fig. 8-1). All species are any of the 55 currently accepted families studied by principally found in the Aleutian Islands except for two McFadden et al. (2022), even though all but Elenanthus —Calcigorgia spiculifera Broch, 1935 and Calcigorgia are common where they occur and many specimens beringi (Nutting, 1912)—that are also found in the east- have been collected for study. The authors indicate that ern Gulf of Alaska. Corals in this family are found at their phylogenetic positions require further confirma- depths between 12 and 2210 m. tion from genomic data and that it would not be sur- Figure 8-1 A map of the North Pacific Ocean showing the distribution of corals in the family Malacalcyo- nacea incertae sedis () in Alaska waters. Chapter 8 119 1. Alaskagorgia aleutiana Sánchez and Cairns, 2004 Description (Adapted from Sánchez and Cairns, sclerite types. The thin surface layer contains small 2004) Colonies are stout, upright, and sparsely double heads; the thick middle layer contains the gastric branched; they have up to eight lateral branches but are cavities of the retracted polyps and oval capstans with typically monopodial in smaller specimens (<30 cm). elaborate ornamentation. The inner layer (axial sheath) The largest known specimen is 84 cm in height with also contains capstans that are less ornamented and typ- branch diameters of 15–22 mm. Branch tips are more ically octoradiate. or less clavate. The holdfast is robust; the axial skele- ton is woody, somewhat flexible but strong, and dark Remarks Alaskagorgia aleutiana is typically found in brown with faint linear striations. The coenenchyme is complex habitats and is an excellent indicator species of thick, can easily be peeled away from the axial skeleton coral garden habitat. Associates include several species in sheets, and is dark yellow to various shades of orange of ophiuroids. The coenenchyme and sclerites are prin- in life, pale orange when dried, and may turn blackish cipally composed of high-magnesium calcite (9.1 mol% when frozen. For best preservation, specimens should MgCO3) with minute amounts of aragonite (senior au- not be frozen for an extended period of time. thor and S. D. Cairns, unpubl. data). Polyps are scattered on all sides of the branches and housed within raised circular and oval calyces that are Distribution Uncommon. In Alaska – known only up to 5.5 mm in diameter and have eight prominent from the Aleutian Islands (south of Yunaska Island to notches. The anthocodiae are fully retractile into a thick east of Stalemate Bank) including Petrel Bank (Fig. 8-2). cortex but are often exsert in preserved specimens. Elsewhere – not reported. The tentacular portion of the polyps is densely armed with pointed, tuberculate spindles (>0.6 mm in length) Habitat In Alaska – found in complex habitat in ar- that may have projecting processes. The thick coenen- eas of moderate current, growing on bedrock, boulders, chyme has three distinct layers, each with predominant cobbles, and siltstone (in deeper waters only). Collected Figure 8-2 A map of the Aleutian Islands showing the distribution of Alaskagorgia aleutiana () in Alaska waters. 120 Professional Paper NMFS 23 specimens range in depth from 87 to 824 m but it has view of a fresh A. aleutiana specimen (USNM 1484146) been observed on video footage to depths between 1030 collected on Petrel Bank, central Aleutian Islands, at and 1044 m in northern Kanaga Pass and to 1255 m on a depth of 154 m. D) An A. aleutiana colony (USNM Amchixtam Chaxsxii (senior author, personal observ.). 1115604) collected off Cape Moffet, Adak Island, cen- tral Aleutian Islands, at a depth of 140 m. E) An A. Photos A) A fresh whole (except the holdfast) A. aleutiana colony photographed northwest of Tanaga aleutiana colony (USNM 1484182) collected near the Island, central Aleutian Islands, at a depth of 180 m. F) Delarof Islands, central Aleutian Islands, at a depth of A close-up view of the lower stalk of the same colony 120 m. B) A fresh whole A. aleutiana colony (USNM in photo E. The distance between the red laser marks in 1484185) collected southwest of Tanaga Island, cen- photos D–F is 10 cm. tral Aleutian Islands, at a depth of 87 m. C) A close-up Chapter 8 121 2. Alaskagorgia splendicitrina Horvath and Stone, 2018 Description The holotype colony is flabellate with nu- Remarks The holotype collected in the far western merous (about 34), robust branches that are somewhat region of the Aleutian Island Archipelago is the only convoluted, turning in all directions, but not anasto- known specimen, despite extensive collections from mosing. Branches are about 5 mm in diameter (not in- fisheries and research surveys throughout the region. cluding polyps); tips are noticeably clavate. The colony This species most likely radiated from the much less is upright (approximately 12 cm in height) but rather explored region to the west, including the Commander squat and broad (approximately 25 cm in width). The (Komandorskie) Islands and Kamchatka Peninsula of holdfast is cup-shaped with irregular margins and is the Russian Far East, and is endemic to the Northwest completely covered with coenenchyme. The axial skel- Pacific region. eton is woody, quite flexible, and black near the base Preliminary genetic analyses have been undertaken (Thoma, 2013) and additional work is ongoing to deter- but olive-brown distally. The coenenchyme is a bright, mine the similarities at the molecular level between this vibrant lemon-yellow when freshly collected and pre- species, A. aleutiana, and an apparent third undescribed sumably in situ, but turns a dull olive-brown when fro- species of Alaskagorgia from deep water (824 m) in the zen and dried. central Aleutian Islands (Horvath and Stone, 2018). The The polyps are large, exposed, and numerous, scat- status of the genus regarding higher level taxa, such as tered all over the branches with few or none on the stalk family, are also being investigated with molecular genet- and holdfast. The anthocodiae are fully retractile into ics (Everett8). a thick coenenchyme but many are preserved exsert in The holotype hosted 23 brittle stars (class Ophiu- the holotype. The cortex is conspicuously raised around roidea) including 20 Ophiosemnotes pachybactra and the polyp openings, forming short, cylindrical protuber- three Astronebris tatafilius (Hendler9). ances as broad, rounded domes or grooved pyramids up to 3.0 mm in height and diameter, round to oblong Distribution Rare. In Alaska – known only from the in shape, and eight-lobed. Tentacles appear to be rather type locality on Stalemate Bank in the far western re- long, extending beyond the aperture edge about 3 mm, gion of the Aleutian Islands (Fig. 8-3). Elsewhere – not and joined together to form a tall grooved column. reported. Sclerites are generally those typical of the genus. Anthocodiae are rather densely armed with somewhat Habitat The holotype was collected at a depth of 184 pointed, tuberculate spindles, many nearly 0.5 mm in m, likely in an area of small cobbles, pebbles, and sand length. The coenenchyme has three layers. The surface with moderate bottom currents. layer is quite thin and contains small (0.07–0.09 mm in Photos A) The dried Alaskagorgia splendicitrina ho- length) oval capstans. The middle layer is thick, contain- lotype (USNM 1498741; whole colony) collected on ing numerous dense gastric cavities, and has oval cap- Stalemate Bank, western Aleutian Islands, at a depth of stans (0.03–0.18 mm in length). The inner layer is quite 184 m. The holdfast is highlighted in the white circle. B) thin, containing the axial sheath and simple or modified A close-up view of the same specimen in photo A. octoradiate sclerites (0.10–0.20 mm in length). Most sclerites are a very pale yellow. 8Everett, M. 2020. Personal commun. National Marine Fisheries Service, Northwest Fisheries Science Center, 2725 Montlake Blvd E, Seattle, WA 98112. 9Hendler, G. 2019. Personal commun. Natural History Museum of Los Angeles County, 900 Expedition Boulevard, Los Angeles, CA 90007. 122 Professional Paper NMFS 23 2. Alaskagorgia splendicitrina Horvath and Stone, 2018 (continued) Figure 8-3 A map of the Aleutian Islands showing the distribution of Alaskagorgia splendicitrina () in Alaska waters. Chapter 8 123 3. Calcigorgia beringi (Nutting, 1912) Description (Adapted from Nutting, 1912) Colonies lands (Akutan Pass to southwest of Buldir Island) and are either flabellate or bushy, somewhat spindly and eastern Gulf of Alaska (Shutter Ridge). Elsewhere – not highly branched in the largest colonies. Branching is ir- reported, but there is one specimen identified by Matsu- regular and to at least the 5th order. The largest colo- moto et al. (2019) from off Cape Flattery, Washington, nies are 55 cm in height and 35 cm in width. Holdfasts as Calcigorgia japonica. However, we tentatively list are strong, circular, and typically covered with coenen- that specimen as C. beringi (which it much more closely chyme. Tubular calyces are very prominent, mainly lat- resembles) until the specimen can be re-examined. eral and often anterior in position. The back of the col- ony is often bare. Typical calyces are 2.5 mm in height Habitat In Alaska – occurs in scattered patches on and 1.3 mm in diameter. Eight broad shallow longitudi- bedrock, boulders, cobbles, and occasionally hexacti- nal folds near the margin extend upward over the bases nellid skeleton; often in areas of moderate current; and of the tentacles and form a thick crust on the dorsal at depths between 87 and 1247 m. Video observations surfaces and eight lobes to the calyx margin. from the central Aleutian Islands indicate that this spe- The sclerites are very small, short terete spindles, and cies is present at depths to 1933 m (senior author, per- densely tuberculate with the tubercles forming regular sonal observ.). The holotype (USNM 30044) was col- whorls around the body of the sclerite. Sometimes there lected southwest of Buldir Island with a beam trawl is a girdle without tubercles around the center, forming from the U.S. Fisheries Steamer Albatross on 6 June double spindles or double heads. 1906 at a reported depth of 1914 m. However, we mea- The central axis is woody, flexible, and black in color sured a depth of only 212 m at the reported collection (nearer the holdfast) to dark brown (distally). The coen- location from modern National Oceanic and Atmo- enchyme is light orange to pinkish orange; in some spec- spheric Administration nautical charts. imens the coenenchyme is so thin that the axial skeleton is visible beneath. Photos A) A fresh whole C. beringi colony (USNM 1481959) collected in southern Amchitka Pass, central Remarks Associated fauna include pleustid and Aleutian Islands, at a depth of 92 m. B) A close-up view stenothoidid amphipods, the hydroid Bonneviella su- of the same specimen in photo A. C) Fresh whole C. perba, and ophiuroids (Watling and Stone7). It is preyed beringi colonies (USNM 1481960) collected on Shutter upon by the spiny red sea star (Hippasteria phrygiana) Ridge, eastern Gulf of Alaska, at a depth of 244 m. D) and calliostomatid snail Otukaia beringensis in the cen- The same colonies in photo C with polyps fully extend- tral Aleutian Islands (Stone, 2014; Tuskes and Clark, ed, just before collection. E) A C. beringi colony photo- 2018). The coenenchyme and sclerites are principally graphed on Amchixtam Chaxsxii in the central Aleutian composed of high-magnesium calcite (8.8–9.0 mol% Islands at a depth of 903 m. F) Two C. beringi colonies MgCO3) with small amounts of aragonite and calcite (left: USNM 1659742; right: USNM 1481957) collect- (senior author and S. D. Cairns, unpubl. data). ed on Amchixtam Chaxsxii at a depth of 861 m. G) A C. beringi colony (USNM 1659742) with polyps fully Distribution Uncommon. In Alaska – this species has retracted. The distance between the red laser marks in a highly disjunct distribution (Fig. 8-4); the Aleutian Is- photos D–G is 10 cm. 124 Professional Paper NMFS 23 3. Calcigorgia beringi (Nutting, 1912) (continued) Chapter 8 125 Figure 8-4 A map of the North Pacific Ocean showing the distribution of Calcigorgia beringi () in Alaska waters. 126 Professional Paper NMFS 23 4. Calcigorgia gigantea Matsumoto et al., 2019 Description Colonies are robust but small; the larg- species given by the authors—gigantea (Latin for giant est of three known colonies (holotype) is 10 cm in or very large). height and 7 cm in width. The central axis is quite ro- bust, about 6 mm wide (without coenenchyme), woody, Distribution Rare. In Alaska – central and western and somewhat flexible. The holdfast is well defined and Aleutian Islands (Fig. 8-5). Known from only three somewhat discoid. The holotype has only two second- specimens collected near Amchitka Pass (Tanaga Pass ary branches. Polyps are non-retractile and quite large to north of Semisopochnoi Island). Elsewhere – not re- (9–10 mm in height by 5–6 mm in width). Tentacles ported. lack sclerites. Polyp sclerites consist of poorly developed Habitat Attaches to hard substrate, most likely cob- clubs (≥0.6 mm in length) and spindles (≥0.9 mm in bles and pebbles, in areas of moderate current, and at length), both with simple tubercles. depths between 135 and 381 m. Ethanol-preserved specimens are light orange. The axial skeleton is a medium brown. Photos A) The preserved (in ethanol) C. gigantea ho- lotype (USNM 1013069; whole colony) collected in Remarks This species is unique among the genus with Tanaga Pass, central Aleutian Islands, at a depth of 381 its very large polyps and sclerites, hence the name of the m. B) A close-up view of the same specimen in photo A. Chapter 8 127 Figure 8-5 A map of the Aleutian Islands showing the distribution of Calcigorgia gigantea () and C. matua (♦) in Alaska waters. 128 Professional Paper NMFS 23 5. Calcigorgia japonica Dautova, 2007 Description (Adapted from Dautova, 2007) Colo- Distribution Locally common. In Alaska – the Aleu- nies are often monopodial, sometimes branched, and tian Islands from the Islands of Four Mountains to Am- somewhat planar. The largest colony is 31 cm in height chixtam Chaxsxii (Fig. 8-6). Elsewhere – Sea of Japan, with 12 branch tips, although most are less than 8 cm Sea of Okhotsk, and a single record identified by Mat- in height with four or fewer branches. Branching is a sumoto et al. (2019) from off Cape Flattery, Washing- mixture of openly lateral and irregularly dichotomous, ton. However, that specimen more closely resembles C. up to 5th order branching. The central axis or stalk and beringi and not C. japonica so we have tentatively listed branches have a width of 2–4 mm and are wiry, with a it as such until it can be re-examined. leathery coenenchyme consisting of a thick outer layer with a smooth surface and a very thin, semi-transparent Habitat In Alaska – found in scattered patches some- inner layer. The naked, bare part of the stalk is highly times densely (up to 15 colonies/m2 in Adak Canyon) variable in length and may range close to almost half of where it attaches to cobbles and pebbles, rarely on the colony. The colony is attached to the substrate with bedrock and boulders. Collected specimens range in a nearly circular holdfast. The central axis is black to depth from 57 to 2095 m but video observations (se- dark brown, woody, and flexible. nior author, personal observ.) indicate this species rang- Large non-retractile polyps (≥10 mm in length by 4 es to depths of 2210 m in the central Aleutian Islands. mm in width) are situated irregularly apart at distances Elsewhere – found in areas of pebbles, sand, and silt at of 2–10 mm. Tentacle sclerites include flattened elon- depths between 228 and 900 m. gated bodies (0.7–0.9 mm in length), clubs (0.09–0.14 mm in length) with plump or elongated heads, and small straight spindles (≥0.12 mm in length). Polyp body wall Photos A) A fresh whole C. japonica colony (CAS sclerites are clubs (0.11–0.14 mm in length), warty club- 234581) collected on Amchixtam Chaxsxii, central like spindles (≥0.15 mm in length), capstans (0.15–0.17 Aleutian Islands, at a depth of 399 m. B) A fresh mm in length) with girdled warts and plump terminal whole C. japonica colony (CAS 234582) collected in tufts, and spindles (0.15–0.17 mm in length). The coen- northern Amchitka Pass, central Aleutian Islands, at enchyme consists of leafy clubs (0.08–0.09 mm in length) a depth of 822 m. C) A preserved (in ethanol) whole and sparse warty clubs (0.13–0.15 mm in length), both C. japonica colony (USNM 1004642) collected north with well-developed heads and blunt warty handles, and of Kanaga Island, central Aleutian Islands, at a depth warty spindles (13 mm in length) and capstans (0.15– near 300 m. D) A preserved (in ethanol) whole C. ja- 0.17 mm in length). ponica colony (USNM 1006154) collected in southern Color of colonies in life is light orange, sometimes Amchitka Pass, central Aleutian Islands, at a depth of with purple hues and light purple to brilliant violet. 280 m. E–G) Calcigorgia japonica colonies observed Specimens preserved in ethanol are light orange. on Amchixtam Chaxsxii at depths of 557 m (E), 862 m (F), and 862 m (G). The distance between the red Remarks Associated organisms include ophiuroids. laser marks in photos E–G is 10 cm. Chapter 8 129 130 Professional Paper NMFS 23 5. Calcigorgia japonica Dautova, 2007 (continued) Figure 8-6 A map of the Aleutian Islands showing the distribution of Calcigorgia japonica () in Alaska waters. Chapter 8 131 6. Calcigorgia matua Dautova, 2018 Description The Alaska specimen, the largest known, species most closely resembles C. beringi, differing is planar and has dimensions 14 cm in height and 15 cm mainly in having very short distinct clubs in the polyps, in width, with branches that are 4–5 mm in diameter. which are entirely lacking in C. beringi. Our observa- Branching is irregular and to the 5th order. The Alaska tions, however, are that this species superficially most specimen was collected without its holdfast but it is closely resembles two other corals in this group, Alas- likely stout and strong. The coenenchyme is quite thick kagorgia aleutiana and Cryogorgia koolsae Williams, and easily peels off in sheets in the preserved specimen. 2005. Prominent tubular calyces are relatively widely spaced and situated irregularly around all sides of the branch- Distribution Rare. In Alaska – known from only a es. Polyps are cylindrical, arranged spirally around the single specimen collected west of Semisopochnoi Island, branches, 2–3 mm in height by 1 mm in width, and western Aleutian Islands (Fig. 8-5). Elsewhere – Sea of slightly widened near the base. Tentacle sclerites are Okhotsk including the Kurile Islands (Russia). scales (0.12 mm in length) with a few simple tubercles; polyp sclerites are small clubs (≥0.10 mm in length) Habitat In Alaska – attaches to hard substrate at with simple tubercles; and coenenchyme sclerites are depths around 300 m. Elsewhere – found in areas of capstans, small spindles, and cylinders (≥0.12 mm in sand and pebbles at depths between 300 and 900 m. length) with simple or complex tubercles. The skeletal axis is woody but relatively stiff, and Photos A) A dried whole (except the holdfast) C. ma- light black in color. The coenenchyme in the dried speci- tua colony (USNM 1006226) collected southwest of men is light orange with a yellow hue. Semisopochnoi Island, western Aleutian Islands, at an approximate depth of 296 m. B) A close-up view of the Remarks Matsumoto et al. (2019) indicate that this same specimen in photo A. 132 Professional Paper NMFS 23 7. Calcigorgia spiculifera Broch, 1935 Description (Adapted from Matsumoto et al., 2019) Distribution Locally common. In Alaska – the eastern Colonies are mostly planar or flabellate but some colo- Gulf of Alaska including the inside waters of Southeast nies are slightly bushy. Maximum colony size is about Alaska and the Aleutian Islands from north of the Kren- 50 cm in height and width. Branching is a mixture of itzen Islands to west of Kiska Island (Fig. 8-7). Else- openly lateral and irregularly dichotomous. Typical where – Sea of Okhotsk (Sakhalin and Kurile Islands) branch width is 6–8 mm, up to 5th order branching, and and northern Vancouver Island and Queen Charlotte large colonies may have up to 65 branch tips. Sound, British Columbia. The cylindrical polyps are up to 3 mm in height and 1.6 mm in width. The polyps are armed with tuberculate Habitat In Alaska – found in small discrete patches, spindles (0.18–0.3 mm in length), more or less club-like, often in areas of high to moderate current where it at- and integrating with the spindles (≥0.36 mm in length) taches to bedrock, boulders, cobbles, and occasion- of the polyp body. The coenenchyme contains small ally bivalve shells. Collected specimens range in depth capstans, double heads, and cylinders (0.07–0.11 mm from 12 to 441 m but video observations in the central in length) with indistinct waists integrating with belted Aleutian Islands (Stone, 2014) indicate that this species spindles (0.12–0.15 mm in length), a few crosses, and ranges to depths of 512 m there. Elsewhere – the type irregular forms. specimen was collected in the Sea of Okhotsk at a depth The axial skeleton is black to dark brown, woody, of 165 m. Found on bedrock at depths between 158 and flexible. The coenenchyme is light to dark orange and 440 m (Sea of Okhotsk) and between 40 and 284 in color but occasionally white in the eastern Gulf of m (British Columbia). Alaska and northern British Columbia. Photos A) A fresh whole Calcigorgia spiculifera colo- ny (USNM 1484124) collected south of Semisopochnoi Remarks This species differs from all congeners in Island, central Aleutian Islands, at a depth of 27 m. B) having polyp spindles and club-like sclerites instead of A fresh whole C. spiculifera colony (USNM 1484107) real clubs. This is the most common shallow-water (<50 collected near Adak Island, central Aleutian Islands, m depth) octocoral in Alaska and consequently its ecol- at a depth of 24 m. C) A close-up view of a fresh C. ogy has been relatively well studied (Stone and Wing, spiculifera specimen (USNM 1484112) collected south 2001; Stone et al., 2017). It is a slow-growing octo- of Semisopochnoi Island at a depth of 22 m. D) A C. coral (linear growth rate of 6.0 mm/year) and, despite spiculifera colony photographed south of Semisopoch- its small size, would require 60 years or more to grow noi Island at a depth of 24 m. Polyps are fully retracted. to maximum size (Stone et al., 2017). This species is a E) A C. spiculifera colony photographed north of Little gonochoristic (i.e., separate sexes) brooder that appears Tanaga Island, central Aleutian Islands, at a depth of to spawn at least annually. The species harbors a diverse 25 m. Polyps are fully expanded. F) A patch of small suite of ophiuroids. The coenenchyme and sclerites are C. spiculifera colonies, including the white morph, on principally composed of high-magnesium calcite (9.5– Shutter Ridge, eastern Gulf of Alaska, at a depth of 88 10.4 mol% MgCO3) with minute amounts of aragonite m. The distance between the red laser marks in photos (senior author and S. D. Cairns, unpubl. data). D–F is 10 cm. Chapter 8 133 134 Professional Paper NMFS 23 7. Calcigorgia spiculifera Broch, 1935 (continued) Figure 8-7 A map of the North Pacific Ocean showing the distribution of Calcigorgia spiculifera () in Alaska waters. Chapter 8 135 8. Cryogorgia koolsae Williams, 2005 Description (Adapted from Williams, 2005) Colo- are small for the family; anthocodial sclerites are lack- nies are upright, stout, branching in one plane, and ing, and all sclerites are colorless. typically candelabra-like. Branching is irregular (lat- eral) and relatively sparse; the largest specimen has Remarks Larger colonies in shallow water often serve only 22 branch tips and 4th order branching. Termi- as an elevated perch for the brittle star Gorgonocepha- nal branches are elongate and slightly clavate. These lus eucnemis and can also be heavily populated by cap- specimens measure up to 44 cm in height and 19.5 rellid amphipods. The microbial ecology of this species cm in width, often with the lower 31–41% of the has been studied from specimens collected in the central colony unbranched. Monopodial colonies are typical- Aleutian Islands (Gray et al., 2011). ly less than 18 cm in height and apparently have not reached the branching stage yet (see photos F and G Distribution Uncommon. In Alaska – known only on next page). Holdfasts are cup-shaped, covered with from the Aleutian Islands (Amukta Pass to south of Bul- coenenchyme, but relatively easy to detach from the dir Island) including Petrel Bank (Fig. 8-8). Elsewhere substrate (i.e., when collecting colonies the holdfast – not reported. will detach before the axis will tear). The axial skel- eton is woody, quite flexible, nearly impossible to tear, Habitat Occurs on hard rock including bedrock, and dark brown to black in color. The coenenchyme is boulders, cobbles, and pebbles in areas of moderate cur- thick, easily removed from the axial skeleton in sheets, rent, and at depths between 18 and 412 m, but rarely bright to medium orange in life, slightly paler orange deeper than 150 m. in ethanol, and yellow when dried. A highly variable section of the stalk is naked or Photos A) A fresh whole (except the holdfast) barren (devoid of polyps); polyps may start near the Cryogorgia koolsae colony (USNM 1484133) collected base or not until the first secondary branches. The north of Kasatochi Island, central Aleutian Islands, at naked section of the stalk is often covered with fila- a depth of 65 m. B) A dried whole C. koolsae colony mentous brown algae in shallow-water specimens (USNM 1484130) collected south of Adak Island, cen- (less than about 80 m depth), and occasionally also tral Aleutian Islands, at a depth of 54 m. C) Two of the covered with demosponges and hydroids. The polyps largest known C. koolsae specimens (USNM 1011357; are retractile but exhibit various states of exsertion in whole colonies; fresh) collected on Petrel Bank, central preserved specimens. Polyps are large (up to 8 mm in Aleutian Islands, at a depth of 175 m. D) A close-up length when fully expanded, see photo F), irregular- view of a fresh C. koolsae specimen (USNM 1484131) ly cover most of the branch surfaces, and are much collected north of Adak Island, central Aleutian Islands, lighter in color than the coenenchyme, almost white, at a depth of 70 m. E) A C. koolsae colony observed in situ (see photo F on next page). The polyp mounds near Kagalaska Island at a depth of 100 m. F) A mono- (not true calyces) are hemispherical to cone-shaped or podial C. koolsae colony with polyps fully extended, somewhat cylindrical. photographed south of Semisopochnoi Island, central Coenenchymal sclerites of the polyp-bearing branch- Aleutian Islands, at a depth of 25 m. G) A monopodial es, branch tips, and polyp mounds are of several dif- C. koolsae colony with polyps mostly retracted, photo- ferent forms: foliates or leaf-clubs (0.07–0.12 mm in graphed north of Little Tanaga Island, central Aleutian length); seven- and eight-radiates (0.07–0.10 mm in Islands, at a depth of 25 m. H) A C. koolsae colony length); clavate forms (0.08–0.15 mm in length); crosses observed northwest of Adak Island, central Aleutian Is- (0.10–0.11 mm in length); and modified radiates, cylin- lands, at a depth of 130 m. The distance between the ders, and spindles (0.07–0.15 mm in length). Sclerites red laser marks in photos E–H is 10 cm. 136 Professional Paper NMFS 23 8. Cryogorgia koolsae Williams, 2005 (continued) Chapter 8 137 Figure 8-8 A map of the North Pacific Ocean showing the distribution of Cryogorgia koolsae () in Alaska waters. 138 Professional Paper NMFS 23 9. Elenanthus cf. violaceus Description (Adapted from Sanamyan and Sanamy- count (Sanamyan and Sanamyan, 2020). Colonies are an, 2020) Elenanthus violaceus Sanamyan and Sanamy- relatively large (up to 8–10 cm in length), typically elon- an, 2020 was recently described from specimens collect- gate (up to 3–4 cm in width), and with up to 36 polyps. ed near the Kurile Islands (Russia). Molecular analysis Polyps are larger than in the original description (up to confirmed that the specimens represented a novel genus 8 mm in length when fully expanded). The Alaska speci- and species of stoloniferan. Colonies in the genus Ele- mens have not yet been examined to verify that they too nanthus are small, typically only several centimeters in completely lack sclerites so accordingly we list the speci- diameter, in the form of a low pad with slightly tucked men as Elenanthus cf. violaceus. Although colonies are edges, and attached to the substrate by the middle part small, they are quite conspicuous due to their brilliant of the lower surface. The holdfast is very short but magenta to fuchsia color. thick, and weakly attached. All polyps are the same, lo- cated evenly over the entire upper surface of the colony, Distribution Uncommon. In Alaska – known from tightly packed together, and fully retractile. The surface four collected specimens and many video observations of the coenenchyme with retracted polyps is perfectly in the central Aleutian Islands (Amlia Island to Amchix- smooth with small elevations. tam Chaxsxii) including Petrel Bank (Fig. 8-9). Else- Colonies are small (up to 30 mm in length and 12 where – the type specimens were collected near Matua mm in width). Fully expanded polyps are up to 5 mm in Island (Kurile Islands, Russia) in the Sea of Okhotsk, length. The most important distinguishing feature of this Northwest Pacific Ocean. species and genus is the complete absence of sclerites in the tissues. Color of colonies in life is a uniform bright Habitat In Alaska – occurs singly or in small patch- purple but turns brown in ethanol. es (up to five colonies together) on bedrock, boulders, The Alaska specimens conform almost perfectly with cobbles, and mollusk shell including the Alaska jingle the description and photos provided in the published ac- (Pododesmus macrochisma) at depths between 22 and Figure 8-9 A map of the Aleutian Islands showing the distribution of Elenanthus cf. violaceus () in Alaska waters. Chapter 8 139 352 m. Elsewhere – occurs singly or in small patches on CAS # pending) with polyps retracted, collected near bedrock at depths between 13 and 14 m. Kagalaska Island, central Aleutian Islands, at a depth of 100 m. The colonies are growing on an Alaska jingle. Photos A) A fresh whole E. cf. violaceus colony (CAS D) An E. cf. violaceus colony (CAS # pending) just be- 234583), with some polyps extended, collected near Bo- fore collection near Bobrof Island, central Aleutian Is- brof Island, central Aleutian Islands, at a depth of 25 m. lands, at a depth of 25 m. The specimen in photo A is An unidentified orange Clavularia species, possibly Cla- visible at the top of the photograph. E) An E. cf. viola- vularia sp. A, is pictured at center. B) A close-up view ceus colony photographed near Bobrof Island at a depth of the same specimen in photo A. C) Two fresh whole of 27 m. F) An E. cf. violaceus colony photographed E. cf. violaceus colonies (indicated by the white circles; near Bobrof Island at a depth of 24 m. 140 CHAPTER 9 Family Paramuriceidae (Adapted from McFadden et al., 2022) Paramuriceidids distribution of the two genera in Alaska is limited to the are octocorals; they almost always have a proteinaceous Aleutian Islands (Fig. 9-1). They are mostly deep-water skeletal axis that is hollow with a wide-chambered cen- fauna occupying depths between 78 and 2087 m and tral core. Colonies are erect and mostly branched but grow directly on hard rock (bedrock, boulders, cobbles, may be planar or bushy. The coenenchyme is typically pebbles, and siltstone). very thin. Polyps are monomorphic and are either re- All species of paramuriceidids in Alaska have defini- tractile into prominent calyces or, if non-retractile, pos- tive taxonomy except Acanthogorgia sp., which was re- sess a polyp wall heavily armored with spindles, giving vealed as a separate species from A. spissa Kükenthal, the appearance of a cylindrical calyx. Calyx sclerites are 1908 using molecular genetics (Thoma, 2013). Addi- thornscales or spindles that are different from coenen- tionally, Muriceides cylindrica Nutting, 1912 is known chymal sclerites. from only two specimens that were poorly cataloged af- Of the 25 known genera of Paramuriceidae, only two ter collection and description. All specimens collected in are found in Alaska waters: Acanthogorgia (two species) Alaska waters that fit the description of either A. spissa and Muriceides (two species). Both genera were former- or M. cylindrica should be retained and properly pre- ly included in the now defunct suborder Holaxonia. The served for morphological and molecular studies. Figure 9-1 A map of the North Pacific Ocean showing the distribution of corals in the family Paramuri- ceidae () in Alaska waters. Chapter 9 141 1. Acanthogorgia spissa Kükenthal, 1908 Description (Adapted from Kükenthal, 1908) Colo- occasionally used as refuge by Verrill’s king crab (Paralo- nies are almost always planar, rarely bushy, and up to mis verrilli). The coenenchyme and sclerites are princi- 50 cm in height and 50 cm in width. Colonies have very pally composed of high-magnesium calcite (7.6 mol% dense branching, with numerous side branches emanat- MgCO3) with minute amounts of aragonite and calcite ing from one side of the branching plane at an acute (senior author and S. D. Cairns, unpubl. data). angle. Branches and side branches are somewhat wavy and bent; all branches except primaries are thin and Distribution Rare but locally common (Fig. 9-2). In delicate. The polyps are non-retractile, densely crowded, Alaska – central Aleutian Islands. Known only from basally preferring the branching plane, with a prominent five specimens collected south of Adak Island, includ- crown of sharp spines (actually spindles) around the top ing Adak Canyon, but also observed on video footage just below the base of the tentacles. The polyps are up collected in southern Amchitka Pass (senior author, per- to 2.5 mm in length, and the upper ends of the heavily sonal observ.). Elsewhere – known only from the type spined sclerites of the polyps stand out of the surface locality in Uruga Channel, Japan, at a depth of about creating eight longitudinal ribs up to 0.8 mm in length. 55 m. Note: the collection site was in the immediate lo- The spined spindles are up to 0.9 mm in length; their cation of a fishing port so we suspect that the specimen, round spines stand out only somewhat above the head given its apparent disparate bathymetric displacement of each polyp and merge slowly into the slightly bent, from the known depth range, might have been translo- widely spined lower leg. The tentacles contain acanthose cated to the collection site during the process of clean- plates, which are wide, up to 0.12 mm in length, and ing fishing gear. often nearly club-shaped. Within the coenenchyme are slim, bent, widely spined sclerites of approximately 0.3 Habitat In Alaska – scattered patches on bedrock and mm in length and single three-rayed and smaller forms large boulders and often on siltstone ridge crests grow- of about 0.06 mm in length, with a middle shaft and at both ends three thick, warty rays. Holdfasts are irregularly shaped, covered with coe- nenchyme, and strongly at- tached to the substrate (i.e., when collecting colonies the axis will almost always tear before the holdfast de- taches). The skeletal axis is woody, pliable, and a dark brown to light black in col- or. The coenenchyme is thin and typically olive-green, occasionally with a golden hue in life, but slightly faded in color when preserved. Remarks This species hosts an extremely diverse suite of associates including am- phipods (caprellids, pleus- tids, and stenothoidids), isopods (antarcturids, mun- nids, and eurycopids), poly- Figure 9-2 chaetes (family Syllidae), and A map of the Aleutian Islands showing the distribution of Acanthogorgia spissa () in Alaska nemerteans (Watling and waters. Stone7). Larger colonies are 142 Professional Paper NMFS 23 ing in linear rows. Collected specimens range in depth of the same specimen in photo A. C) A close-up view from 1692 to 1728 m, but video observations in the of the polyps on the same specimen in photo A. D) An central Aleutian Islands (senior author, unpubl. data) A. spissa colony observed in Adak Canyon at a depth indicate this species ranges at depths between 1092 and of 1894 m. E) An A. spissa colony observed in Adak 2087 m. Co-occurs with Acanthogorgia sp. in the same Canyon at a depth of 1896 m. A Verrill’s king crab uses patches where their depth ranges overlap. Elsewhere – the colony as refuge. The distance between the red laser no data available. marks in Photos D and E is 10 cm. F) A patch of A. spissa colonies on a siltstone ridge crest in Adak Can- Photos A) A dried whole A. spissa colony (USNM yon at a depth of 1958 m. The distance between the red 1484165) collected in Adak Canyon in the central Aleu- laser marks is 20 cm. tian Islands at a depth of 1714 m. B) A close-up view Chapter 9 143 2. Acanthogorgia sp. Description This species appears similar to A. spissa Distribution Rare but locally common. In Alaska – but molecular analysis (Thoma, 2013) indicates enough central Aleutian Islands from southwest of Amlia Island differentiation to warrant a separate designation at this to Amchixtam Chaxsxsii (Fig. 9-3). Elsewhere – Acan- time. Colonies are bushier than A. spissa, rarely planar, thogorgia sp. has been reported in the northern North and up to 25 cm in height and 25 cm in width. Color is Pacific Ocean only from off northern Vancouver and typically olive-green with a golden hue. Queen Charlotte Islands, British Columbia. Remarks The World Register of Marine Species lists Habitat In Alaska – scattered patches on bedrock 58 valid species of Acanthogorgia. They are found in and large boulders and often on siltstone ridges. Col- all oceans in both shallow and deep waters. However, lected specimens range in depth from 843 to 1692 m. they are rare in the northern North Pacific Ocean and Co-occurs with A. spissa in the same patches where represented only by A. spissa and this species. This their depth ranges overlap. Elsewhere – Acanthogorgia species is also similar to A. paradoxa Nutting, 1912, sp. have been reported at depths between 876 and 1468 which is known only from the East China Sea (174– m in northern British Columbia and at depths between 247 m depth) and Suruga Gulf, Japan (198 m depth). 372 and 480 m on the Emperor Seamounts in the cen- Several specimens identified only as Acanthogorgia sp. tral-west North Pacific Ocean. have been collected from northern British Columbia at depths between 876 and 1460 m and may be the same Photos A) A large portion of a dried Acanthogorgia species presented here. sp. colony (USNM 1484162) collected southwest of This species hosts a rich suite of associates, including Amlia Island, central Aleutian Islands, at a depth of 843 caprellid and stenothoidid amphipods and munnid and m. B) A close-up view of the same specimen in photo A. antarcturid isopods (Watling and Stone7). C) An Acanthogorgia sp. colony (J2100-4-1; indicated Figure 9-3 A map of the Aleutian Islands showing the distribution of Acanthogorgia sp. () in Alaska waters. 144 Professional Paper NMFS 23 by the white circle) collected in Adak Canyon, central the red laser marks is 10 cm. E) A sloped habitat of silt- Aleutian Islands, at a depth of 1692 m. The distance be- stone and sand in Adak Canyon at a depth of 1712 m, tween the red laser marks is 20 cm. D) A close-up view with both Acanthogorgia sp. and A. spissa present. The of the same colony in photo C. The distance between distance between the red laser marks is 20 cm. Chapter 9 145 3. Muriceides cylindrica Nutting, 1912 Description (Adapted from Nutting, 1912) The holo- and relatively more slender than those of the coenen- type colony is flabellate but quite straggly with an obvi- chyme, which are often terete forms and are densely/ ous main stalk and two secondary branches originating closely tuberculated. The coenenchyme is covered with close to the base. The colony is 15 cm in height, quite small, stout, warty spindles irregularly disposed but po- flexible, and delicate in appearance. The branches are sitioned longitudinally. The polyp crown is very strong somewhat flattened in one plane, and branching is ir- and composed of several transverse rows of small spin- regular and to the 3rd order only. The photo of the ho- dles concentrated in a conspicuous band. The opercu- lotype (photo B) does not show a holdfast but the axial lum is robust and composed of numerous small spindles skeleton is woody, quite fibrous, and brown in color. arranged en chevron on basal parts of the tentacles. All The color of the coenenchyme in the ethanol-preserved sclerites are colorless. holotype is a dull yellowish-brown; the color in situ is unknown. Remarks The information provided with the original Calyces are irregularly distributed on the branches species description is vague and apparently incomplete. but are generally lateral and distributed more on all This species is described from two specimens collected sides nearer the distal ends. They appear to be alter- during the 1906 Northwest Pacific Ocean Expedition nately positioned and generally sparse and well spaced of the U.S. Fisheries Steamer Albatross. The holotype (≥2 mm). The calyces are tubular, typically 2.0 mm in (USNM 30046) was collected near Agattu Island in the height and 1.7 mm in diameter, and surmounted with western Aleutian Islands and is pictured in the original a relatively large polyp. The polyps are apparently not description (Nutting, 1912, plate 11, figs. 1 and 1a). The completely retractile; most are resting above the margin second specimen was collected off southeastern Honshu of the calyx in the preserved specimen. Island, Japan, and the deposition of that specimen is un- The sclerites are exclusively spindles and are small known. Aside from the holotype, the species is known for the family. The spindles of the calyx walls are longer from only a single specimen collected in southern Kana- ga Pass and identified by Dr. F. M. Bayer in 2001 as Muriceides cf. cylindrica (see photo A on next page). Given the amount of sam- pling effort in the western Aleutian Islands, this spe- cies is either extremely rare or some collected speci- mens have been misiden- tified as M. nigra (see M. nigra Remarks). Associated fauna in- clude ophiuroids. Distribution Rare. In Alas- ka – known only from the holotype collected south- east of Agattu Island in the western Aleutian Islands and one specimen collected in southern Kanaga Pass, central Aleutian Islands (Fig. 9-4). Elsewhere – a single specimen collected Figure 9-4 by the U.S. Fisheries Steam- er Albatross near Ose Saki A map of the Aleutian Islands showing the distribution of Muriceides cylindrica () in Alaska waters. Light, southeast Honshu Island, Japan. 146 Professional Paper NMFS 23 Habitat In Alaska – likely attaches to cobbles and m. B) A preserved (in ethanol) large portion of a M. pebbles in areas of moderate current at depths between cylindrica colony (USNM 30046; holotype) collected 393 and 881 m. Elsewhere – depth of 173 m. southeast of Agattu Island in the western Aleutian Is- lands at a depth of 881 m. C) A close-up view of the Photos A) Branches of a large portion of a dried M. same specimen in photo B. D) A close-up view of the cylindrica colony (USNM 1006330) collected south of same specimen in photo B showing a polyp with promi- Kanaga Pass, central Aleutian Islands, at a depth of 393 nent spindles. Chapter 9 147 4. Muriceides nigra Nutting, 1912 Description (Adapted from Nutting, 1912) Colonies parameters were equally represented in either haplotype are flabellate and upright, with robust branches that are (senior author, unpubl. data). So for now, we list them all somewhat convoluted, turning in all directions, but not as M. nigra, but acknowledge that there is intraspecific anastomosing. Colonies are generally uniplanar but the variation in some gross morphological parameters and twisting and turning nature of the branches gives the urge that additional molecular work be undertaken. colony a bushy appearance. The largest known colony This species hosts an extremely rich suite of associates, is 30 cm in height and 21 cm in width but larger colo- including amphipods (families Acanthonotozomellidae, nies have been observed in situ. The branches are highly Caprellidae, Ischyroceridae, Pleustidae, and Stenothoi- variable in width; the proximal main axis is up to 13 dae), isopods (families Janiridae and Munnidae), poly- mm in diameter and distal branches are up to only 2 chaetes (family Nereididae), ophiuroids that are often mm in diameter (excluding polyps). The axial skeleton is cryptically colored and bare sections of the axial skeleton woody, fibrous, quite flexible, and brown under a thick, are often substrate for hydroids (including Bonneviella shiny olive-green sheath, regardless of the color of the superba), small bivalves (family Limidae), and actiniar- coenenchyme. The coenenchyme is thick and is easy to ians (Watling and Stone7). The coenenchyme and scler- peel away in sheets. ites are principally composed of high-magnesium calcite Calyces are distributed all over the main stalk and (8.6 mol% MgCO3) with minute amounts of aragonite branches without any obvious order; they may be less (senior author and S. D. Cairns, unpubl. data). abundant on the lower main stalk but are present on the holdfast. Calyces are tubular, generally crowded, Distribution Locally common. In Alaska – Aleutian and 2–3 mm in diameter. The polyps are only partially Islands (south of Unalaska Island to east of Attu Island) retractile, with the crowns resting at the margins. including Petrel Bank (Fig. 9-5). Video observations (se- Sclerites are mostly spindles, quite small for the fam- nior author, personal observ.) indicate that this species ily, and considerably smaller than those of the conge- ranges to near the northwest end of Bowers Bank. Else- ner M. cylindrica. The coenenchyme bears small, warty where – not reported. spindles arranged longitudinally. The calyce walls are Habitat In Alaska – grows singly or in low-density filled with small spindles that are irregularly arranged patches on bedrock, boulders, cobbles, and pebbles at and vertically positioned. The operculum is dome- depths between 78 and 824 m, often in association with shaped and composed of sclerites arranged en chevron hexactinellid sponges (in particular those in the family on the tentacle bases. Rossellidae). Video observations in the central Aleutian This is one of the most colorful corals found in Alas- Islands indicate this species ranges to depths of 1195 m ka. The coenenchyme is predominantly olive green or there (Stone, 2014). blue, but may range from brilliant purple to dark-or- ange crimson and, rarely, golden-light green. The polyps Photos A) A fresh whole M. nigra colony (USNM are often a different color than the coenenchyme, with 1481899) collected south of Tanaga Island, central bluish/purple highlights most common, and present a Aleutian Islands, at a depth of 489 m. B) A fresh whole striking visual contrast. In comparison, the coloration M. nigra colony (AB02-0116A) collected on Petrel in dried and ethanol-preserved specimens could not be Bank, central Aleutian Islands, at a depth of 87 m. C) more plain and monochromatic. The highlights are faded A close-up view of a fresh M. nigra specimen (USNM and many specimens turn a dark brown or even black. 1481891) collected near Kanaga Island, central Aleu- The ethanol-preserved holotype was one such specimen, tian Islands, at a depth of 139 m. D) A close-up view hence the name nigra given by Nutting (1912). of a fresh M. nigra specimen (USNM 1481897) col- lected south of Tanaga Island, central Aleutian Islands, Remarks Thoma (2013) performed molecular analy- at a depth of 489 m. E) A M. nigra colony (USNM sis on 15 M. nigra specimens that we collected in the 1481897) collected south of Tanaga Island at a depth Aleutian Islands and isolated two mtMutS haplotypes, of 489 m. F) A M. nigra colony (USNM 1481895) col- representing M. nigra (eight specimens) and Muriceides lected south of Bobrof Island, central Aleutian Islands, sp. (seven specimens). Careful study of these specimens at a depth of 160 m. G) A M. nigra colony (USNM from photographs taken immediately after collection in- 1481896) collected south of Bobrof Island at a depth dicated that there are differences regarding the color of of 119 m. H) A close-up view of the same specimen in the coenenchyme and anthocodiae, the general thickness photo G, showing the arrangement and details of the of the stalk and branches, the density of calyces (polyps), polyp structure. The distance between the red laser and the depth of collection between specimens, but these marks in photos E–G is 10 cm. 148 Professional Paper NMFS 23 4. Muriceides nigra Nutting, 1912 (continued) Chapter 9 149 Figure 9-5 A map of the Aleutian Islands showing the distribution of Muriceides nigra () in Alaska waters. 150 CHAPTER 10 Order Scleralcyonacea The Scleralcyonacea includes the former order Pennat- axes of solid calcium carbonate or sclerites fused to- ulacea (now superfamily Pennatuloidea) and the order gether with calcitic material. Helioporacea, most taxa in the former suborder Cal- The World Register of Marine Species currently rec- caxonia, and a morphologically heterogeneous group ognizes 21 families and one superfamily in the order of taxa assigned to the former suborders Alcyoniina, Scleralcyonacea, only six of which are represented in Stolonifera, Scleraxonia, and Holaxonia (McFadden Alaska waters—Chrysogorgiidae, Coralliidae, Kera- et al., 2022). The new order contains taxa with a wide toisidae, Primnoidae, Sarcodictyonidae, and the super- range of growth morphologies, and most taxa have family Pennatuloidea. Family Chrysogorgiidae (Adapted from McFadden et al., 2022 and Cairns et in otherwise featureless, deep-water habitats. Here we al., 2021) Chrysogorgiids are octocorals with a solid, list seven chrysogorgiid taxa: three tentatively identified non-scleritic skeletal axis that is concentrically lay- simply as Chrysogorgia, three tentatively listed as un- ered with calcium carbonate. The axis is smoothly lay- known species of Pseudochrysogorgia, and the recently ered, elliptical or circular in cross section, often brittle, described R. stonei which had long been known from and has a metallic luster. Colonies are erect, typically the region but considered to be the more cosmopolitan branched (sympodial, dichotomous, bottlebrush, or R. pleurocristatus Stearns, 1883. pinnate), and attached by a heavily calcified holdfast or Seventy-seven species of Chrysogorgia are presently anchored with a root-like process if inhabiting soft-sed- known worldwide (WoRMS Editorial Board, 2022). iment habitats. Polyps are monomorphic, non-retractile They are known from all ocean basins and are rela- but contractile, and predominantly arranged uniserially tively common in the central North Pacific and central along the axis and branches. The sclerite complement Indo-Pacific regions, but are notably rare in the northern includes scales, plates, spindles, and rods. North Pacific Ocean where they are represented by only Of the seven accepted genera of Chrysogorgiidae, a few known species: C. japonica (Wright and Studer, only three are found in Alaska waters: Radicipes (a sin- 1889), known from off Japan at a depth of 1875 m, and gle species), and Chrysogorgia and Pseudochrysogorgia C. pinnata Cairns, 2007, known from Davidson and (each represented by three unknown taxa). All three Vance Seamounts off California and Oregon (depths genera were formerly included in the now defunct sub- 1968–3114 m), and near Hawaii (3957 m). Pseu- order Calcaxonia. The distribution of the three genera dochrysogorgia, on the other hand, was described from in Alaska is limited to several seamounts in the Gulf a single species collected in the Coral Sea (Pante and of Alaska Seamount Province and the Aleutian Islands France, 2010), but more recently the original authorities (Fig. 10-1). They are a eurybathic fauna but only oc- and others have identified additional specimens, includ- cupy deep water (1359–4768 m) where they grow di- ing several from Alaska, as Pseudochrysogorgia. rectly on hard rock (bedrock, boulders, cobbles, and The main difference between Chrysogorgia and siltstone) or anchor in soft-sediment habitats. Pseudochrysogorgia is that the former have sympodi- All chrysogorgiid specimens collected in Alaska wa- al branching and the latter have monopodial branch- ters should be retained and properly preserved for mor- ing; that is the principal character that we have used phological and molecular studies. to designate the six unknown Alaska taxa to genus. Chrysogorgiidae occur worldwide and are a eury- Specimens have been collected for five of the six taxa bathic (100–4768 m depth) fauna, but in the North Pa- and we provide simple macro- and microscopic mor- cific Ocean they only inhabit deep-water habitats where phological descriptions here but mitochondrial DNA they attach to hard substrate. Colonies are generally analyses could facilitate future identification to spe- small, often wispy, pale to gold-colored, and all-around cies. The sixth and most distinctly bottlebrush-shaped inconspicuous, so are easily overlooked especially in situ. taxon, Pseudochrysogorgia sp. B, is known only from Radicipes stonei Cordeiro et al., 2017 is exceptional in video observations and we list it here to highlight it as that it often forms dense fields that are notable features a priority for future collection. Chapter 10 151 Genus Chrysogorgia Duchassaing and Michellotti, 1864 (Adapted from Cairns, 2001) Chrysogorgia have Chrysogorgia species based on the complement and branches that spiral around the main stem, or form location of sclerites. Group A (Spiculosae) are those parallel fans at the top of a short trunk. Branches sub- species that have rods and/or spindles in the body divide dichotomously, either from a regular single, as- wall and tentacles. Group B (Squamosae aberrentes) cending spiral around the main stem or forming two are those species that have rods and/or spindles in parallel fans above a short main stem. The polyps are tentacles but not in the body wall. Group C (Squa- few in number, well separated, and large compared mosae typicae) are those species that have only scales; to the branches they emanate from. Sclerites consist rods and spindles are not present. All three species of of spindles, rods, and scales. The axial skeleton has a Chrysogorgia reported here from Alaska belong to brilliant metallic luster; some species are quite flexible. Group C. Versluys (1902) established three distinct groups of Figure 10-1 A map of the North Pacific Ocean showing the distribution of corals in the family Chrysogor- giidae () in Alaska waters. 152 Professional Paper NMFS 23 1. Chrysogorgia sp. A Description The branches are sympodial in shape, This species has a prominent (1.8 cm maximum in forming two parallel fans at the top of a short trunk, diameter), discoidal calcareous holdfast that is creamy and arranged in equal dichotomous fashion as is typi- white in color. The color of the thin coenenchyme and cal for the genus. One fan (photo A, left) was damaged polyps is very pale orange. or preyed upon and has no living tissue but rather a light growth of hydroids. The right fan is largely intact Remarks The tentacular and coenenchymal sclerite but obviously delicate and has dimensions of 10 cm in complement place this taxon in Group C sensu Ver- height and 7.2 cm in width. The coenenchyme is very sluys, 1902. thin and barely noticeable on the main branches. The thick branches are brown, the thinner branches are ol- Distribution Uncommon. In Alaska – known from ive-green, and the axis presents a metallic luster near two specimens collected in the Aleutian Islands; one in the base along the stalk (see photo D). Adak Canyon and the other southeast of Agattu Island The polyps are sparse, relatively large (about 1.4 mm (Fig. 10-2). Examination of video footage at the Adak in length), and present along the branches and appar- Canyon collection site indicated that there were mul- ently always at the branch tips. Body wall sclerites are tiple patches of three to five colonies in the immediate smooth platelets 0.20–0.24 mm in length, about 2.5 vicinity. Elsewhere – unknown. times as long as wide, and arranged transversely across the body wall in an imbricating arrangement. Tentacular Habitat In Alaska – found on siltstone ridges in linear sclerites also appear to be platelets, slightly smaller in rows and in small patches with the primnoids Plumarel- length than the body wall sclerites, and also imbricating. la profunda Cairns, 2011 and Parastenella doederleini Coenenchymal sclerites are also platelets. All platelets (Wright and Studer, 1889) at depths between 1913 and are slightly constricted medially. 2514 m. Figure 10-2 A map of the Aleutian Islands showing the distribution of Chrysogorgia sp. A (), Chrys- ogorgia sp. B (♦), and Chrysogorgia sp. C (▲) in Alaska waters. Chapter 10 153 Photos A) A large portion of a fresh Chrysogorgia two preserved (in ethanol) Chrysogorgia sp. A colonies sp. A colony (USNM 1679741) collected in Adak Can- (USNM 30152) collected southeast of Agattu Island, yon, central Aleutian Islands, at a depth of 2514 m. B) central Aleutian Islands, at a depth of 1913 m. E) A A close-up view of the most distal branch of the same small Chrysogorgia sp. A colony (indicated by the white specimen in photo A. C) A close-up view of the polyps circle) at the Adak Canyon collection site at 2514 m. A of the same specimen in photo A. D) Large portions of Plumarella profunda colony is at right. 154 Professional Paper NMFS 23 2. Chrysogorgia sp. B Description The species is known from five collected more polyps and often having a slightly bushy appear- specimens and multiple video observations made in the ance. immediate vicinity of the collections. The branches are sympodial in shape, with a relatively short (approxi- Distribution Uncommon. In Alaska – known only mately 6 mm) but stout unbranched stem (compared from Derickson Seamount in the Gulf of Alaska Sea- to Chrysogorgia sp. A), and then branched in a single mount Province (Fig. 10-2). Elsewhere – unknown. plane but with multiple small fans that are often in slightly offset planes and giving the appearance of be- Habitat In Alaska – found singly and in small patches ing slightly bushy. The largest colony has dimensions of on boulder, fractured bedrock, and occasionally on ver- 31 cm in height and 32 cm in width. Polyps are rela- tical bedrock at depths between 3358 and 4768 m. tively large and abundant compared to the other two Chrysogorgia species listed here and are located along Photos A) A mostly whole preserved (in ethanol) all sides of the branches including termini. Polyps are Chrysogorgia sp. B colony (USNM 1081178) collect- unifacial (i.e., pointed towards one of the faces), stout, ed on Derickson Seamount at a depth of 4162 m. B) and measure about the same in height and diameter A mostly whole preserved (in ethanol) Chrysogorgia (2.5 mm). Body wall sclerites are thin, smooth, elon- sp. B colony (USNM 1081181) collected on Derick- gate, irregularly shaped platelets up to 0.35–0.45 mm son Seamount at a depth of 4329 m. C) A close-up in length. Tentacular and coenenchymal sclerites are the view of the branching pattern and polyps of the same same but smaller (approximately 0.20 mm in length). specimen in photo A. D) A close-up view of the stalk This species has a small (approximately 6 mm in and main branches of a preserved (in ethanol) Chrys- diameter), circular but flat, calcareous holdfast that is ogorgia sp. B specimen (USNM 1081176) collected strongly attached to the substrate and white in color. on Derickson Seamount at a depth of 3991 m. E) A The color of the axis, visible beneath the very thin coe- Chrysogorgia sp. B colony (USNM 1081176; left of nenchyme, is golden with a brilliant metallic sheen. The the red laser marks) prior to collection on Derickson color of the coenenchyme is light pink to white in life Seamount at a depth of 3991 m. F) A Chrysogorgia and light orange in ethanol. sp. B colony (USNM 1081175; above the red laser marks) prior to collection on Derickson Seamount at Remarks The tentacular and coenenchymal sclerite a depth of 3358 m. G) A close-up view of the polyps complement place this taxon in Group C sensu Ver- (contracted) and holdfast of the same colony in photo sluys, 1902. This species co-occurs with Pseudochrys- F. The distance between the red laser marks in photos ogorgia sp. C but differs in being more robust with far E and F is 10 cm. Chapter 10 155 156 Professional Paper NMFS 23 3. Chrysogorgia sp. C Description Branches are arranged in a sympodial the arrangement of sclerites in the body wall. It may be fashion in one plane, slightly spiraling and bottlebrush- an unknown genus represented by four specimens col- shaped but not like a typical Chrysogorgia. The largest lected at two sites; one was collected by the U.S. Fisher- colony is 14.7 cm in height and about 12 cm in width. ies Steamer Albatross in 1906 and identified by Nutting Colonies are delicate, wispy, and flimsy with dichoto- as Chrysogorgia sp. and the second lot of three speci- mous branching alternately from the main axis. Branch- mens (collected by us in 2004) was also identified as ing begins close to the inconspicuous holdfast. The Chrysogorgia sp. (Pante et al., 2012). largest branches subdivide in multiple planes. All of Associated fauna include large caprellid amphipods the polyps appear to be on the top side of the branches (Caprella sp.; see photo C). and leaning distad. Polyps are elongate; the neck can be slightly narrower than the head, up to 2.3 mm in length Distribution Rare. In Alaska – known only from near and about one-third that in diameter. Body wall, coe- Amlia Island in the Aleutian Islands (Fig. 10-2). Else- nenchymal, and tentacular sclerites are elongate plate- where – unknown. lets (0.20 mm in length and about four times longer than width), arranged longitudinally and not imbricate. Habitat In Alaska – found in small patches on silt- The holdfast is very small and delicate. The stem and stone exposures in areas of low current and at depths branches are golden-green; stem and thicker branches near 1359 m. are slightly metallic. The coenenchyme is relatively thick, pinkish-white and slightly reflective in situ, but Photos A) A fresh whole Chrysogorgia sp. C colony light orange when brought to the surface. (USNM 1659740) collected southwest of Amlia Island in the central Aleutian Islands at a depth of 1359 m. B) Remarks Here we list an unknown species that con- A cluster of three Chrysogorgia sp. C colonies, includ- forms more to the description of Chrysogorgia than ing the colony in photo A (center), just before collec- Pseudochrysogorgia but not completely to either. We tion. The distance between the red laser marks is 10 cm. cannot determine completely how the specimens com- C) A close-up view of the same cluster of the three colo- pare to Pseudochrysogorgia since the description for the nies in photo B. D) A caprellid amphipod (black circle) genus (Pante et al., 2012) provided no information on perched on the same colonies in photo B. Chapter 10 157 158 Professional Paper NMFS 23 Genus Pseudochrysogorgia Pante and France, 2010 (Adapted from Pante and France, 2010) Pseudochrys- sclerites of irregular shape, in the form of plates, scales, ogorgia are monopodial colonies that are often bottle- and rods. The branch coenenchyme contains sclerites in brush-shaped and slightly zigzagging. They have abun- the form of scales and plates that are mostly parallel to dant, dichotomously subdivided branches in multiple the main branch axis. The axis and larger branches are planes that originate in an irregular or regular spiral typically a dull black but characterized by a dark metal- around a relatively tall, upright main stem. The distance lic luster. between branches along the stem is short (<2 cm) and The three species of Pseudochrysogorgia as we list regular. Most polyps are oriented slightly distad, en- them here appear to be strictly limited to Dickens, Pratt, larged somewhat distally at the head, and most often Quinn, and Derickson Seamounts in the Gulf of Alaska equally wide and tall. Polyps have slightly ornamented Seamount Province at depths between 1854 and 4712 m. Chapter 10 159 4. Pseudochrysogorgia sp. A Description Colonies are monopodial, not spiraling are the same but smaller, about 0.25 mm in length, with or bottlebrush-shaped, with alternate pinnate, uniplanar two or three sclerites across a tentacle at the base. branching. Colonies are up to 19 cm in height and 10.5 cm in width, delicate but rigid and easily broken. Branch- Remarks Pseudochrysorgia is currently monotypic, ing does not begin close to the small holdfast, which represented only by Pseudochrysogorgia bellona Pante is partially calcified. The largest branches subdivide in and France, 2010 from the Coral Sea. Here we list an multiple planes, sometimes at perfect right angles. Polyps unknown species represented by one collected specimen occur on both sides of the main axis in no particular pat- and several video observations; given the vast distance tern, including on the stem below the first branching. The between collection sites it is unlikely P. bellona. coenenchyme is relatively thin, and pinkish-white and slightly reflective in situ but light orange to white when brought to the surface or in ethanol. The unbranched Distribution Uncommon. In Alaska – known only stem of some colonies may have an olive-green hue when from Pratt and Quinn Seamounts in the Gulf of Alaska alive. Seamount Province (Fig. 10-3). Elsewhere – unknown, Polyps stand perpendicular to the stem, are spaced 4 but there is one other Pseudochrysorgia specimen mm apart, point upward, and have very long, pointed (USNM 1112696) collected from the North Pacific tentacles. Polyps are slender (twice as high as wide), Ocean (Brooks Bank near Midway Island at 835 m cylindrical, and up to 3 mm in length. The base of depth). the expanded polyp contains eggs and is covered with transversely oriented rods. Body wall sclerites are rods Habitat In Alaska – occurs singly on bedrock, frac- covered with low granules, 0.35–0.40 mm in length, ar- tured bedrock including basalt, and boulders at depths ranged longitudinally on the body. Tentacular sclerites between 1854 and 2227 m. Figure 10-3 A map of the North Pacific Ocean showing the distribution of Pseudochrysogorgia sp. A (), Pseudochrysogorgia sp. B (♦), and Pseudochrysogorgia sp. C (▲) in Alaska waters. 160 Professional Paper NMFS 23 Photos A) A preserved (in ethanol) whole Pseu- colony in photo A in situ showing the branching pattern dochrysogorgia sp. A colony (USNM 1075801) col- and large polyps. E) A Pseudochrysogorgia sp. A colony lected on Pratt Seamount at a depth of 1854 m. B) A photographed on Quinn Seamount at a depth of 2136 close-up view of the same specimen in photo A. C) The m. The distance between the red laser marks in photos same colony in photo A prior to collection. D) The same C and E is 10 cm. Chapter 10 161 5. Pseudochrysogorgia sp. B Description This taxon is known from only a single Dickens Seamount in the Gulf of Alaska Seamount colony photographed on Dickens Seamount in the east- Province (Fig. 10-3). Elsewhere – unknown. ern Gulf of Alaska. We list it here as a separate taxon to highlight it for future collection. It is the only chrysogor- Habitat Grows on bedrock at a depth of 2787 m with giid known from Alaska that is classically bottlebrush- the primnoid Calyptrophora laevispinosa Cairns, 2007. shaped, and it appears to be monopodial with elaborate branching around a stout main stem that is unbranched Photos A) A Pseudochrysogorgia sp. B colony (indi- for the lower 4–5 cm. The approximate dimensions of the cated by the white ellipse) photographed on Dickens colony are 19 cm in length and 8 cm in diameter. Polyps Seamount at a depth of 2787 m. The colony is grow- appear to be abundant and large. The color of the colony ing on the same bedrock outcrop as a large C. laevispi- in situ appears to be white with a slight metallic hue. nosa colony. B) An alternate view of the same colony in photo A (indicated by the white ellipse). The distance Distribution Rare. In Alaska – known only from between the red laser marks in photos A and B is 10 cm. 162 Professional Paper NMFS 23 6. Pseudochrysogorgia sp. C Description This species is known from only two col- fewer polyps and never having a slightly bushy appear- lected specimens and multiple video observations. Colo- ance. This species also occurs with the keratoisidids nies are monopodial, with relatively long unbranched Bathygorgia profunda and Isidella sp. B. stems (compared to the other species) and then branch- ing to multiple small, planar fans. The largest colony Distribution Uncommon. In Alaska – known only has dimensions 23 cm in height and 22 cm in width. from Derickson Seamount in the Gulf of Alaska Sea- Polyps are relatively large (up to 4.7 mm in length) but mount Province (Fig. 10-3). Elsewhere – unknown. sparse (spaced 11–12 mm apart) and located along the top and termini of branches. Polyps consist of a nar- Habitat In Alaska – found singly on bedrock includ- row cylindrical upper portion and a bulbous lower por- ing steep vertical walls, fractured bedrock, and boulders tion up to 2 mm in diameter that contains eggs. Body at depths between 4103 and 4712 m. wall sclerites are slender rods, 0.42–0.50 mm in length, and longitudinally arranged. Tentacular sclerites are Photos A) Preserved (in ethanol) branch fragments of also rods but smaller (0.35 mm in length); the base of a Pseudochrysogorgia sp. C colony (USNM 1081180) the polyp (covering the egg mass) contains irregularly collected on Derickson Seamount at a depth of 4168 m. shaped platelets 0.25 mm in length. B) A close-up view of the same specimen in photo A. C) Colonies have a small, inconspicuous calcareous A Pseudochrysogorgia sp. C colony (USNM 1081177; holdfast that is strongly attached to the substrate. The indicated by the white circle) collected on Derickson axis and branches are light olive-green with a slight me- Seamount at a depth of 4162 m. D) A close-up view of tallic sheen. The color of the coenenchyme in life is light the same specimen in photo C. The holdfast is indicated pink to white and in ethanol-preserved specimens it is by the white circle. The distance between the red laser light orange-yellow. marks in photos C and D is 10 cm. E) The same colony in photo C during collection. Note that the colony was Remarks This species co-occurs with Chrysogorgia growing in a mixed field with B. profunda and Isidella sp. B but differs in being much more delicate with far sp. B. Chapter 10 163 164 Professional Paper NMFS 23 7. Radicipes stonei Cordeiro et al., 2017 Description (Adapted from Cordeiro et al., 2017) (see photo H). Interestingly, although the large ophiu- Colonies are delicate, unbranched, flagelliform, and roid Asteronyx sp. is common at depths where R. stonei spirally twisted or coiled in either direction but more was present, none were observed using the colonies as commonly counterclockwise. The largest colonies are elevated perches, whereas they were common on the up to at least 150 cm in length; axis diameter is up co-occurring pennatuloidean Anthoptilum sp. A. The to 2.9 mm; and the axis tapers slightly to the tip. The congener R. pleurocristatus often serves as a perch for axis is stiff enough to maintain colonies off the sea- Asteronyx loveni in Japanese and Indonesian waters floor and is thus an excellent indicator of water cur- (Fujita and Ohta, 1988). rent direction and strength. Smaller colonies (<70 cm) have polyps near the base but the lower part (up to 12 Distribution Locally abundant. In Alaska – Aleutian cm) of larger colonies is sterile (devoid of polyps). The Islands (south of Unimak Pass to northwest of Tanaga coenenchyme is somewhat thin; the dark axis is visible Island) and on Derickson Seamount in the Gulf of Alas- distally. Colonies arise from a somewhat flexible, calci- ka Seamount Province (Fig. 10-4). Elsewhere – recently fied holdfast that is modified with root-like processes collected west of Cordell Bank in northern California. for attaching to pebbles lodged in the sediment (see This highly disjunct distribution suggests that the spe- photo C). cies could range along the west coast of North America Polyps (1.0–3.0 mm in length) are inclined 45–90° in deeper areas not yet explored. in relation to the axis and arranged uniserially along one side of the axis, producing obvious polypar and ab- Habitat In Alaska – found on siltstone exposures polypar sides of the colony. Polyps are closely packed, and attaches to pebbles in low-relief sand/silt habitats ranging to 5 polyps/cm, but are less closely packed at depths between 1612 m (Stone, 2014) and 3580 m distally. The sclerites are rods and scales located in the (Cordeiro et al., 2017). Elsewhere – depths between polyp walls, coenenchyme, tentacles, and pinnules and 2072 and 2646 m. are described in detail in the original description of the Photos A) The fresh whole R. stonei holotype species. (USNM 1418007) collected northwest of Tanaga Is- Colonies are light orange, light pink, or cream-col- land, central Aleutian Islands, at a depth of 2107 m. B) ored in situ; light orange on deck; and light olive-green A close-up view of the polyps on a preserved (in etha- in ethanol with the characteristic iridescence of other nol) R. stonei specimen (USNM 1418006) collected chrysogorgiids. north of Atka Island in the central Aleutian Islands at a depth of 2153 m. C) A close-up view of the root-like Remarks This species is most similar to Radicipes holdfast of the same specimen in photo B. D) A field pleurocristatus, known from the eastern North Pa- of R. stonei colonies in Adak Canyon, central Aleu- cific (Japan to Indonesia) and from shallower depths tian Islands, at a depth of 2479 m. E) A zoomed-in (629–1301 m). The body wall sclerites for both species view of the same field of colonies in photo D. F) A R. are long, rounded rods but are smaller (<0.8 mm) in stonei colony (J2095-2-1-1) collected southwest of Am- R. stonei than in R. pleurocristatus ( ≥2.0 mm). The lia Island, central Aleutian Islands, at a depth of 2827 coenchymal sclerites of the latter species are also more m. G) A close-up view of the same colony in photo F. strongly sculpted and have what we call a “woody” H) The tip of a R. stonei colony (USNM 1418006) appearance. with an associated pycnogonid (Colossendeis sp.). The Larger colonies are occasionally used by the sea spi- distance between the red laser marks in photos D–F is der (Pycnogonida, Colossendeis sp.) as a resting perch 10 cm. Chapter 10 165 166 Professional Paper NMFS 23 7. Radicipes stonei Cordeiro et al., 2017 (continued) Figure 10-4 A map of the Aleutian Islands showing the distribution of Radicipes stonei () in Alaska waters. 167 CHAPTER 11 Family Coralliidae (Adapted from McFadden et al., 2022) Coralliids are genera found in Alaska waters include Hemicorallium octocorals with or without a skeletal axis of fused or (one species), Paragorgia (six taxa), and Sibogagorgia unfused sclerites. Colonies with axes are usually erect, (one species). The former two genera were previously planar, and variably branched. Colonies without axes included in the now defunct suborder Alcyoniina and are typically hemispherical to digitiform, capitate with the later three were formerly included in the now de- a conspicuous sterile stalk, or on occasion sparsely funct suborder Scleraxonia. Coralliidae are found in all branched or lobate. Polyps are dimorphic with feed- regions of Alaska except the Arctic and some genera are ing autozooids and reproductive siphonozooids and are locally abundant (Fig. 11-1). They are a eurybathic fau- retractile directly into the coenenchyme or cortex sur- na mostly found in deep water (21–2766 m) where they rounding the axis. Polyps may lack sclerites or have a grow directly on hard rock (bedrock, fractured bedrock, mixed complement of spindles, ovals, rods, or radiates. boulders, cobbles, and large pebbles), siltstone, hexac- Coenenchyme sclerites are radiates and spheroids with- tinellid sponge skeleton, and shell. All Hemicorallium, out tubercles and occasionally with blunt rods and spin- Sibogagorgia, and Pseudoanthomastus specimens col- dles. Sclerites are often brightly colored shades of red. lected in Alaska waters should be retained and properly The family Coralliidae contains 14 genera with four preserved for morphological and molecular studies. of them designated in the subfamily Anthomastinae, in- The taxonomy and systematics of octocorals in the cluding the familiar Alaska genera Heteropolypus (three former family Paragorgiidae from the North Pacific taxa) and Pseudoanthomastus (one species). Other Ocean are best described as uncertain and in a state of Figure 11-1 A map of the North Pacific Ocean showing the distribution of corals in the family Coralliidae () in Alaska waters. 168 Professional Paper NMFS 23 flux. Phenotypically they are incredibly diverse. For ex- Here we have adopted the most recent systematics ample, Alaska bubblegum corals, previously designated for the former family Paragorgiidae and list four ac- as Paragorgia arborea (Linnaeus, 1758) until recently cepted species: Paragorgia jamesi Herrera and Shank, and now Paragorgia arborea var. pacifica Verrill, 1922, 2016; Paragorgia arborea var. pacifica and morph are by far the most diverse in Alaska in terms of gross nodosa; Paragorgia stephencairnsi Sánchez, 2005; colony morphology (e.g., size range, branching pattern, and Sibogagorgia cauliflora Herrera et al., 2010. We surface features, and color), zoogeography, and depth additionally list two suspected undescribed species range. However, for the family Paragorgiidae, the mi- (Paragorgia sp. A and Paragorgia sp. B) to highlight croscopic characteristics traditionally used to differen- them for future taxonomic work. Paragorgia arborea tiate species (sclerites) are subtle, and molecular DNA var. pacifica is one of the most abundant, widespread, techniques have proven to be somewhat unreliable and and ecologically important corals in Alaska waters, so conflicting (Herrera and Shank, 2016). Future work we highlight that species by designating two morphs: using new methodologies, such as restriction site-as- arborea and nodosa. Historically, these two morphs sociated DNA sequencing in combination with careful were considered separate species, thus we present study of gross morphological characteristics and ecol- them in that way to retain information regarding dif- ogy, should help elucidate the systematics of the genera ferences in morphology, biogeography, and ecology. Paragorgia and Sibogagorgia. Chapter 11 169 1. Hemicorallium sp. Description This taxon is known in Alaska from only mount but have separate, non-overlapping depth ranges two collected specimens and a third specimen (most- there. ly a skeleton) observed very close to one of the col- Corals in the genera Hemicorallium and Corallium lected specimens (see photo D). The largest of the two are also known as precious corals and are commercially specimens (USNM 1082614) is uniplanar and 20 cm harvested in parts of the world for the jewelry trade. in height and 38 cm in width. The following detailed Coral fisheries are managed by local jurisdictions in description is based on the smaller specimen (USNM some regions and some are regulated under the author- 1075800). The collected corallum (a large branch com- ity of the Convention on International Trade in Endan- plex) is uniplanar and quite fragile, 23 cm in height and gered Species of Wild Fauna and Flora (CITES, 2007; 13 cm in width; the broken basal stem is 8×9 mm in di- Appendix III); consequently, they are one of the most ameter. The whole colony was measured with lasers in well-studied groups of corals worldwide. All coralliids situ at 30 cm in height and 42 cm in width. The branch- are azooxanthellate species and most have life-history ing is irregularly dichotomous; the short distal branch- characteristics that make them particularly vulnerable lets are rarely more than 10 mm in length. Branch to over-exploitation (i.e., K-selected species), including anastomosis does not occur. The axis is smooth; both extreme longevity (75–200 years), late age of maturi- the axis and the coenosteum are pale pink. The distal ty (7–12 years), slow growth (2–20 mm in length and branchlets bear squat, cylindrical (about 1.2 mm in 0.24–1.32 mm in diameter per year, with growth rates height and 1.4 mm in diameter in the contracted state), declining with age), low fecundity, and low natural mor- longitudinally grooved calices, 9–11 on each branch- tality rates (Grigg, 2002; Baco and Shank, 2005; Bra- let occurring in random order but not on the posterior manti et al., 2005; Roark et. al., 2006). Hemicorallium side. Small papillae (siphonozooids) cover the coenos- specimens, possibly H. regale, from Davidson Seamount teum on all sides of the branches. Elongate, slender rods off central California showed a linear growth rate of measuring up to 0.12 mm in length occur, presumably approximately 2.5 mm/year, indicating a colony age of from the calyx; however, the predominant sclerite type about 115 years; however, based on the radial growth is eight-radiates, which are 0.08–0.10 mm in length. rate, an age of up to 200 years is possible (Andrews et The color of the corallum is a uniform light pink; the al., 2005). polyps are very light pink to clear. Distribution Rare. In Alaska – Pratt and Patton Sea- Remarks These specimens are placed in the recent- mounts in the Gulf of Alaska Seamount Province (Fig. ly resurrected genus Hemicorallium (see Ardila et al., 11-2). Elsewhere – unknown. 2012) because they have slender calicular rods, cylin- drical autozooids, and a smooth axis. The closest geo- Habitat In Alaska – occurs singly or in small patches graphical congeners are H. regale (Bayer, 1956) and on bedrock and fractured bedrock (boulder) at depths H. laauense (Bayer, 1956), both from the Hawaiian re- between 1677 and 1779 m. Elsewhere – Hemicoralli- gion. Based on Bayer’s (1956) key to the genus, these um regale is found on hard rock at depths between 300 specimens are most similar to H. laauense, but detailed and 1815 m and H. laauense is found on hard rock at study of additional specimens is needed to make a de- depths between 366 and 1807 m. finitive identification. Herrera and Shank (2016) used molecular genetics techniques (restriction site-associated Photos A) A dried branch from a Hemicorallium sp. DNA sequencing analysis) to assign specimen USNM colony (USNM 1075800) collected on Pratt Seamount 1075800 to H. imperiale-laauense (an unaccepted com- at a depth of 1677 m. B) A close-up view of the same bination); however, if identification to either species (H. specimen in photo A. C) The same colony in photos A imperiale or H. laauense) is correct, it would constitute and B photographed in situ. D) Alternate view of the a huge geographical range extension (about 2800 km) same colony in photo C. A second colony consisting of from the known distribution in the Hawaiian region. mostly a skeleton attached to the substrate is visible to This species is very similar in gross morphology (e.g., the right of the live colony. E) A Hemicorallium sp. col- branching pattern) and color to Paragorgia sp. B and ony (USNM 1082614) photographed during collection the two species could easily be mistaken for each other on Patton Seamount at a depth of 1779 m. The distance in situ. Fortunately, they only co-occur on Patton Sea- between the red laser marks in photos C–E is 10 cm. 170 Professional Paper NMFS 23 1. Hemicorallium sp. (continued) Chapter 11 171 Figure 11-2 A map of the North Pacific Ocean showing the distribution of Hemicorallium sp. () in Alaska waters. 172 Professional Paper NMFS 23 2. Paragorgia jamesi Herrera and Shank, 2016 Background Specimens examined by Sánchez (2005) branched including occasional whiplike colonies. The to describe Paragorgia stephencairnsi were re-examined largest colonies are about 55 cm in height and 30 cm using restriction site-associated DNA sequencing analy- in width with up to 50 branch tips. This morph is not sis and the species complex differentiated into a shal- as robust as the pink form. All branch tips are clavate low and a deep-water sub-clade (Herrera and Shank, and some of the outer branches and tips appear to be 2016). Based on the specimens examined (all from Brit- flattened. Two colonies often grow together with a com- ish Columbia except two from southern California), mon holdfast (see photos A and E). Holdfasts are small those authors somewhat arbitrarily designated the deep but strongly attached; the branches are fragile and easily (1168–1194 m) specimens as P. jamesi and the shal- broken. The colony is generally a light yellow color; the low (32–350 m) specimens as P. stephencairnsi. They autozooid apertures are red to crimson-red, numerous, additionally listed five specimens from Alaska (Herrera not raised, and often aligned in rows; and the polyps are and Shank, 2016, suppl. table 1) ranging in depth from light orange. 171 to 751 m as P. stephencairnsi. Otherwise they not- The pink morph is more robust but typically has ed that the morphology of the two species was nearly fewer branches and is often scraggly in appearance, oc- identical based on the original description (see page 182 casionally planar. The largest colonies are about 75 cm for P. stephencairnsi) except that the lobulated rays of in height and 60 cm in width; some colonies have up to the seven- and eight-radiate surface sclerites in P. jamesi 45 branch tips. All branch tips are clavate and some of have mostly rounded edges, whereas those from P. ste- the outer branches and tips appear to be flattened. This phencairnsi have mostly sharp edges. morph has fewer autozooids that are more randomly Neither the original description for P. stephencairnsi situated on the branches, slightly raised, and with dark- (Sánchez, 2005), the revised morphological diagnosis for er pink autozooid apertures. The coenenchyme is light P. stephencairnsi (Herrera and Shank, 2016), nor the pink and the polyps are light red. Holdfasts are small new description for P. jamesi (Herrera and Shank, 2016) but strongly attached; the branches are fragile and eas- provided much detail on colony morphology, including ily broken. branching pattern and color. Our microscopic examina- tion of one of the Alaska specimens (USNM 1484230) Remarks Seamount colonies host hippolytid shrimps, confirmed that it conformed to the sclerite morphology the sternostylid crab Sternostylus iaspus, and ophiu- described for the species complex but also indicated that roids including Asteronyx species. the characteristic (geometry of the rays of the surface sclerites) used to differentiate the two species was too un- Distribution Uncommon but locally abundant. In reliable to be used to designate the specimens to species. Alaska – specimens have been collected in the eastern Additionally, we found no clear demarcation regarding GOA and the region where the western GOA meets bathymetric distribution (171–944 m) for the Alaska the eastern Aleutian Islands. Specimens and many vid- specimens. However, close examination of museum eo rec ords have been collected from Dickens, Welker, specimens and archived in situ video footage revealed Pratt, Quinn, Murray, Giacomini, and Patton Sea- the presence of two distinct forms with regard to exter- mounts in the Gulf of Alaska Seamount Province (Fig. nal morphological characteristics. Based on that exami- 11-3). This species is particularly abundant on Dick- nation, we separated the complex into the two species ens Seamount. Elsewhere – known only from the type previously described (P. jamesi and P. stephencairnsi). specimens collected in northern (holotype; RBCM 010-00234-004; 2344) and southern (paratype; USNM Description Paragorgia jamesi displays two mor- 1007316) British Columbia. phological forms (morphs) in Alaska: a yellow morph This species appears to be spatially discrete from P. (photos A, B, E, and F) that occurs on the continental stephencairnsi but does overlap spatially with P. arbo- slope and seamounts, and a pink morph (photos C, D, rea var. pacifica, Paragorgia sp. A, and Paragorgia sp. B G, and H) that appears to be restricted to seamounts in on seamounts in the Gulf of Alaska Seamount Province. the Gulf of Alaska Seamount Province (Pratt, Quinn, and Giacomini). Both morphs overlap spatially on the Habitat In Alaska – occurs in patches, rarely singly, seamounts where they co-occur. in rough habitat with moderate current. Attaches to The yellow morph is the dominant one; it often has bedrock, fragmented bedrock, boulder, cobble, large a candelabrum shape, is mostly planar, and is variably pebbles, and siltstone at depths between 372 and 944 Chapter 11 173 Figure 11-3 A map of the North Pacific Ocean showing the distribution of Paragorgia jamesi () in Alaska waters. m. Elsewhere – continental slope habitats at depths be- view of the same specimen in photo C. E) A P. jamesi tween 1168 and 1194 m. colony (yellow morph) photographed on Dickens Sea- mount at a depth of 751 m. F) A close-up view of the Photos A) A P. jamesi specimen (yellow morph) col- same colony in photo E with polyps fully extended. G) lected in the eastern GOA at a depth of approximate- A P. jamesi colony (USNM 1661898; pink morph) sam- ly 500 m. B) A close-up view of the same specimen in pled on Giacomini Seamount at a depth of 762 m. H) A photo A. C) A preserved (in ethanol) branch from a P. close-up view of the same colony in photo G with pol- jamesi colony (USNM 1075764; pink morph) collected yps fully extended. The distance between the red laser on Pratt Seamount at a depth of 920 m. D) A close-up marks in photos E and G is 10 cm. 174 Professional Paper NMFS 23 2. Paragorgia jamesi Herrera and Shank, 2016 (continued) Chapter 11 175 3. Paragorgia arborea var. pacifica Verrill, 1922 Background At one time three separate Paragorgia location and approximate depth where P. stephen- taxa were recognized in the North Pacific Ocean in- cairnsi have more recently been documented (see that cluding Alaska—Paragorgia arborea, P. pacifica Ver- species’ profile, photo F). rill, 1922, and P. nodosa Koren and Danielssen, 1883. The most recent description for P. arborea by Sánchez Paragorgia arborea was originally described from the (2005) was written for the species worldwide, including North Atlantic Ocean (Linnaeus, 1758), then later Alaska, and described robust colonies up to several me- Kinoshita (1913), Hickson (1915), and Verrill (1922) ters high with bubble-like concentrations of autozooids. designated specimens collected in the North Pacific Dense and regular accumulations of autozooid nod- Ocean, including Alaska waters, as P. arborea. Verrill ules are largely restricted to distal and lateral branches, (1922) described P. pacifica from British Columbia at whereas the distal main stem and branches are without the same time that he recognized P. arborea from Alas- nodules. The inter-nodular surface is covered with nu- ka waters and expressed uncertainty whether P. paci- merous and uniformly distributed tiny siphonozooids, fica was actually a variety of P. arborea; he noted that giving the colony a granular texture when the apertures additional examination of the sclerites was warranted. are closed. The medulla is perforated by 5–7 main-stem Herrera et al. (2012) found genetic differentiation of the canals in terminal branches, surrounded by both red North Pacific Ocean P. arborea populations relative to and colorless spindles; the outer medulla has colorless all other world populations (South Pacific, Indian, and sclerites and numerous smaller canals. Polyps are com- North Atlantic Oceans) and suggested that they may pletely retractable into small conical calyces. Tentacle represent a sub-species. Finally, Herrera and Shank sclerites are blunt, stubby ovals with some pointed-like (2016) synonymized P. arborea (in the North Pacific spindles. Surface sclerites are small, uniform 6-radiates; Ocean region only) with P. pacifica. medulla sclerites are long, thin spindles. The WoRMS Editorial Board (2022) currently rec- Paragorgia arborea var. pacifica is one of the most ognizes only P. pacifica as a valid species. However, colorful Alaska corals, along with Muriceides nigra. since arborea was synonymized with pacifica, arborea The color of the coenenchyme is uniform and the pre- has precedence as the senior synonym. Accordingly, we dominant color is pink to light red, but Aleutian Island adopt the designation by Horvath (2019) for the species specimens may also be white, salmon orange, or yel- as P. arborea var. pacifica. low. The latter three color-phases are more common in We do not dispute the validity of a single designation the shallower part (<305 m depth) of the species’ range as P. arborea var. pacifica but have chosen to provide a in the Aleutian Islands, and those specimens appear to separate species profile for the morph that would have have a contagious distribution (senior author, unpubl. traditionally been listed as P. nodosa to highlight both data). The medulla is typically creamy white but occa- the differences and overlap in form, biogeography, and sionally the same color as the coenenchyme or a slight- ecology. Additionally, since the systematics for this spe- ly lighter shade. Similarly, the polyps and tentacles are cies complex seems incomplete, recalcitrant, and likely typically a creamy white but are often pink in speci- to be refined again with advances in molecular DNA mens with pink or reddish coenenchyme. Seamount techniques (see Herrera and Shank, 2016), this presen- colonies are strictly light red to crimson red. Specimens tation might aid in facilitating future work with the spe- retain their natural coloration in ethanol but the color cies complex. dulls somewhat in dried specimens. Description The original description by Verrill (1922) Remarks Paragorgia arborea var. pacifica is often for P. pacifica indicated that colonies are more delicate found in complex habitats and is an excellent indicator and smoother than North Atlantic P. arborea specimens species of coral garden habitat. Paragorgia arborea var. and that the sclerites differ considerably. The medulla is pacifica is also the second largest coral species in Alas- finer and more compact and the longitudinal canals are ka, growing to more than 2 m in height and width with relatively smaller. The stem and axis are quite distinct, basal diameters up to 20 cm, but massive specimens harder, more compact, and a lighter color than the sur- have been collected near New Zealand (Tracey et al., rounding middle layer, with larger and longer sclerites, 2003) measuring approximately 75 cm in basal diam- some forked, and with fewer warts. The coenenchyme eter. Growth rates are largely unknown for Paragorgia contains an abundance of smaller sclerites, mostly very species given the lack of a calcified skeleton, but limited irregular warty spindles of various sizes. The type speci- evidence suggests axial (linear) growth rates for North men was collected in Jervis Inlet, southern British Co- Atlantic P. arborea specimens of 1 cm/year (Mortensen lumbia, at a depth of 18 m. Note that this is the same and Mortensen, 2005), while Sherwood and Edinger 176 Professional Paper NMFS 23 (2009) estimate axial growth rates of 1.62 cm/year and Habitat In Alaska – grows on bedrock, fractured bed- ages of up to 80 years old. rock and boulders, and occasionally on large cobbles Paragorgia arborea var. pacifica produces copious and hexactinellid sponge skeleton (Farrea spp. and Het- amounts of mucus and consequently supports a rich mi- erochone sp.), in areas of moderate current, at depths crobial community (Gray et al., 2011) and numerous between 21 and 1115 m. These observations are both associated fauna including ophiuroids, amphipods, and the shallowest and deepest (Stone, 2014) for the species. the shrimp Heptacarpus moseri (in Zhemchug Canyon). Elsewhere – likely similar habitat to that in Alaska at Seamount specimens are often used by the sternostylid depths near 102 m near the Commander Islands (Kam- crab Sternostylus iaspus as platforms for suspension chatka, Russia), 30–1168 m off British Columbia, 750 feeding and support large numbers of ophiuroids in- m off Oregon, and 274–650 m off California. cluding the large Asteronyx sp. (up to 15 per colony) and polychaetes (often the same color as the host colony Photos A) A fresh whole P. arborea var. pacifica col- coenenchyme). ony (partial view only) (USNM 1106086) collected in The coenenchyme and medulla is composed almost Adak Strait, central Aleutian Islands, at a depth of 90 entirely of high-magnesium calcite (9.1–10.7 mol% m. B) A close-up view of a fresh P. arborea var. paci- MgCO3; senior author and S. D. Cairns, unpubl. data). fica specimen (USNM 1011097) collected northwest of Adak Island in the central Aleutian Islands at a depth Distribution Widespread and locally abundant. In of 160 m. C) A P. arborea var. pacifica colony photo- Alaska – specimens have been collected in the west- graphed near Little Tanaga Island in the central Aleu- ern Gulf of Alaska (near Chirikof Island to Shumagin tian Islands at a depth of 156 m. A Pacific cod (Gadus Bank), the Aleutian Islands (near Unalaska Island to macrocephalus) takes shelter under the colony. D) A P. west of Attu Island) including Bowers Bank, the eastern arborea var. pacifica colony photographed near Adak Bering Sea (north of Unimak Pass to Pervenets Canyon), Island at a depth of 138 m. E) A P. arborea var. pacifica and on Dickens (video observations only), Welker, colony photographed northwest of Tanaga Island in the Pratt, Giacomini, Murray, and Patton Seamounts in the central Aleutian Islands at a depth of 115 m. F) A P. Gulf of Alaska Seamount Province (Fig. 11-4). Else- arborea var. pacifica colony (USNM 1484211) photo- where – the North Pacific Ocean from the Command- graphed on Amchixtam Chaxsxii in the central Aleutian er Islands (Kamchatka, Russia) to Monterey Bay and Islands at a depth of 212 m. A darkfin sculpin (Mala- Davidson Seamount (California). Paragorgia arborea cocottus zonurus) takes shelter under the colony. The is reported in the North Atlantic Ocean from Trond- distance between the red laser marks in photos C–F is heim Fjord (Norway), the Faroe Islands, Iceland, and 20 cm. G) A close-up view of a P. arborea var. pacifica Greenland (Tendal, 1992). Additional records indicate colony (USNM 1075766) prior to sample collection on a patchy distribution from southern Greenland (Davis Welker Seamount at a depth of 1114 m. H) A P. arbo- Strait) to off North Carolina. This species also occurs in rea var. pacifica colony photographed with polyps fully the South Pacific Ocean near New Zealand. extended near the Delarof Islands (Amatignak Island) in Paragorgia arborea var. pacifica overlaps spatially the central Aleutian Islands at a depth of approximately with P. jamesi, Paragorgia sp. A, and Paragorgia sp. B 160 m. on seamounts in the Gulf of Alaska Seamount Province. Chapter 11 177 178 Professional Paper NMFS 23 3. Paragorgia arborea var. pacifica Verrill, 1922 (continued) Figure 11-4 A map of the North Pacific Ocean showing the distribution of Paragorgia arborea var. paci- fica () in Alaska waters. Note that records for the nodosa morph are illustrated in Fig. 11-5; together the two figures show the whole distribution for P. arborea var. pacifica. Chapter 11 179 3A. Paragorgia arborea var. pacifica morph nodosa Background The original description for Paragorgia by large conspicuous longitudinal canals. Sclerites are nodosa by Koren and Danielssen (1883) indicates that mostly small, irregular tuberculate spindles and a few the type specimen was collected off Norway at a depth irregular minute double heads and crosses. The polyp of 549 m. Subsequently, Nutting (1912) provided a de- sclerites are red and the medullar sclerites are white. tailed re-description of the species for a specimen col- We have no doubt why Nutting (1912) regarded the lected on Bowers Ridge (Aleutian Islands) at a depth specimen as P. nodosa since most specimens collected of 655 m during the 1906 U.S. Fisheries Steamer Al- deeper than about 500 m in the Aleutian Islands, in batross expedition. After later re-examination of Nut- general, are stouter, less arborescent, and much more ting’s specimen, Broch (1957) synonymized P. nodosa nodulated. Nodules are more prominent, well defined, with P. arborea, concluding that the two species were and not restricted to the branch tips or branch nubs on identical. Broch (1957) did not explicitly state how they the secondary branches. The branches and nodules have were identical but we assume he meant with regard to a rough appearance because the autozooid apertures are the complement and morphometry of the sclerites. large and raised on conical calyces. The coenenchyme on the primary branches is sometimes coarsely textured Comparison Paragorgia arborea var. pacifica and the and linear striations are apparent. Colonies rarely exceed morph nodosa have very similar geographical distribu- 1 m in height and 1 m in width. Colonies are predomi- tions in Alaska (Figs. 11-4 and 11-5) and do overlap nantly various shades of red, sometimes with a purple slightly in bathymetric distribution, especially on several hue in situ (Aleutian Islands) and orange to crimson red seamounts in the Gulf of Alaska Seamount Province, elsewhere. Polyps are almost always the same color as but otherwise are fairly spatially discrete. The two taxa the coenenchyme. Specimens generally retain their color (one is a morph, the other a variety) differ from each in ethanol, but the color commonly fades when dried; other with regard to several gross morphological char- red specimens fade to orangish-red. acteristics. In general, Paragorgia arborea var. pacifica is more robust, arborescent, obviously fan-shaped, and Remarks Paragorgia arborea var. pacifica morph no- much less nodulated than the morph nodosa. Nodules dosa is often found in complex habitats and is an ex- are mostly restricted to the branch tips or branch nubs cellent indicator species of coral garden habitat. The on the secondary branches. The branches and nodules scarlet king crab (Lithodes couesi) is occasionally ob- are smooth and the autozooid apertures are relatively served on colonies on seamounts in the Gulf of Alaska small and not raised. Colonies often exceed 1 m in Seamount Province (see photo F on page 181), and eel- height and 1 m in width. pouts (including Puzanovia rubra) deposit and guard egg masses on colonies in the Aleutian Islands (see pho- Description (Adapted from Koren and Danielssen, to G on page 181). Other associated fauna include poly- 1883) The original description for the type specimen noid polychaetes and pleustid amphipods (Watling and collected off Norway (549 m depth) indicated that it Stone7), the ophiuroids Ophiacantha sp. and Asteronyx was tree-like with two main branches. The branches sp., and the sternostylid crab Sternostylus iaspus. were knotty and densely populated with polyps and zo- oids. The branches are naked and smooth and slender Distribution Widespread and locally common. In between the nodes or nodules. The polyps are short, re- Alaska – specimens have been collected in the western tractile, and contain numerous sclerites. Gulf of Alaska off Chirikof Island to Shumagin Bank, Nutting’s (1912) description of an additional colo- the Aleutian Islands south of Amlia Island to west of ny indicated that it consisted of a thick stem bearing Buldir Island including Bowers Bank, the eastern Ber- branches on all sides; some were short clavate branches ing Sea from Pribilof Canyon to Pervenets Canyon, and while others were merely irregular nodules (up to 14 Dickens, Welker, and Pratt Seamounts in the Gulf of mm in diameter) seated immediately on the stem. Most Alaska Seamount Province (Fig. 11-5). Elsewhere – un- polyps were situated on the nodules; he suggested that known. a few on the main stem or branches might be the begin- Morph nodosa overlaps spatially with P. jamesi, P. ning of a new branch or nodule. Calyces are low verru- arborea var. pacifica, Paragorgia sp. A, and Paragorgia cae with eight-lobed margins and 3 mm in height and 6 sp. B on seamounts in the Gulf of Alaska Seamount mm in diameter at the base. The polyps are completely Province. retractile and the tentacles bear longitudinal spindles. Siphonozooids are numerous but very small. The me- Habitat In Alaska – grows on bedrock, fractured dulla consists almost entirely of small spindles traversed bedrock, boulders, large cobbles, and occasionally on 180 Professional Paper NMFS 23 Figure 11-5 A map of the North Pacific Ocean showing the distribution of Paragorgia arborea var. paci- fica morph nodosa () in Alaska waters. hexactinellid sponge skeleton (Farrea spp.), in areas of Aleutian Islands at a depth of 746 m, showing the detail moderate current, at depths between 299 and 2022 m. of the polyps and tentacles. E) A coral garden domi- Depth ranges for each region of Alaska are 691–1143 nated by P. arborea var. pacifica morph nodosa colo- m in the western GOA, 470–2022 m in the Aleutian nies on Amchixtam Chaxsxii in the central Aleutian Is- Islands, 299–1077 m in the eastern Bering Sea, and lands at a depth of 921 m. F) A P. arborea var. pacifica 754–1761 m in the Gulf of Alaska Seamount Province. morph nodosa colony (USNM 1075745) prior to sam- ple collection on Dickens Seamount at a depth of 849 Photos A) A fresh whole P. arborea var. pacifica m. A scarlet king crab uses the colony as a perch. G) morph nodosa colony (USNM 1123935) collected A P. arborea var. pacifica morph nodosa colony pho- southwest of Amlia Island in the central Aleutian Is- tographed in northern Amchitka Pass, central Aleutian lands at a depth of 844 m. B) A fresh branch complex Islands, at a depth of 798 m. An eelpout (probably Pu- from a P. arborea var. pacifica morph nodosa colony zanovia rubra) uses the colony as shelter and displays (USNM 1484212) collected in Zhemchug Canyon in cryptic coloration. H) A P. arborea var. pacifica morph the eastern Bering Sea at a depth of 987 m. C) A dried nodosa colony photographed growing on siltstone whole P. arborea var. pacifica morph nodosa colony southwest of Amlia Island, central Aleutian Islands, at (USNM 1484223) collected east of Chirikof Island in a depth of 2022 m—the deepest known P. arborea var. the western GOA at a depth of 691 m. D) A fresh P. pacifica morph nodosa colony in the world. The dis- arborea var. pacifica morph nodosa specimen (USNM tance between the red laser marks is 10 cm in photos E 1123938) collected in Amchitka Pass in the central and G and 20 cm in photo H. Chapter 11 181 182 Professional Paper NMFS 23 4. Paragorgia stephencairnsi Sánchez, 2005 Description (Revised morphological diagnosis from diameter) immediately above the holdfast and then the Herrera and Shank, 2016, amended from Sánchez, 2005) branches are of uniform dimension (1.3–1.5 cm diam- Colonies have robust branches with a white, pink, or red eter), not flattened, with clavate branch tips and some cortex; white or pink medulla; white, pink, red, or purple swellings along the branch length. Autozooids are lo- autozooid apertures. Numerous conical, semi-closed, au- cated over the entire colony surface but are much more tozooid polyp apertures are uniformly/randomly distrib- abundant on the front or outer side, presumably the side uted on all branch surfaces. Siphonozooid apertures are of the colony facing the predominant current. Siphono- tightly closed and not observable to the naked eye. The zooids are not obvious to the naked eye. Anthocodiae medulla in terminal branches has six or seven major are randomly arranged throughout the colony, not obvi- canals. Surface sclerites are mostly seven- and eight-radi- ously in linear rows, only slightly raised, and the polyps ates, with long (>0.01 mm) lobulated, smooth rays. Me- are completely retractile but the apertures are variably dulla sclerites are elongated, forked or irregular spindles, closed. The colony is supported by a large (about 11 cm highly ornated, and usually less than 0.3 mm in length. in height and width), stout asymmetrical holdfast that is With regard to the Alaska specimens, two distinct white in color. The color of the colony is a uniform dark morphs conform to this species. One specimen (see pho- pink to red; the polyps are white. Holdfasts are strong tos A and F) is without doubt the same as the shallow- but the branches are rather brittle and easily broken. water (≤41 m) specimens from British Columbia, but the The other/latter morph is similar in every way to others found in deeper water are similar in morphologi- the former except colonies are sparsely and irregularly cal characteristics but quite different in form. The former branched (see photos G and H). Colonies have a stout morph is almost perfectly fan-shaped, planar but obvi- appearance and branch in a single plane and only to the ously concave, with branching to the 4th order and up to 3rd order. The color of the colony is a uniform crimson- 35 branch tips. The colony is about 48 cm in height and red in situ and when freshly collected, but dried colo- width. The colony divides into two main stems (5 cm nies are orange-red; the color of the polyps is unknown but presumed to be white. Holdfasts are strong but the branches are rather brittle and easily broken. Remarks Associates include pandalid shrimp (see photo D). Distribution Rare. In Alaska – found mostly on the conti- nental slope and upper shelf in the eastern GOA but there are a few records from the west- ern GOA and the easternmost part of the Aleutian Islands (Fig. 11-6). This species does not appear to occur in the Gulf of Alaska Seamount Province. Elsewhere – the holotype was collected in Georgia Strait, southern British Columbia, at a depth of approximately 350 m. Specimens have been collected throughout much Figure 11-6 of British Columbia and on A map of the North Pacific Ocean showing the distribution of Paragorgia stephencairnsi () Fieberling Guyot and Piggy in Alaska waters. Bank Seamount off southern California. Chapter 11 183 Habitat In Alaska – occurs singly in rough, rocky ar- men in photo B, dried. D) The same colony in photo eas (bedrock, fragmented bedrock, and cobbles) with A in situ prior to collection, with polyps retracting. E) moderate current, at depths between 171 and 741 m. The same colony in photo A in situ prior to collection, Elsewhere – occurs singly or in small thickets in rocky with polyps fully extended. F) A small thicket of P. ste- areas at depths between 32 and 490 m. phencairnsi at a depth of 40 m in Jervis Inlet, southern British Columbia. G) A fragmented but whole dried P. Photos A) A P. stephencairnsi colony (USNM stephencairnsi colony (USNM 1484201) collected in the 1484215) prior to sampling on Shutter Ridge, eastern eastern GOA at a depth of 513 m. H) Large dried frag- GOA, at a depth of 171 m. The distance between the ments of a P. stephencairnsi colony (USNM 1484203) red laser marks is 10 cm. B) A fresh branch tip collected collected in Dixon Entrance, eastern GOA, at a depth from the same colony in photo A. C) The same speci- of 427 m. 184 Professional Paper NMFS 23 4. Paragorgia stephencairnsi Sánchez, 2005 (continued) Chapter 11 185 5. Paragorgia sp. A Description Colonies are almost always planar and other, providing additional evidence that they may be fan-shaped; the largest colonies are about 50 cm in different species, subspecies, or varieties and not eco- height and 60 cm in width. Colonies appear rubbery morphs (i.e., a locally adapted population of an intra- but are actually quite fragile. Colonies are not robust, specific group having distinctive characteristics resulting but are typically highly branched, with branches that from selective pressures of the local environment). rarely exceed 1 cm in diameter. Branching starts very This species hosts hippolytid shrimps, sternostylid close to a strong holdfast that is irregularly shaped, crabs (Sternostylus iaspus), and ophiuroids including large (8 cm in diameter), white in color, and occasional- Asteronyx species. ly covered with a golden-brown stain. Nodules envelope the entire branch or are located terminally on very short Distribution Locally common. In Alaska – found branches and contain large (up to 3.6 mm in height and only in the Gulf of Alaska Seamount Province (Dickens, diameter), characteristically conical-shaped calyces that Welker, Pratt, Giacomini, and Murray Seamounts; Fig. are irregularly spaced. The anthocodiae are often white- 11-7). Elsewhere – unknown. tipped in situ and in preserved specimens; the tips of Paragorgia sp. A overlaps spatially with P. jamesi, the white (or very light pink) polyps appear to be com- P. arborea var. pacifica, P. arborea var. pacifica morph pletely retractable but often are not. The siphonozooids nodosa, and Paragorgia sp. B on seamounts in the Gulf are relatively large (approximately 0.8 mm in diameter) of Alaska Seamount Province. and abundant on all surfaces, giving the colony a rough or warty appearance. The sclerites for colonies we have Habitat In Alaska – grows mostly on bedrock and designated as Paragorgia sp. A have not yet been ex- fractured bedrock and occasionally on boulders, large amined. Colonies are a uniform, brilliant pinkish or cobbles, and hexactinellid sponge skeletons (Hetero- crimson-red color in situ and a slightly lighter hue in chone sp.). Found singly or in tight patches (up to 10 ethanol. colonies together) in generally rough habitat and at Remarks This species is known from five collected specimens and several ob- servations made in situ. Herrera and Shanks (2016) used restriction site-associ- ated DNA sequencing tech- niques to identify four of the collected specimens as P. arborea var. pacifica and one only to genus. Parago- rgia sp. A does bear a mor- phological likeness to P. arborea var. pacifica morph nodosa (that is included in the P. arborea complex of Herrera and Shanks, 2016), but there are obvious mor- phological differences that we believe warrant listing it separately until additional taxonomic work is done. Furthermore, Paragorgia sp. A and P. arborea var. paci- fica morph nodosa are not Figure 11-7 geographically or bathymet- A map of the North Pacific Ocean showing the distribution of Paragorgia sp. A () in Alaska rically discrete, even occur- waters. ring within meters of each 186 Professional Paper NMFS 23 depths between 746 and 1112 m. Observed as shallow es of a Paragorgia sp. A colony (USNM 1661895) col- as 716 m on video footage collected on Giacomini Sea- lected on Giacomini Seamount at a depth of 746 m. D) mount. A Paragorgia sp. A colony (USNM 1075751) collected on Welker Seamount at a depth of 785 m. The colony Photos A) Preserved (in ethanol) branches of a Par- is surrounded by hexactinellid skeleton (Farrea sp.) and agorgia sp. A colony (USNM 1075746) collected on hosts a sternostylid crab (Sternostylus iaspus). The dis- Welker Seamount at a depth of 781 m. B) A close-up tance between the red laser marks is 10 cm. E) A close- view of the same specimen in photo A. C) Fresh branch- up view of the same colony in photo D. Chapter 11 187 6. Paragorgia sp. B Description Colonies are almost always planar, fan- the two species could easily be mistaken for each other shaped, and convex, with an obvious front and back in situ. Fortunately, they only co-occur on Patton Sea- side; the largest colonies are about 47 cm in height and mount and have separate, non-overlapping depth ranges 58 cm in width. Colonies are often highly branched there. and appear delicate but are actually somewhat rubbery This species hosts hippolytid shrimps and ophiuroids and difficult to tear when collected. Main branches are including Asteronyx species. Sections of damaged speci- robust and often exceed 1 cm in diameter. Branching mens are often covered with hydroids, small actiniar- starts very close to a strong holdfast that is irregularly ians, and zoantharians (Zibrowius sp.). shaped, medium-sized (up to 6 cm in diameter), and white in color, occasionally covered with a golden- Distribution Uncommon. In Alaska – known only brown stain. Nodules are absent in Paragorgia sp. B; from the Gulf of Alaska Seamount Province (Welker, autozooids are directly on the branches and predomi- Pratt, Quinn, Giacomini, and Murray Seamounts; Fig. nantly on the front and lateral sides. Autozooids are 11-8). Particularly abundant on Pratt and Welker Sea- rarely present on the back side of the colony. Siphono- mounts. Elsewhere – unknown. zooids are moderately abundant and located all over Paragorgia sp. B overlaps spatially with P. jamesi, P. the branches. The short round calyces are irregularly arborea var. pacifica, P. arborea var. pacifica morph no- placed and bear large polyps that do not appear to be dosa, and Paragorgia sp. A on seamounts in the Gulf of completely retractile. The anthocodiae are often white- Alaska Seamount Province. tipped in situ and in preserved specimens; undoubtedly these are the tips of the exposed white (or light pink) Habitat In Alaska – typically found singly on bedrock polyps. Linear striations are visible in the coenenchyme but also fractured bedrock, boulders, large cobbles, and on the back side of the colony. The sclerites for colonies hexactinellid skeleton (Farrea sp.), at depths between we have designated as Paragorgia sp. B have not yet 730 and 1119 m. Observed on video footage collected been examined. Colonies are pink in situ and a slightly on Murray Seamount as deep as 1376 m. lighter hue in ethanol. The branches are a lighter shade of pink than the calyces and are lighter on the back side Photos A) Preserved (in ethanol) branches of a Para- than on the front side, even appearing white in some gorgia sp. B colony (USNM 1075759) collected on Pratt specimens. Specimens hold their natural colors when Seamount at a depth of 1067 m. B) The back side of preserved in ethanol. the same colony in photo A. C) A large preserved (in ethanol) branch complex of a Paragorgia sp. B colony Remarks This species is known from 11 collected (USNM 1075750) collected on Welker Seamount at a specimens and about 90 observations made in situ. Of depth of 783 m. D) The back side of the same colo- the 11 collected specimens, Herrera and Shanks (2016), ny in photo C. E) A Paragorgia sp. B colony (USNM using restriction site-associated DNA sequencing tech- 1075759) just prior to collection. F) A close-up view of niques, identified eight as an unknown species of Para- the same colony in photo E. A small actiniarian is at- gorgia and two as P. arborea. This species is quite dif- tached to the colony. G) The back side of a Paragorgia ferent from any other Paragorgia species known from sp. B colony (USNM 1075750) prior to collection. H) Alaska. It co-occurs with all other Paragorgia species A close-up view of a Paragorgia sp. B colony (USNM found on the same seamounts and in particular with 1075760), showing the detail of the polyps. A single zo- Paragorgia sp. A. antharian (Zibrowius cf. ammophilus) is attached to the This species is very similar in gross morphology (e.g., colony (center, top). The distance between the red laser branching pattern) and color to Hemicorallium sp., and marks in photos E and G is 10 cm. 188 Professional Paper NMFS 23 6. Paragorgia sp. B (continued) Chapter 11 189 Figure 11-8 A map of the North Pacific Ocean showing the distribution of Paragorgia sp. B () in Alaska waters. 190 Professional Paper NMFS 23 7. Sibogagorgia cauliflora Herrera et al., 2010 Description (Adapted from Herrera et al., 2010) Col- Distribution Rare. In Alaska – known from only one onies are attached to substrate with a strong, prominent collected specimen and another observed on video foot- holdfast. Branches are stout but rubbery, more so than age immediately adjacent to the collected specimen (see all Paragorgia species except maybe Paragorgia sp. B, photos C and D) on Derickson Seamount in the Gulf and emanate immediately above the holdfast. Branch- of Alaska Seamount Province (Fig. 11-9). Elsewhere – ing is irregular in a single plane. Colonies are large, up known from Davidson Seamount off northern Califor- to 150 cm in height and 130 cm in width; the Alaska nia and the Straits of Florida north of Cuba. specimen is about 50 cm in height and 52 cm in width. Main branches are only slightly thicker than clavate Habitat In Alaska – grows on large, angular boulders terminal branches. Autozooids are irregularly located at a depth of 2766 m. Found in close association with throughout the branches whereas the tiny granular si- the octocorals Parastenella gymnogaster and Narella ar- phonozooids are located uniformly on all branches. buscula (see photos C and D). Elsewhere – grows singly Sclerite morphology and ornamentation is included in on boulders at depths between 1698 and 3042 m. the original description (Herrera et al., 2010). Color is creamy white to light yellow in situ and light Photos A) Preserved (in ethanol) branches of a S. cau- yellowish-orange in freshly collected and ethanol pre- liflora colony (USNM 1081143) collected on Derickson served specimens; described as beige to red elsewhere. Seamount at a depth of 2766 m. B) A close-up view of the same specimen in photo A. C) Two S. cauliflora colo- Remarks Sibogagorgia are very similar to Paragorgia nies growing together on a large boulder on Derickson in almost all aspects except the noticeable lack of pol- Seamount at a depth of 2766 m. The white arrow points yp sclerites and the presence of boundary canals rather to the collected specimen (USNM 1081143). D) A close- than only a few larger canals separating the cortex of up view of the same specimens in photo C. The distance the branches from the medulla (Herrera et al., 2010). between the red laser marks in photos C and D is 10 cm. Chapter 11 191 Figure 11-9 A map of the North Pacific Ocean showing the distribution of Sibogagorgia cauliflora () in Alaska waters. 192 Professional Paper NMFS 23 Subfamily Anthomastinae 8. Heteropolypus japonicus (Nutting, 1912) Description (Adapted from Nutting, 1912) The holo- situated over the capitulum including the margin. Sipho- type measures 6 cm in height and the capitulum is 4.1 nozooids are densely packed between autozooids over cm in diameter. The polyps (autozooids) are relatively the entire upper surface of the capitulum (see photo C). infrequent, large, sterile and retractile, and arranged The stalk is prominent and relatively long (approxi- uniformly over the capitulum including the margin. The mately 60% as long as the capitulum diameter) with a body cavities extend directly to join the stem cavity. large attachment point. The color of the capitulum and There are a number of minute, smooth, bar-like spicules stalk in the dried state is a very pale pink to orange; in the polyps and also minute crosses, stars, and double color of the polyps is pale red. stars. All spicules are smaller than usual for the genus. Siphonozooids are fertile and densely crowded be- Remarks This taxon is known in Alaska from only tween autozooids over the entire upper surface of the two specimens (USNM 1006252 and USNM 1006300) capitulum. They appear as closely packed, rounded, or identified by F. M. Bayer in 2001. Closer examination cone-shaped verrucae with a pit in the center. The color of the specimens is warranted to determine if they are of the type specimen (USNM 30038) is dark red with indeed Heteropolypus japonicus or represent an unde- slightly darker polyps and a slightly lighter stalk. scribed species. The largest of the two known Alaska specimens The coenenchymal scales are composed principally (USNM 1006252) has a perfectly round capitulum, 4.8 (98.6%) of high-magnesium calcite (8.4 mol% MgCO3) cm in diameter, with a relatively short but stout stalk with small amounts of aragonite (senior author and S. without autozooids. The colony has a total height of D. Cairns, unpubl. data). about 8 cm. The large autozooids (about 28 total) are in various states of retraction and more or less uniformly Distribution Uncommon. In Alaska – Tanaga Pass Figure 11-10 A map of the Aleutian Islands showing the distribution of Heteropolypus japonicus () and Het- eropolypus ritteri (♦) in Alaska waters. Chapter 11 193 and Delarof Islands, central Aleutian Islands (Fig. 11- Photos A) A dorsal view of a dried whole H. japoni- 10). Elsewhere – H. japonicus is documented from cus colony (USNM 1006252) collected near the Delar- Japan, southwest of Hokkaido and south and east of of Islands, central Aleutian Islands, at a depth of about Honshu. The type specimen (USNM 30038) is from a 400 m. B) An oblique lateral view of the same speci- depth of 584 m (Molodtsova, 2013). men in photo A. C) A close-up view of the same speci- men in photo A. D) A dried whole H. japonicus colony Habitat In Alaska – likely found in scattered patches (USNM 1482137) collected in Tanaga Pass, central in areas of small cobbles, pebbles, sand, and shell at Aleutian Islands, at a depth of 424 m. This specimen is depths between about 400 and 424 m. Elsewhere – re- attached to a shell, most likely that of a large thoraci- ported at depths between 486 and 996 m. can barnacle. 194 Professional Paper NMFS 23 9. Heteropolypus ritteri (Nutting, 1909) Description (Adapted from Nutting, 1909) Colonies cies have been studied in the laboratory (Cordes et al., have a round, expanded disk-shaped holdfast, with a 2001). sterile, relatively short and stout stalk, and a somewhat flattened mushroom-shaped capitulum. The capitulum Distribution Common. In Alaska – Aleutian Islands is up to 10 cm in diameter. Autozooids are sparse and from Amukta Pass to south of Amchitka Island (Fig. irregularly positioned over the entire capitulum. Polyps 11-10). Elsewhere – along the west coast of North are completely retractile but the verrucae do not close America from British Columbia (west of the Queen over the retracted polyps, leaving a sunken pit 6 mm in Charlotte Islands) to off San Diego including several diameter. Siphonozooids are densely crowded between seamounts off California. The type specimen (USNM the autozooids over the entire upper surface of the ca- 25422) was collected off California at a depth of 801 m pitulum, giving it a granular appearance. (Molodtsova, 2013). Sclerites include abundant needle-like forms embed- ded in the surface of the capitulum, abundant stars and Habitat In Alaska – likely found singly or in scattered double stars embedded on the surface of the capitulum patches in areas of small cobbles, pebbles, and sand at between the siphonozooids, and star-like forms densely depths between 241 and 429 m. Elsewhere – reported in packed in the polyp walls. Tufts of needle-like sclerites rocky areas at depths between 55 and 1646 m. surround the autozooids. The polyps are generally a deep crimson-red with Photos A) A preserved (in ethanol) whole Hetero- lighter stalks; the color of the capitulum and stalk is polypus ritteri colony (USNM 43179) collected near the variable and typically the same color (red or pink to Queen Charlotte Islands, northern British Columbia, at creamy white). a depth of 373 m. B) A preserved (in ethanol) whole H. ritteri colony (USNM 43466) collected south of Mon- Remarks The reproductive biology, larval dynam- terey Bay, California, at a depth of 640 m. C) A close- ics including recruitment, and growth rate of this spe- up view of the same specimen in photo B. Chapter 11 195 10. Heteropolypus sp. Description (Adapted from Molodtsova, 2013) Col- of Alaska, Aleutian Islands to west of Attu Island and onies are mushroom-shaped to obconic and distinctly including Petrel Bank, the eastern Bering Sea to near separated into a capitulum and sterile stalk. Polyps are St. Lawrence Island, and Pratt, Quinn, Giacomini, and dimorphic or trimorphic and have sclerites. Autozooids Derickson Seamounts in the Gulf of Alaska Seamount are relatively scarce, large, sterile and retractile, ar- Province (Fig. 11-11). This taxon is also present on ranged evenly over the capitulum or only at the mar- Patton Seamount at depths between 393 and 3209 m gin. Anthocodial armature is often asymmetrically de- (Hoff and Stevens, 2005), and examination of archived veloped, with ridges of more-developed sclerites on the video footage (senior author, unpubl. data) confirmed dorsal side of the autozooid. the presence of the taxon on most of the seamounts that Siphonozooids are fertile, numerous, strongly ar- have been explored including Dickens (depth 746 m), mored, and densely set between the autozooids (see Welker (depths 784–1885 m), Pratt (depths 912–1935 photo B on next page). Mesozooids, when present, pos- m), Quinn (depth 2316 m), Giacomini (depths 730– sess feebly developed, retractile tentacles and are scat- 2730 m), Murray (depths 657–1851 m), Patton (depths tered between the siphonozooids. Sclerites are flanged 592–1754 m), Marchand (depth 2465 m), and Chirikof rods, radiates, clubs, rods, plates, and spindles. Pharyn- (depth 2778 m). Notably, there are no reports of this geal sclerites are predominantly platelets. Tentacular taxon from the western Gulf of Alaska or the Arctic re- sclerites are rods, clubs, spindles, multi-radiates, plates, gion. Elsewhere – Heteropolypus spp. are broadly dis- and crosses. tributed throughout the Pacific, Indian, and North At- The largest colonies observed in Alaska may be 30 lantic Oceans (Molodtsova, 2013). cm in height and 45 cm in diameter (with fully extended autozooids), possess more than 50 large autozooids, Habitat In Alaska – found singly or in scattered and form dense patches of more than 20 colonies/m2. patches in many habitat types including areas of cob- Colonies exhibit a broad range of colors in life includ- bles, pebbles, and sand and in moderate-current areas ing the full range of red and pink, orange, and creamy white. Stalks, autozooids, and polyps may be the same or different color as the ca- pitulum. Remarks This group in- cludes 72 collected speci- mens (and many observa- tions from video footage) identified only to genus and represents an ideal collec- tion of specimens to un- dertake a much needed re- examination of the genus in the North Pacific Ocean. The collection undoubtedly includes specimens of H. ja- ponicus, H. ritteri, Pseudo- anthomastus, Anthomastus, and likely one or more un- described species. Heteropolypus spp. typi- cally harbor few associated fauna. Figure 11-11 A map of the North Pacific Ocean showing the distribution of Heteropolypus sp. () in Alaska Distribution Common. waters. In Alaska – eastern Gulf 196 Professional Paper NMFS 23 of bedrock at depths between 29 and 4216 m. Re- a depth of 915 m. C) A habitat heavily populated by cently discovered growing on carbonate mounds on Heteropolypus sp. in Adak Canyon at a depth of 1734 the flanks of a mud volcano in the southeastern Gulf m. The distance between the red laser marks is 20 cm. of Alaska (see photo D; Greene10). Elsewhere – re- D) Heteropolypus sp. recently discovered growing on a ported at depths between 55 and 1646 m. Examina- carbonate mound on a mud volcano in the southeastern tion of archived video footage (senior author, unpubl. Gulf of Alaska at a depth of 1000 m. E) Two Hetero- data) confirmed the presence of the taxon on polypus sp., with polyps fully retracted, photographed Warwick Seamount off Washington at depths on Amchixtam Chaxsxii, central Aleutian Islands, at a between 871 and 1719 m. depth of 1254 m. A juvenile Verrill’s king crab (Paralo- mis verrilli) takes refuge between the two specimens. Photos A) A fresh whole Heteropolypus sp. colony F) A Heteropolypus sp. colony (J2103-4-1; CAS # (CAS 234576) collected in Adak Canyon, central Aleu- pending) collected on Amchixtam Chaxsxii at a depth tian Islands, at a depth of 1717 m. A juvenile colony is of 1339 m. G) A Heteropolypus sp. colony (USNM highlighted within the white circle. B) A close-up view 1075806) photographed on Giacomini Seamount, Gulf of a Heteropolypus sp. specimen (USNM 1116850) of Alaska Seamount Province, at a depth of 2710 m. collected in Zhemchug Canyon, eastern Bering Sea, at H) A close-up view of the same colony in photo G. I) A Heteropolypus sp. colony photographed on Amchixtam 10Greene, G. 2016. Personal commun. University of California, Chaxsxii at a depth of 559 m. The distance between the 1850 Research Park Dr., Ste. 300, Davis, CA 95618-6134. red laser marks in photos D–I is 10 cm. Chapter 11 197 198 Professional Paper NMFS 23 11. Pseudoanthomastus sp. Description (Adapted from Molodtsova, 2013) Colo- Remarks Pseudoanthomastus is a small genus con- nies are mushroom-shaped and capitate with a dome- taining only seven species, of which only one, Pseudo- shaped capitulum sharply separated from a conspicuous anthomastus fisheri (Bayer, 1952), is known to occur in stalk. Autozooids are few in number, large, and retrac- the North Pacific Ocean (in Hawaii at depths between tile (but rarely completely), and arranged evenly over 388 and 876 m). The four Alaska specimens should be the capitulum. Polyps are on very long stalks, dimor- more closely examined to determine if they are P. fish- phic, and contain sclerites. Sclerites are equally devel- eri ranging northward from the Hawaiian region or an oped around the autozooids. Anthocodial armature is unknown species. more developed near the base of the tentacles. Pseudoanthomastus elsewhere are reported to harbor Siphonozooids are fertile, usually conspicuous, feebly polynoid polychaetes (Watling et al., 2011). armored, few in number, and scattered among the large autozooids. Sclerites are principally radiates, warty Distribution Rare. In Alaska – known from only four clubs, and clubs with distal heads, and additionally a specimens collected on two seamounts (Densen and few tubercules, rods, spindles, and crosses. Pharyngeal Welker) in the Gulf of Alaska Seamount Province and sclerites are predominantly rodlets. Tentacular sclerites from video observations made elsewhere on Welker are needles, rods, clubs, and radiates. Seamount (Fig. 11-12). Elsewhere – unknown. Alaska specimens have between 5 and 10 very large autozooids; the largest specimen is approximately 5 cm Habitat In Alaska – always found in small clusters of in height and 3 cm in width at the base. The capitulum 2–3 colonies in areas of basalt and fractured basalt at and autozooids including the polyps are uniform red to depths between 1122 and 1410 m, but observed in situ dark pink in both life and in ethanol. The stalk is a uni- to a depth of 2642 m (senior author, personal observ.). form lighter shade both in life and in ethanol. Figure 11-12 A map of the North Pacific Ocean showing the distribution of Pseudoanthomastus sp. () in Alaska waters. Chapter 11 199 Photos A) Two preserved (in ethanol) whole Pseudo- colonies (USNM 1075805; lower left, inside the white anthomastus sp. colonies (USNM 1075803) collected circle) collected on Welker Seamount in the Gulf of on Densen Seamount in the Gulf of Alaska Seamount Alaska Seamount Province at a depth of 1122 m. The Province at a depth of 1410 m. The specimens are not distance between the red laser marks is 10 cm. E) A attached at the base. B) A close-up view of the same close-up view of the same colonies in photo D (center). specimens in photo A. C) The view from the top of the Note the smaller specimen to the left, inside the white same specimens in photo A. D) Pseudoanthomastus sp. circle. 200 CHAPTER 12 Family Keratoisididae Of the 13 genera in the family Keratoisididae currently referred to as bamboo corals since their axial skeleton accepted worldwide (WoRMS Editorial Board, 2022), consists of alternating white calcareous internodes and only four are known from Alaska waters: Bathygorgia, narrower dark proteinaceous nodes that resemble bam- Isidella, Keratoisis, and Orstomisis. The family Kera- boo. They are known from all ocean basins and are toisididae was known until just recently as the family quite common throughout the North Pacific and central Isididae (Saucier et al., 2021). The original description Indo-Pacific regions. They are common in all regions of of the family (Gray, 1870), amended by Verrill (1922), Alaska except the Arctic (Fig. 12-1) and are quite con- described corals with simple or variably branched axes spicuous given their often large size and unique skel- consisting of long calcareous and often hollow sec- eton. They are abundant in some regions, particularly tions that alternate with shorter horny joints consisting on seamounts in the Gulf of Alaska Seamount Province of pure gorgonin. Branches, if present, typically arise (Fig. 12-1), and provide important fisheries habitat in from the horny nodes but occasionally from the cal- some regions (Stone, 2014). They are deep-water fauna careous internodes. The coenenchyme is typically thin in Alaska, with few records shallower than 400 m, and with long conspicuous sclerites and occasionally small found at depths to 4784 m. The status of the taxonomy scale-like sclerites near the surface. Calyces are large of bamboo corals in Alaska is poor and is largely unre- and prominent, filled with large fusiform sclerites, of solved for most of the taxa we list here. Archived speci- which eight or more are larger than the rest and com- mens of all taxa are available for study with several new monly project as sharp marginal spines between the species descriptions currently in preparation (Watling11); bases of the tentacles, forming an armature for the pro- however, all bamboo coral specimens encountered, ex- tection of the tentacles that are typically not completely cept the distinctive Isidella tentaculum, should be prop- retractile. erly preserved and retained for taxonomic study. A revised description for the family, based on DNA Five morphological characteristics have traditionally sequence data (amplification of mtMutS and 18S) and been used to classify and identify bamboo corals (Bayer consequent reconstructed phylogeny, contains all four and Stefani, 1987): 1) colony branching, 2) sclerite mor- genera found in Alaska and follows from Saucier et al. phology and arrangement, 3) degree of polyp retracta- (2021). Colonies have an articulated skeleton of hol- bility, 4) structure of the axial skeleton, and 5) the pres- low or solid calcium carbonate internodes interrupted ence of an operculum on the polyps. Branching pattern, by brown to dark brown proteinaceous and sclerite-free particularly with respect to origin at the nodes (Isidella) nodes. Colonies may be unbranched (“whip-like”) or or internodes (Keratoisis), has traditionally been used with branches originating at the nodes or internodes, as an obvious characteristic to differentiate between and from either immediately distal to the nodes, or from the genera of the Keratoisididae (Bayer, 1990; France, mid-way along the internode. The coenenchyme is usu- 2007). Undoubtedly, that character has been largely ally thin but sometimes slightly thickened and covered used by experts to identify many of the museum speci- in a fleshy tegument containing nematocysts. Polyps mens from Alaska. Recent mitochondrial DNA genetics are monomorphic and non-retractile but can contract analysis has indicated that branching pattern is not a to varying degrees, never within the coenenchyme, and reliable character to distinguish among genera (France, the tentacles may only contract over the polyp oral cav- 2007; Dueñas et al., 2014). Because genetic analyses ity. Sclerites are needles, spindles, rods, or scales that and microscopic examination of sclerites (another diag- are arranged longitudinally, transversely, or obliquely nostic characteristic) have not yet been undertaken for along the polyp body and in the coenenchyme. A mes- most of the Alaska specimens, we have retained use of enterial arrangement of one or a cluster of needle-like branching pattern with other gross colony characteris- sclerites protrudes between the bases of the tentacles in tics and ecology to partition unknown specimens (and many species; other species may not possess needles but observations) generically for the purposes of this guide. a mesenterial arrangement of rod-like sclerites may still These generic listings serve as placeholders to guide fu- be present. Pharyngeal sclerites are usually present and include tuberculated or spiny rodlets, and double stars. 11Watling, L. 2021. Personal commun. School of Life Sciences, Uni- Corals in this family (keratoisidids) are commonly versity of Hawai´i at Ma |noa, Honolulu, HI 96822. Chapter 12 201 ture collections, highlight taxa for taxonomic study us- specimens (USNM 75055) with a few branches at the ing both morphological and genetics analyses, and addi- nodes were collected in the eastern Gulf of Alaska in tionally provide an updated, albeit unconfirmed, species 1942 and identified as Keratoisis paucispinosa Wright count for the region. and Studer, 1889 by E. Deichmann but unfortunately Bathygorgia is represented by two species in Alaska: the record contains no specific location or depth infor- B. profunda and Bathygorgia sp. A. Colonies are of- mation. Keratoisis paucispinosa is known from near ten unbranched or with only a few branches; the scler- Hawaii and Japan at depths between 347 and 539 m ites of the polyps and coenenchyme are bar-like forms so the identification may be a valid range extension to with turgid-rounded ends and are often bi-clavate. the north, but the original description of K. paucispi- Bathygorgia is quite similar to Keratoisis but differs in nosa (Wright and Studer, 1889) details an unbranched the complement and shape of the sclerites (Lapointe and coral so it is likely that the Alaska specimens were really Watling, 2015). Isidella is planar, often perfectly fla- Bathygorgia. bellate colonies with dichotomous branching from the Four additional species of Keratoisis have been re- nodes (Kükenthal, 1919; Bayer, 1990), and is represent- ported from the North Pacific Ocean: K. flabellum Nut- ed by three species in Alaska: Isidella tentaculum and ting, 1908 (Hawaii, 347–417 m depth), K. japonica two unknown species (Isidella sp. A and Isidella sp. B) Studer, 1878 (near Japan, 549 m depth), K. philippinen- from very deep water on Derickson Seamount. Alaska sis Wright and Studer, 1889 (Sea of Okhotsk, Russia, specimens that branch at the internodes have been des- 805 m), and K. squarrosa Kükenthal, 1915 (Kükenthal, ignated to date as Keratoisis. This might be a taxonomic 1915a) (near Japan, depth unknown). Among the seven simplification and not necessarily accurate, but without Keratoisis taxa we list here from Alaska, only Keratoi- detailed examination of sclerites and genetics analysis, sis sp. F is found within the same depth zone as any of this is the only characteristic to distinguish them from these species and should be carefully compared to those Isidella (specimens branching at the nodes). congeners. Until that time, we list the Alaska Keratoisis There are currently 22 accepted species of Keratoisis generically as distinct taxa that conform to the general worldwide (WoRMS Editorial Board, 2022); only one description of the genus but differ from each other in is reported from Alaska waters. Two large, whip-like morphology and/or ecology. Figure 12-1 A map of the North Pacific Ocean showing the distribution of corals in the family Keratoisidi- dae () in Alaska waters. 202 Professional Paper NMFS 23 1. Bathygorgia profunda Wright, 1885 Description The type specimen was reportedly only mostly discoid holdfast. The coenenchyme is relatively a fragment of the axis measuring 6 cm in length and 1 thin; the brown nodes are of variable length and can cm in diameter (3 cm in diameter including preserved often be seen through the tissue. Polyps are large, on all polyps) and consisted of three calcareous internodes sides of the branches but predominantly on two sides, and two very small horny nodes. The calcareous in- randomly arranged, and relatively sparse. ternodes were glassy, solid, smooth, bent or curved Our examination of several B. cf. profunda speci- inward, slightly quadrangular in section but more cir- mens (USNM 30076, 1081184, and 1081187) indicates cular distally. The coenenchyme was thin and densely that the polyps are somewhat squat between 3.6 and 5.0 covered with oblong, warty sclerites that were denser mm in height and 2.5 mm in diameter. The body wall near the base of the polyps. The polyps were large (5 sclerites are large (0.53–1.05 mm in length by 0.10–0.23 mm in height and 2.5 mm in diameter) and projected at mm in diameter), smooth, opaque, white, blunt-tipped right angles from the axis. Non-retractile tentacles were rods. There are also smaller (0.30–0.70 mm in length folded over each other in a somewhat irregular manner. by 0.04–0.08 mm in diameter), opaque body wall rods Large clavate, spinulate sclerites covered the body wall that may be a developing stage of the larger rods or fully of the polyp; and small, oblong sclerites densely covered grown but smaller body wall sclerites. Tentacular scler- the outer portions of the tentacles. ites are platelets (0.11–0.19 mm in length by 0.03–0.05 Subsequent descriptions by Wright and Studer (1889) mm in diameter). and Nutting (1912) indicated that the axis had long, ir- regularly curved, and somewhat square calcareous inter- Remarks Originally described as B. profunda by nodes and small horny nodal joints. The coenenchyme Wright (1885), this is the type species for the genus was thin and covered with oblong warty sclerites. The that was later synonymized with Keratoisis (Küken- polyps were large and uniserial, club-shaped (larger at thal, 1919). With the discovery of two new species of the base and top than in the middle), with non-retractile bamboo corals from seamounts off Tasmania, the ge- tentacles; and the polyp body wall was composed of nus Bathygorgia was resurrected with this species re- large, spiny, clavate and bi-clavate sclerites. Sclerites did tained as the type species (Lapointe and Watling, 2015). not protrude beyond the tentacles. Bathygorgia differs from Keratoisis in the complement The Alaska specimen (USNM 30076) is also only a and shape of the sclerites (Wright and Studer, 1889; fragment of the axis measuring 10 cm in length with Lapointe and Watling, 2015). a mean diameter of approximately 2.5 mm. It has one Both B. abyssicola Lapointe and Watling, 2015 and very small side branch with two polyps that measures B. tasmaniensis Lapointe and Watling, 2015, the only 1.3 cm in length and is very thin and delicate. There other species in the genus, are unbranched. There are is no evidence of additional branches. The polyps are no known in situ images of B. profunda and little evi- around the whole axis branch but sparsely populated dence from the two known specimens to indicate that and with prominent stalks. The color of the coenen- it is branched, aside from the small auxiliary branch on chyme in ethanol is light orange. the Alaska specimen. In addition to the type specimen and Alaska specimen Colonies on seamounts occasionally host ophiuroids (USNM 30076), nine specimens have been collected in and the chirostylid crab, Sternostylus iaspis, and at- Alaska that conform well to the above descriptions and tached skeletons are used by crinoids and brisingid sea we designate them as B. cf. profunda. Since the above stars as platforms for suspension feeding. descriptions were based on fragments of colonies, only these additional specimens along with the accompany- Distribution Apparently common (Fig. 12-2). In ing in situ observations provide detailed information Alaska – specimens have been collected in the Gulf of on colony morphology and ecology. Colonies can be Alaska Seamount Province (Giacomini, Pratt, Patton, quite large (up to 230 cm in length) and occasionally and Derickson Seamounts) and the eastern Aleutian Is- bent, twisted, and with a full spiral distally in larger in- lands (south of Davidson Bank) to the central Aleutian dividuals. Colonies are mostly whiplike or monopodial Islands (north of Atka Island) on the island arc slope but occasionally may bear as many as 10 branches that along the eastern Bering Sea. Additionally, video obser- emanate from either the nodes or internodes. Branched vations have been made on Dickens, Densen, Welker, colonies tend to be more prevalent at the shallower part Pratt, and Murray Seamounts and south of Amlia Is- of their depth range. The lower 10–25% of the stalk is land in the central Aleutian Islands (senior author, per- naked (without polyps) and colonies are supported by sonal observ.). Elsewhere – the only other known speci- a prominent but relatively small, white and reflective, men, the type specimen, was collected at Challenger Chapter 12 203 Figure 12-2 A map of the North Pacific Ocean showing the distribution of Bathygorgia profunda () in Alaska waters. station 241 in the central Pacific Ocean between Japan Atka Island in the central Aleutian Islands at a depth and Hawaii. of 3230 m. B) A close-up view of the same specimen in photo A. C) A large portion of a preserved (in ethanol) Habitat In Alaska – grows on hard substrate includ- B. cf. profunda colony (USNM 1081184) collected on ing bedrock vertical walls, fractured bedrock, pillow Derickson Seamount at a depth of 4784 m. D) A close- lavas, conglomerate, siltstone, boulders, and cobbles in up view of the same specimen in photo C. E) The same deep water (2372–4784 m). This is one of the deepest colony in photo C during collection. F) A B. cf. profun- known records for the family. Elsewhere – attaches to da colony photographed on Welker Seamount at a depth hard rock in areas of soft sediment (red clay) at a depth of 2778 m. G) A B. cf. profunda colony photographed of 4206 m. on Densen Seamount at a depth of 2488 m. H) A multi- branched B. cf. profunda colony photographed on Gia- Photos A) A fragment of a preserved (in ethanol) B. comini Seamount at a depth of 2484 m. The distance profunda colony (USNM 30076) collected north of between the red laser marks in photos F–H is 10 cm. 204 Professional Paper NMFS 23 1. Bathygorgia profunda Wright, 1885 (continued) Chapter 12 205 2. Bathygorgia sp. A Description Colonies are whip-like, infrequently ences we tentatively list this taxon as a separate species branched, never with more than four side branches until genetic analyses can be undertaken. that originate from the internodes, and up to 127 cm Colonies occasionally host pycnogonids (Colossend- in length. Whip-like colonies are often curved or spi- eis sp.). Colonies were observed with large sections of raled, especially distally, sometimes with two complete bare skeleton (up to 50% of the total colony) near the spirals and more often in a clockwise direction. The base that were undoubtedly the result of predation, like- axis is quite flexible but strong and difficult to break. ly by an unknown sea star (see photo G on next page). Internodes are variable in length (3.1–5.1 cm); the first Actiniarians often colonize those bare sections of skel- node is often immediately above the holdfast. Nodes eton. are relatively small (approximately 2–3 mm in length), somewhat recessed from the internodes, and black in Distribution Uncommon. In Alaska – known only color. Holdfasts are relatively small, cup-shaped, typi- from seamounts in the Gulf of Alaska Seamount Prov- cally without coenenchyme, and strongly attached to ince and a single location in the central Aleutian Islands the substrate. Polyps are large (8 mm in length), densely (Fig. 12-3). Specimens have been collected on Welker, populated on all branch surfaces and the axis to the Pratt, Quinn, and Giacomini Seamounts; and video holdfast, and characteristically with a bright red mouth. observations have additionally been made on Murray Tentacles are relatively short (1.5 mm) with lighter-col- Seamount and northwest of Adak Island in the central ored tips. Polyps retract very quickly when colonies are Aleutian Islands. Elsewhere – unknown. disturbed during collection. The coenenchyme is rela- tively thin; the nodes are visible in fresh specimens. The Habitat In Alaska – occurs singly or a few together color of the coenenchyme in life and in ethanol is a uni- on bedrock, fractured bedrock, boulders, and cobbles form light orange. in areas of low to moderate current, and at depths be- Our critical examination of several specimens (USNM tween 603 and 1925 m. The shallowest observations 1082131, 1082132, 1082133, and 1082146) indicates are from the Aleutian Islands. According to Lapointe that the polyps are larger (up to 5.5 mm in length) and and Watling (2015) all known Bathygorgia specimens more slender (>2.5 mm in diameter) than those in B. cf. worldwide are from depths greater than 2000 m, high- profunda. The body wall of each polyp has eight very lighting the importance of additional taxonomic work long (1.80–3.50 mm in length) and slender (0.05–0.15 with the Alaska specimens. mm in diameter) rods that protrude beyond the tenta- cles. The rods are smooth, slightly curved, and opaque. Photos A) The fresh top half of a Bathygorgia sp. The body wall sclerites proximal to the larger ones are A colony (USNM 1661888) collected on Quinn Sea- smaller (0.07–1.00 mm in length by 0.05 mm in diam- mount at a depth of 900 m. B) A close-up view of the eter). Tentacular sclerites are platelets (0.18–0.23 mm in same specimen in photo A. C) A close-up view of a pre- length by 0.02–0.05 mm in diameter). served (in ethanol) Bathygorgia sp. A specimen (USNM 1082133) collected on Pratt Seamount at a depth of Remarks This species is represented by five specimens 1873 m. D) The same specimen in photo C showing the and more than 120 video observations. Colonies oc- holdfast and lower part of the stalk that was denuded cur in the same geographical areas as B. cf. profunda when collected, likely from predation. E) A Bathygor- but occupy shallower water areas that do not appear to gia sp. A colony (USNM 1082132) collected on Welker overlap with each other. Based on our examination of Seamount at a depth of 1130 m. F) A close-up view of sclerites, this species is clearly different from B. cf. pro- the same specimen in photo E. G) A Bathygorgia sp. A funda. Additionally, the polyps are larger, more slender, colony photographed on Quinn Seamount at a depth of and situated equally on all sides of the branch axis. Col- 895 m. H) A multi-branched Bathygorgia sp. A colony onies have little or no naked section of the axis basally, (USNM 1082131) collected on Welker Seamount at a have a greater tendency to curve or spiral distally, and depth of 1132 m. The distance between the red laser have much less tendency to branch. Due to these differ- marks in photos E, G, and H is 10 cm. 206 Professional Paper NMFS 23 2. Bathygorgia sp. A (continued) Chapter 12 207 Figure 12-3 A map of the North Pacific Ocean showing the distribution of Bathygorgia sp. A () in Alaska waters. 208 Professional Paper NMFS 23 3. Isidella tentaculum Etnoyer, 2008 Description (Adapted from Etnoyer, 2008) Colonies pink in life. Colonies exude copious amounts of thick are typically planar and often perfectly formed like mucus upon collection and this mucus supports a rich candelabra when mature. The largest colonies are up bacterial fauna (Penn et al., 2006). to 150 cm in height and 150 cm in width, with up to 76 branch tips. Colonies are typically unbranched or Remarks Sweeper tentacles are unique to this species monopodial until about 18–20 cm in height and then of bamboo coral in Alaska and their presence is an eas- start to branch dichotomously at fairly regular inter- ily recognized diagnostic feature for accurate identifica- vals. Branching is from the nodes and to the 5th order tion in situ. The antipatharian Bathypathes alaskensis is in the largest colonies. The nodes are at highly irregular the only other coral found in Alaska that typically bears intervals, sometimes separated by millimeters only, es- sweeper tentacles. Sweeper tentacles are an obvious fea- pecially on the stem (see photo A) and not every node ture of the corals and provide an excellent indication produces a branching. The color of the nodes appears of bottom current direction, and to some degree, mag- to be browner than in other bamboo corals; the nodes nitude. The sweeper tentacles are considered modified are also comparatively more recessed (i.e., not level polyps rather than modified tentacles since there is no with the surface of the internodes) than those in other apparent intermediate peristome (Etnoyer, 2008). bamboo corals. The internodes are robust and solid Elongate sweeper tentacles occur in some scleractini- proximally but less so distally where they are delicate ans and other octocorals and serve an agonistic function and hollow. for space and substratum on which to grow (Sebens and When small monopodial colonies start to branch, the Miles, 1988). In some stony corals, sweeper tentacles polyps (autozooids) of the unbranched stem apparently serve as probes to detect the approach of competitors, develop into elongated basal zooids called sweeper ten- including conspecifics, and suppress encroachment by tacles; these tentacles are restricted to the stem and on those competitors by causing injury and extensive tis- the lower parts of the primary branches in some colo- sue necrosis (Lapid and Chadwick, 2006). Nematocysts nies. Sweeper tentacles are obvious, numerous (up to are present in the sweeper tentacles of Isidella tentacu- 100), densely packed, and to 40 cm or more in length. lum, implying that they too have an agonistic function Sweeper tentacles have light-colored clavate tips (acro- (Etnoyer, 2008), and our extensive in situ observations spheres) similar to the tentacles of some corallimorphar- support this premise. Colonies rarely have other seden- ian species. The holdfasts are robust, circular, medi- tary fauna growing nearby, and conspecifics are rarely um-sized (up to 5 cm in diameter), never covered with closer than 30 cm to each other. Surprisingly, this spe- coenenchyme, and strongly attached to hard substrate. cies is heavily preyed upon by at least three mobile pred- The first internode sometimes has a sharp curve along ators (see below) that are capable of denuding entire its length indicative of colonies that settled on vertical colonies including the sweeper tentacles (Stone, 2014). surfaces (e.g., sides of boulders) and then turned verti- So it is unlikely that the tentacles develop as a predator cally in the water column. defensive mechanism, per se, but we have also observed The polyps (autozooids) are large (6–9 mm in length them growing on broken branches where the colony has and 2–3 mm in diameter), tightly packed in whorls of sustained wounds. 4–5, and droopy in appearance. The tentacles are rela- The age and growth of this species have been studied tively short and stubby (2–3 mm in length), each sepa- from specimens collected on seamounts in the Gulf of rated by a projecting long (2.5 mm in length) needle-like Alaska (Andrews et al., 2009; USNM 1484154) and off sclerite. The calyces contain small cylindrical rod-shaped Washington (Roark et al., 2005). Ages can range to 208 sclerites and flattened scales with medial constriction ±42 years, with radial-axis growth rates between 0.09 that are arranged en chevron along the adaxial ridge of and 0.16 mm/year and linear growth rates of approxi- each tentacle. Small thornstar sclerites are found in the mately 1.4 cm/year. Another study (Saenger et al., 2017) pharyngeal wall and there are no sclerites in the coenen- examined the skeletal composition and biomineraliza- chyme. The polyps are capable of quick retraction when tion mechanisms of this species and demonstrated that they are disturbed in situ. the skeletons are potential paleoceanographic archives The coenenchyme is a very light golden-orange color that can be used to reconstruct temperature variations and so thin and translucent that the skeleton beneath throughout the water column over multiple centuries. is visible on all branches. The polyps, including the The skeleton (internodes) is composed almost en- mouth, are a uniform bright to light orange in life, a tirely (98.2–99.6%) of high-magnesium calcite (8.1–9.3 lighter shade in ethanol, and a darker shade when fro- mol% MgCO3) with minute amounts of aragonite and zen or dried. The sweeper tentacles are a light orange or low-magnesium calcite (senior author and S. D. Cairns, Chapter 12 209 unpubl. data). Gulf of Alaska Seamount colonies are of- This species likely ranges along the west coast of North ten used as platforms for suspension feeding by crinoids America from Alaska to southern California. (<600 m depth) and the sternostylid crab Sternostylus iaspus (>600 m depth). Other associated fauna include Habitat In Alaska – seldom found singly, more often the ophiuroid Ophiolebes brachygnatha and colonies in patches (up to 3 colonies/m2 for adults and 5 colo- occasionally host snailfish (Careproctus spp.) and the nies/m2 for juveniles) on bedrock including lava flows, golden king crab (Lithodes aequispinus). fractured bedrock, and ridges of siltstone at depths be- This species is preyed upon by the spiny red sea star tween 115 and 1608 m. Records shallower than 400 m (Hippasteria phrygiana), calliostomatid snails (Akoya are rare and the shallowest records are from the west- platinum), and nudibranchs (Tritonia sp.; Stone, 2014). ern Aleutian Islands. Elsewhere – found on hard rock at The three predators appear to operate in tandem and depths between 634 and 907 m. together can completely denude large colonies of flesh including sweeper tentacles. The snails and nudibranchs Photos A) A fresh whole I. tentaculum colony (USNM lay their egg cases on the denuded skeletons. Predation 1484153), except the holdfast, collected west of the Fair- by the spiny red sea star on stands of I. tentaculum on weather Ground, eastern Gulf of Alaska, at a depth of several Gulf of Alaska seamounts is extensive (senior 729 m. B) A fresh branch of an I. tentaculum colony author, personal observ.). (ZC-07-02-ROV-04) collected in Zhemchug Canyon in the eastern Bering Sea at a depth of 909 m. C) A large Distribution This species is one of the most wide- I. tentaculum colony photographed near Tanaga Island, spread octocorals in the North Pacific Ocean (Fig. 12- central Aleutian Islands, at a depth of 1325 m. D) A 4) and is locally abundant. In Alaska – specimens have large I. tentaculum colony photographed on Dickens been collected in the Gulf of Alaska (Dixon Entrance Seamount at a depth of 758 m. At least six sternostylid to southwest of Albatross Bank), the Gulf of Alaska crab (Sternostylus iaspus) are perched in the colony. The Seamount Province (Dickens, Welker, Pratt, Giacomi- distance between the red laser marks in photos C and ni, and Patton Seamounts), the central Aleutian Islands D is 20 cm. E) A close-up view of the same specimen in (southwest of Amlia Island to Stalemate Bank), and the photo D with the polyps fully extended. F) A close-up eastern Bering Sea (south of Pribilof Canyon to Nava- view of polyps contracting in an I. tentaculum colony rin Canyon). Colonies have also been observed on video (USNM 1076664) sampled on Pratt Seamount at a depth footage collected on Dickens (743–857 m depth), Welk- of 913 m. G) A close-up view of the sweeper tentacles er (773–1071 m depth), Pratt (911–1062 m depth), adorning the stem of an I. tentaculum colony (USNM Giacomini (715–890 m depth), Murray (672–984 m 1082136) sampled on Pratt Seamount at a depth of 1049 depth), and Patton (286–993 m depth) Seamounts. Else- m. H) A close-up view of the sweeper tentacles on an I. where – specimens have been collected from Warwick tentaculum colony (USNM 1082145) sampled on Gia- Seamount (Washington), Monterey Bay (northern Cali- comini Seamount at a depth of 727 m. The distance be- fornia), and Rodriguez Seamount (southern California). tween the red laser marks is 10 cm. 210 Professional Paper NMFS 23 3. Isidella tentaculum Etnoyer, 2008 (continued) Chapter 12 211 Figure 12-4 A map of the North Pacific Ocean showing the distribution of Isidella tentaculum () in Alaska waters. 212 Professional Paper NMFS 23 4. Isidella sp. A Description Colonies are flabellate and mostly pla- tentaculum is the only named species known to occur nar with irregular branching starting on or immediately in the northern North Pacific Ocean. Isidella sp. A is above the holdfast. The largest colonies measure about very different morphologically and ecologically from I. 35 cm in height and 74 cm in width, with up to 75 tentaculum and undoubtedly represents an undescribed branch tips (see photo E). Branching is from the nodes species. It co-occurs with Isidella sp. B (described on and typically to the 3rd and 4th order in the largest colo- page 214) and conspicuously differs from it. nies. Branches are quite fragile and easily broken dur- Some colonies host crinoid associates and small ac- ing collection. Polyps mostly alternate in one plane and tiniarians attach to bare sections of the skeleton. are long and slender, about three times longer than the width of the branch from which they emanate. Polyps Distribution Uncommon. In Alaska – known only are sparse and moderately spaced. The branches are from Derickson Seamount in the Gulf of Alaska Sea- hollow except for near the base, and the coenenchyme mount Province (Fig. 12-5). Elsewhere – unknown. is moderately thick such that the brown nodes are barely visible beneath. Holdfasts are relatively small, circular, and notably discoid and strongly attached to Habitat In Alaska – occurs singly (rarely more than the substrate. The sclerites have not yet been critically one) on bedrock and fractured bedrock, including boul- examined for this taxon. The coenenchyme in life is ders and cobbles. Co-occurs with Bathygorgia profun- a pinkish orange and a yellowish orange preserved in da, Isidella sp. B, Keratoisis sp. E, and Narella bayeri ethanol. at the shallow end of its depth range. Specimens have been collected at depths between 3993 and 4488 m but Remarks There are six accepted species of Isidella colonies have been observed as shallow as 3839 m and worldwide (WoRMS Editorial Board, 2022); Isidella as deep as 4643 m (senior author, personal observ.). Figure 12-5 A map of the Aleutian Islands showing the distribution of Isidella sp. A () and Isidella sp. B () in Alaska waters. The two taxa have nearly coincident distributions. Chapter 12 213 Photos A) A large branch complex of a preserved (in D) A close-up view of the same colony in photo C. The ethanol) Isidella sp. A colony (USNM 1081182) col- distance between the red laser marks in photos C and D lected on Derickson Seamount at a depth of 4471 m. B) is 10 cm. E) An Isidella sp. A colony (USNM 1081183) A close-up view of the same specimen in photo A show- collected on Derickson Seamount at a depth of 4488 m. ing the polyp structure and arrangement. C) The same The distance between the red laser marks is 20 cm. colony in photo A photographed just before collection. 214 Professional Paper NMFS 23 5. Isidella sp. B Description Colonies have a somewhat scraggly ap- Habitat In Alaska – typically found in patches, some- pearance. Largest colonies are up to 38 cm in height, times moderately dense patches (up to 8 colonies/m2) with 25 branch tips starting on the holdfast or imme- on flat bedrock and fractured bedrock with Bathygorgia diately above it; they are mostly branched only to the profunda, Isidella sp. A, Keratoisis sp. E, Chrysogorgia 2nd order and rarely to the 3rd. Colonies have very thin sp. B, and Pseudochrysogorgia sp. C, and at depths be- coenenchyme and are extremely fragile. Polyps appear tween 3909 and 4643 m. to be slightly alternate and the short stem, if present, has polyps. Lower branches on the stem are almost op- Photos A) An Isidella sp. B colony (USNM 1081179; posite. The sclerites have not yet been critically exam- indicated by the white circle) just before collection on ined for this taxon. Prominent, circular holdfasts are Derickson Seamount at a depth of 4162 m. The colony strongly attached and without coenenchyme. is in a field of congeners, B. profunda, and chrysogor- gids. B) An Isidella sp. B colony photographed on De- Remarks There are six accepted species of Isidella rickson Seamount at a depth of 4527 m. C) A close- worldwide (WoRMS Editorial Board, 2022); Isidella up view of the same colony in photo B. The distance tentaculum is the only species known to occur from the between the red laser marks in photos A–C is 10 cm. northern North Pacific Ocean. Isidella sp. B co-occurs D) A patch of Isidella sp. B colonies on a large boulder with Isidella sp. A and conspicuously differs from it. on Derickson Seamount at a depth of 4025 m. The dis- tance between the red laser marks is 20 cm. E) A field Distribution Uncommon but locally abundant. In of Isidella sp. B colonies mixed with B. profunda on Alaska – known only from Derickson Seamount in the Derickson Seamount at a depth of 4163 m. Gulf of Alaska Seamount Province (Fig. 12-5). Else- where – unknown. Chapter 12 215 216 Professional Paper NMFS 23 6. Keratoisis sp. A Description This taxon was determined to be a dif- Distribution Uncommon but locally abundant. In ferent species than Keratoisis sp. B and Keratoisis sp. Alaska – known only from a series of volcanic cones C based on the complement and sizes of sclerites and and lava flows in Tanaga Pass in the central Aleutian analysis of mitochondrial msh1 DNA sequences and is Islands (Fig. 12-6). Elsewhere – unknown. currently queued for formal description (Watling12 and France13). Colonies are polymorphic, ranging from pla- Habitat In Alaska – colonies are found on a series of nar and candelabra-like to slightly bushy or even scrag- heavily eroded pinnacles and crags forming a ridge at gly. The largest collected specimen has dimensions 64 depths between 1584 and 2035 m. Colonies are restrict- cm in height, 55 cm in width, with a stem measuring 9 ed to the volcanic ridge and were not observed on boul- cm in length, an irregular-shaped holdfast measuring 9 ders scattered in the soft-sediment habitat surrounding mm in diameter, and 39 branch tips. The smallest speci- the volcanic features. Colonies occur in patches up to men is 28 cm in height with 8 branch tips. The largest 8 colonies/m2 in areas of moderate current and are as- colonies observed in situ measured 92 cm in height, 82 sociated with hexactinellid sponges (Caulophacus adak- cm in width, with more than 60 branches. Branching is ensis and Farrea aspondyla) and the demosponge Axo- from the internodes, typically only to the 2nd order but niderma corona in the deeper part of its range. At the occasionally to the 3rd order in larger individuals. Pri- shallower part of its range, it co-occurs with Plumarella mary branches start at or on the holdfast or on a very echinata Cairns, 2011 and briefly overlaps with Isidella short stem. Branches are quite flexible but fragile and tentaculum where that species becomes the dominant very easy to break when sampling. The internodes are octocoral. relatively close together (about every 4 cm); the nodes are relatively short and black in color. Photos A) Fresh branches of a Keratoisis sp. A col- Polyps are principally aligned in two lateral rows on ony (USNM 1659769) collected northwest of Tanaga either side of the branch but are scattered all around. Island, central Aleutian Islands, at a depth of 2005 m. Colonies have large, robust, irregularly shaped holdfasts B) Fresh large fragments of a Keratoisis sp. A colony that are white, without coenenchyme, and strongly at- (USNM 1484161) collected northwest of Tanaga Is- tached. The skeletal axis can easily be seen beneath a land at a depth of 2031 m. C) A close-up view of the relatively thin coenenchyme. The sclerites have not yet polyps and branching pattern of the same specimen in been critically examined for this taxon. The color of the photo B, preserved in ethanol. D) The same colony in coenenchyme in life and in ethanol is a very light orange photo B just before collection. The distance between or translucent; the color of the polyps is a darker shade the red laser marks is 20 cm. E) A Keratoisis sp. A col- of orange. ony (USNM 1659769) photographed just before col- lection. F) A Keratoisis sp. A colony photographed in Remarks The skeleton (internodes) is composed al- Tanaga Pass at a depth of 1719 m. G) A close-up view most entirely (98.3–99.3%) of high-magnesium calcite of extended polyps in the same colony in photo F. The (7.4–8.9 mol% MgCO3) with minute amounts of ara- distance between the red laser marks in photos F and gonite and low-magnesium calcite (senior author and S. G is 10 cm. H) A large Keratoisis sp. A colony photo- D. Cairns, unpubl. data). graphed in Tanaga Pass at 1618 m. The distance be- tween the red laser marks is 20 cm. 12Watling, L. 2021. Personal commun. School of Life Sciences, Uni- versity of Hawai´i at Ma-noa, Honolulu, HI 96822. 13France, S. 2021. Personal commun. University of Louisiana at Lafayette, Lafayette, LA 70504. Chapter 12 217 218 Professional Paper NMFS 23 6. Keratoisis sp. A (continued) Figure 12-6 A map of the Aleutian Islands showing the distribution of Keratoisis sp. A (), Keratoisis sp. B (♦), and Keratoisis sp. C (▲) in Alaska waters. Chapter 12 219 7. Keratoisis sp. B Description This taxon was determined to be a dif- Adak Canyon, central Aleutian Islands. Of the three lo- ferent species than Keratoisis sp. A and Keratoisis sp. cations in Adak Canyon that have been explored within C based on the complement and sizes of sclerites and the known depth range of this species, it was common analysis of mitochondrial msh1 DNA sequences and is at one site; four specimens were observed at the second currently queued for formal description (Watling12 and site; and it was not observed at the third site. These ob- France13). Colonies are straggly but appear to be planar servations indicate that this species has a very limited in broad view, with irregular branching that starts close geographical range. Elsewhere – unknown. to the holdfast; there are up to 22 branch tips to the 3rd order. Maximum size is 35 cm in height and 35 cm Habitat In Alaska – occurs in relatively rough habitat in width. Colonies are somewhat flexible and not par- with moderate currents and grows on fractured bed- ticularly fragile. Branching occurs along the internodes; rock, boulder, conglomerate, and siltstone at depths the nodes are very short and dark black in color. Pol- between 1716 and 2044 m. It occurs in dense patches yps are quite crowded, principally on three sides of the (up to 7 colonies/m2) with the congener Keratoisis sp. branches, and are slightly darker in color than the rela- C, Acanthogorgia spissa, and an unidentified species of tively thin coenenchyme that reveals the axial skeleton Heteropolypus. underneath. The slightly conical polyps are small (6 mm in height by 3–4 mm in width) and appear to be mostly Photos A) A fresh large portion of a Keratoisis sp. B retractable in fresh specimens. The sclerites have not yet colony (USNM 1484160) collected in Adak Canyon in been critically examined for this taxon. The holdfast is the central Aleutian Islands at a depth of 1715 m. B) relatively small but robust, irregularly cup-shaped, and The same colony in photo A in situ with polyps fully white and reflective in situ. Color of the coenenchyme is extended. A partially skeletonized colony (at left) shows light to medium orange in life. the branching pattern with nodes and internodes. The distance between the red laser marks is 10 cm. C) A Remarks The skeleton (internodes) is composed al- close-up view of the same colony in photo A during most entirely (96.5–99.3%) of high-magnesium calcite collection. D) A fresh whole Keratoisis sp. B colony (7.6–9.4 mol% MgCO3) with minute amounts of ara- (USNM 1659765) collected in Adak Canyon at a depth gonite and low-magnesium calcite (senior author and S. of 1717 m. E) A close-up view of the same colony in D. Cairns, unpubl. data). photo D with polyps in various states of contraction. F) The same colony in photo D in situ (at center) just Distribution Uncommon but locally abundant (Fig. before collection. The distance between the red laser 12-6). In Alaska – known only from two locations in marks is 20 cm. 220 Professional Paper NMFS 23 7. Keratoisis sp. B (continued) Chapter 12 221 8. Keratoisis sp. C Description This taxon was determined to be a dif- mol% MgCO3) with minute amounts of aragonite and ferent species than Keratoisis sp. A and Keratoisis sp. low-magnesium calcite (senior author and S. D. Cairns, B based on the complement and sizes of sclerites and unpubl. data). analysis of mitochondrial msh1 DNA sequences and is currently queued for formal description (Watling12 and Distribution Uncommon. In Alaska – known only France13). It co-occurs with Keratoisis sp. B but appar- from a single location in Adak Canyon, central Aleutian ently differs in being more robust, having fewer branch- Islands (Fig. 12-6). It occurs at that site with Keratoisis es and more polyps that are spiraled on all sides of the sp. B but appears to be much less common than the for- branches, and having a darker orange color (see photo mer species in patches where they co-occur. Elsewhere B on next page). Colonies are mostly planar, with ir- – unknown. regular branching that starts close to the holdfast; they have up to 15 branch tips to the 3rd order. Maximum Habitat In Alaska – occurs in relatively rough habitat size is 42 cm in height and 34 cm in width. Colonies with moderate currents and grows on fractured bed- are somewhat flexible and not particularly fragile. rock, conglomerate, and siltstone at depths between Branching occurs along the internodes; the nodes are 1715 and 1988 m. It occurs in dense patches (up to 7 short and dark black in color. Polyps are quite crowded colonies/m2) with the congener Keratoisis sp. B, Acan- on all sides of the branches and are slightly darker in thogorgia spissa, and an unidentified species of Hetero- color than the relatively thin coenenchyme that reveals polypus. the axial skeleton only along the stem. The cylindrical polyps are small (7 mm in height by 4 mm in width) Photos A) A fresh whole (except the holdfast) Kera- and appear to be mostly retractable in fresh specimens. toisis sp. C colony (USNM 1659767) collected in Adak The sclerites have not yet been critically examined for Canyon, central Aleutian Islands, at a depth of 1715 m. this taxon. The holdfast is moderately large and robust, B) A close-up of the same specimen in photo A. C) The more or less cup-shaped or discoid, and white and re- same colony in photo A prior to collection. A Keratoisis flective in situ (see photo D on next page). This species sp. B colony is at the lower left. D) A close-up view of secretes copious amounts of mucous upon collection. the same colony in photo C prior to collection showing Color of the coenenchyme is medium to dark orange the highly reflective holdfast. E) An alternate close-up in life. view of the same colony in photo D showing the polyps. The distance between the red laser marks in photos C–E Remarks The skeleton (internodes) is composed al- is 10 cm. most entirely (97.0%) of high-magnesium calcite (8.9 222 Professional Paper NMFS 23 8. Keratoisis sp. C (continued) Chapter 12 223 9. Keratoisis sp. D Description Colonies are bushy with a straggly ap- are visible along most of the branches basally. The color pearance; branching typically starts from a very short of the coenenchyme is light orange to cream in life and stem; branching is to the 3rd order only. The largest light orange in ethanol-preserved specimens. colonies are to about 30 cm in height and width with no more than 20 branch tips. Branching is at the in- Remarks Associated fauna include sternostylid (Ster- ternodes; the nodes are notably narrow and occur at nostylus iaspus) and munidopsid crabs (Munidopsis semi-irregular intervals (about 16–21 mm apart). Hold- sp.); ophiuroids and actiniarians occasionally attach to fasts are small (up to 13 mm in diameter) and circular, bare sections of the skeleton. This taxon overlaps with typically without coenenchyme, but may be robust and the primnoid Calyptrophora laevispinosa in the deeper heavily calcified. Colonies are fragile and easily broken part of its depth range and has similar morphology with during sampling. juvenile colonies of the latter species. Its fragility sets Polyps appear to be alternately or randomly situated it apart, however, as C. laevispinosa is very difficult to on all sides of the branches, giving the appearance of be- tear upon collection. ing spiraled. Polyps (expanded) are long and slender (up to 10.0 mm in length by 3.0 mm in width), taper slightly Distribution Uncommon. In Alaska – known only distally, have a red-colored mouth, and are thicker than from seamounts in the Gulf of Alaska Seamount Prov- the slender branches that bear them. Tentacles are blunt- ince (Fig. 12-7). Specimens have been collected from tipped, about 1.4 mm in length, and bear 8–9 pairs of Dickens, Densen, Welker, and Pratt Seamounts and pinnules. video observations have additionally been collected on The sclerites have not yet been critically examined for Giacomini, Murray, and Patton Seamounts. It is par- this taxon. The coenenchyme is variably thick but often ticularly common on Murray Seamount. Elsewhere – translucent enough that the narrow, dark brown nodes unknown. Figure 12-7 A map of the North Pacific Ocean showing the distribution of Keratoisis sp. D (), Keratoisis sp. E (♦), and Keratoisis sp. F (▲) in Alaska waters. 224 Professional Paper NMFS 23 Habitat In Alaska – these corals typically occur sin- just before collection. D) A close-up view of the same gly, occasionally 2–4 colonies together, on bedrock, colony in photo A. E) A Keratoisis sp. D colony (USNM fractured bedrock, boulders and cobbles, and occasion- 1076659) photographed just before collection on Welk- ally at the base of hexactinellid sponges, at depths be- er Seamount at a depth of 1096 m. An actiniarian is tween 598 and 2689 m. attached to a bare section of the skeleton and a muni- dopsid crab (Munidopsis sp.) uses the colony as refuge. Photos A) A large portion of a preserved (in etha- F) A Keratoisis sp. D colony photographed on Giaco- nol) Keratoisis sp. D colony (USNM 1076661) col- mini Seamount at a depth of 2477 m. Note the juvenile lected on Pratt Seamount at a depth of 1210 m. B) A colony in the background (at far right). The distance close-up view of the same specimen in photo A showing between the red laser marks in photos C, E, and F is 10 details of the polyps. C) The same colony in photo A cm. Chapter 12 225 10. Keratoisis sp. E Description The single collected specimen (USNM Distribution Locally abundant. In Alaska – known 1081149) is bushy but somewhat fan-shaped and rela- only from Derickson Seamount in the Gulf of Alaska tively planar. It measures 34 cm in height and 44 cm Seamount Province (Fig. 12-7). Elsewhere – unknown. in width with 16 branch tips; larger colonies have been observed in situ that have between 20 and 25 branch Habitat In Alaska – almost always occurs singly but tips. Branching is from the internodes and starts at the occasionally in patches of 5 colonies/m2 in cobble field holdfast or from a very short stem; branching is typi- habitat (fractured bedrock) with Isidella sp. A, I sidella cally only to the 2nd order and rarely to the 3rd order. sp. B, and a few Bathygorgia cf. profunda. Found at Branches appear rubbery and are more robust than depths between 3990 and 4373 m and is more common those in Isidella sp. A and Isidella sp. B, with which in the shallower part of its depth range. it co-occurs, but they are easily broken upon collec- tion. The polyps are tightly packed on all sides of the Photos A) A Keratoisis sp. E colony (USNM branches. The holdfast is small, discoid, firmly attached 1081149; indicated by the white circle) photographed to the substrate, and devoid of coenenchyme. The coe- just before collection on Derickson Seamount at a depth nenchyme is thick except near the holdfast, and is light of 4098 m. The colony is in a field dominated by Isidel- orange to light pink in color. The sclerites have not yet la sp. B. B) A close-up view of the same colony in photo been critically examined for this taxon. A. C) A Keratoisis sp. E colony (indicated by the white circle) photographed on Derickson Seamount at a depth Remarks This taxon is known from only one collect- of 4115 m. The distance between the red laser marks in ed specimen and many video observations made nearby. photos A–C is 20 cm. There are no other Keratoisis known from these bathyal depths in Alaska waters. 226 Professional Paper NMFS 23 11. Keratoisis sp. F Description Colonies are large, generally bushy or marine ecosystems including coral gardens. This species straggly, and even scraggly as they are often damaged co-occurs in the central Aleutian Islands with Primnoa or preyed upon and exhibit large areas of bare or bro- wingi Cairns and Bayer, 2005 and the two species are ken skeleton. The largest colonies are to at least 120 easily mistaken for each other in situ. Primnoa wingi cm in height and 100 cm in width, with more than 100 has a very stout holdfast and stem that are completely branch tips. The main axis and lower branches are ro- covered with coenenchyme whereas Keratoisis sp. F bust and solid; distal branches are delicate, fragile, and has a massive, irregular holdfast that is almost always hollow. Branching is highly irregular and from the in- bare skeleton. Colonies often present evidence of pre- ternodes that provide for twisting branches that occa- dation, likely from the spiny red sea star (Hippasteria sionally anastomose (see photo B). The black nodes are phrygiana) that has been observed preying on nearby extremely short (more so than any other taxon listed Isidella tentaculum (see photo G), and bare skeletons of here), especially basally, but gradually increasing in Aleutian Island colonies are occasionally covered with length distally. Primary branches emanate from the Clavularia sp. A. holdfast or immediately above it. The axial skeleton has obvious linear striations. The holdfast is large, irregular, Distribution Uncommon but locally abundant. In devoid of coenenchyme, and strongly attached to the Alaska – this taxon is known from the continental slope substrate (i.e., the primary branch will typically break of the eastern GOA and the island arc slope of the cen- near the base before the holdfast). tral Aleutian Islands (Fig. 12-7). Elsewhere – unknown. The polyps are large, tightly packed all around the Habitat In Alaska – occurs on hard substrate includ- branches, not completely retractile, and with obvious ing bedrock, fractured bedrock, boulders, and large fusiform, sharp marginal spines projecting at the base cobbles at depths between 574 and 1563 m. It occurs of the tentacles. The coenenchyme is generally thick in low density patches with I. tentaculum in the eastern but the holdfast and lower branches are often devoid of GOA and is found in coral garden habitat in northern flesh. The sclerites have not yet been critically examined Amchitka Pass in the central Aleutian Islands. for this taxon. The coenenchyme including the polyps is a uniform bright to medium orange in life and a lighter Photos A) A dried large medial branch complex of orange shade when dried. Keratoisis sp. F (USNM 1484159) collected in the east- ern GOA at a depth of 760 m. B) A dried and varnished Remarks This is a very large and conspicuous bam- Keratoisis sp. F specimen (USNM 1484157) collected boo coral that somehow largely escaped notice until re- in the eastern GOA at a depth of 697 m. The photo cently, despite being locally common within the depth shows the colony base (just above the holdfast, at cen- range of historical fishing activities and traditional re- ter) and several branch sections. C) Dried branch tips search surveys. of Keratoisis sp. F (USNM 1484158) collected in the The age and growth of this species have been stud- eastern GOA at a depth of 578 m. D) A large Kera- ied from a specimen (USNM 1485149) collected in the toisis sp. F colony photographed in northern Amchitka eastern Gulf of Alaska with age estimated at 116 ± 29 Pass in the central Aleutian Islands at a depth of 849 m. years, a radial-axis growth rate of approximately 0.056 The distance between the red laser marks is 20 cm. E) A mm/year, and a linear growth rate of approximately 10 close-up view of the same colony in photo D. The dis- mm/year (Andrews et al., 2009). Another study (Saenger tance between the red laser marks is 10 cm. F) A cluster et al., 2017) examined the skeletal composition and of three Keratoisis sp. F colonies (top) photographed biomineralization mechanisms of this species and dem- northwest of Adak Island in the central Aleutian Islands onstrated that the skeletons are potential paleoceano- at a depth of 725 m. An I. tentaculum colony is in the graphic archives that can be used to reconstruct tem- foreground. The distance between the red laser marks perature variations throughout the water column over is 50 cm. G) A Keratoisis sp. F colony with evidence of multiple centuries. predation photographed northwest of Adak Island at a Keratoisis sp. F is typically found in complex habi- depth of 1462 m. The distance between the red laser tats and is an excellent indicator species of vulnerable marks is 20 cm. Chapter 12 227 228 Professional Paper NMFS 23 12. Keratoisis sp. G Description Colonies are bushy but somewhat fan- northern California (see Figure 1B in Andrews et al., shaped and relatively planar with irregular branching. 2009). The largest colonies measure 96 cm in height and 77 cm in width, with almost 100 branch tips. Branching in Habitat In Alaska – grows on basalt bedrock (lava the largest specimens is to the 4th order but in smaller flows) and fractured bedrock (boulders and cobbles) at specimens typically only to the 2nd order. Branching is depths between 1689 and 1726 m. Colonies occur sin- from the internodes and starts at the holdfast or from gly, spaced out by 5 m or more, and occasionally grow a very short stem. Branches are quite flexible but are at the bases of hexactinellid sponges. Elsewhere – on easily broken upon collection, and larger colonies often fractured bedrock at 1400–1455 m depth. have broken and bare sections of skeleton that likely result from unobserved predation. Nodes are relatively Photos A) Branches (preserved in ethanol) of Kera- short and dark brown in color. The polyps are tightly toisis sp. G (USNM 1076666) sampled on Giacomini packed on all sides of the branches, apparently in four Seamount at a depth of 1726 m. B) A close-up view of rows. The short, stout, and conical polyps bear short the same specimen in photo A showing branching at the and stubby tentacles. The sclerites have not yet been internodes. C) A zoomed-in view of the same specimen critically examined for this taxon. The holdfast is mod- in photo A. D) The same colony in photo A growing at erately large and robust, irregularly shaped, firmly at- the base of a hexactinellid sponge (Chonelasma oreia) tached to the substrate, and devoid of coenenchyme as photographed just before collection. E) A zoomed-in are many of the lower branches. The coenenchyme is view of the same colony in photo D. F) A Keratoisis sp. thick where present, and is orange to light pink in life G colony (USNM 1082147) sampled on Giacomini Sea- and a yellowish-orange in ethanol. mount at a depth of 1689 m. G) A Keratoisis sp. G col- ony photographed on Giacomini Seamount at a depth Remarks This species is known from two collected of 1692 m. The spider crab Macroregonia macrochira specimens and multiple video observations on the southeast flank of Giaco- mini Seamount. It is a dif- ferent morphotype than all other Keratoisis listed in this guide. Growth rate and age were studied for a morpho- type collected on Davidson Seamount off northern Cali- fornia at a depth of 1455 m, with age estimated at >145 years and a moderate linear growth rate of 14–28 mm/ year (Andrews et al., 2009). The spider crab Macrorego- nia macrochira occasionally uses colonies as refuge. Distribution Uncommon. In Alaska – known only from Giacomini Seamount in the Gulf of Alaska Sea- mount Province (Fig. 12-8). Elsewhere – unknown but a Figure 12-8 morphotype and video ob- A map of the North Pacific Ocean showing the distribution of Keratoisis sp. G () in Alaska servations were collected waters. on Davidson Seamount off Chapter 12 229 uses the colony as refuge. The distance between the view of a Keratoisis sp. G colony photographed on Gia- red laser marks in photos D–G is 20 cm. H) A close-up comini Seamount at a depth of 1690 m. 230 Professional Paper NMFS 23 13. Orstomisis sp. Description The single collected specimen consists of recently collected from the Mid-Karin Ridge near Ha- five branches that emanate from two main stalks off a waii and was generically identified by S. D. Cairns as very short stem above the holdfast; the longest branch Orstomisis. is about 18 cm in length and 3.3 mm in average width. The skeleton (internodes) is composed almost entirely Branching is apparently from the barely visible nodes; (98.2%) of high-magnesium calcite (8.0 mol% MgCO3) in this genus the nodes are typically overgrown by the with minute amounts of aragonite and low-magnesium calcareous internodes especially near the base (Bayer, calcite (senior author and S. D. Cairns, unpubl. data). 1990). The branches are fairly flexible and a good in- dicator of water current direction. Polyps are sparsely Distribution Rare. In Alaska – known only from a populated, principally on two sides of the branches, and small area south of Amlia Island, central Aleutian Is- are relatively long and narrow when fully expanded, lands (Fig. 12-9). Elsewhere – unknown. but short and squat when retracted. The polyps form Habitat In Alaska – occurs singly or in small patches short, cylindrical calyces into which the tentacles can be (up to 3 colonies/m2) on siltstone, bedrock, and frac- fully retracted. The coenenchyme is thin and the hold- tured bedrock (cobble) at depths between 2674 and fast and bottom branches are typically without coenen- 2826 m. Co-occurs with the the antipatharian Trisso- chyme. Colonies have prominent white, cup-shaped, but pathes pseudotristicha, and the octocorals Radicipes often irregular, holdfasts that are reflective in situ. The stonei and Plumarella aleutiana Cairns, 2011. Else- coenenchyme is medium to light orange in color. where – the Hawaiian specimen grew singly on basalt In addition to the collected specimen, we observed ap- table rock at a depth of 2255 m. proximately 50 colonies in the immediate area with mor- phology that conformed to this same species. Colonies Photos A) A fresh whole Orstomisis sp. colony were mostly planar, with up to 45 branches emanating (USNM 1484147/1659770) collected south of Amlia Is- mostly from near the base, and were rarely branched to land, central Aleutian Islands, at a depth of 2826 m. B) the 3rd order. The largest colonies are to 150 cm in width A close-up view of the same specimen in photo A show- and at least 70 cm in height. The largest colonies have ing the arrangement of the polyps. C) The same colony massive holdfasts (up to 13 cm in diameter) that were in photo A just before collection. D) An Orstomisis sp. covered with coenenchyme and polyps (see photo G). colony photographed on the same submarine ridge at a depth of 2826 m. E) A close-up view of the same speci- Remarks This genus was erected to accommodate men in photo D. F) Two large Orstomisis sp. colonies a new species from New Caledonia, Orstomisis cros- photographed on the same submarine ridge as the speci- nieri Bayer, 1990, found at depths between 542 and mens in photos C and E. G) An Orstomisis sp. colony 600 m. The Alaska specimen is currently queued for photographed on the same submarine ridge as the speci- formal species description (Watling12 and France13). mens in photos C, E, and F. The distance between the The only other specimen assigned to the genus was red laser marks in photos C, D, F, and G is 20 cm. Chapter 12 231 232 Professional Paper NMFS 23 13. Orstomisis sp. (continued) Figure 12-9 A map of the Aleutian Islands showing the distribution of Orstomisis sp. () in Alaska waters. 233 CHAPTER 13 Family Primnoidae Corals in the family Primnoidae are octocorals with a skel- of Alaska Seamount Province. One species ranges fur- etal axis of solid, non-scleritic material and continuous ther to the east (Asia), three species range further to the calcium carbonate (Fig. 13-1). A cross-section of the axis south (including the variant), and two species range to shows undulating concentric layers of calcified material both the east and south. variably embedded in gorgonin (Fig. 13-1). Colonies are Primnoids are found in all regions of Alaska except erect, typically branched, firmly attached to hard substrate the Arctic; none are found further north than Zhemchug with a robust calcareous holdfast, and sometimes referred Canyon in the eastern Bering Sea (Fig. 13-2). The records to as gorgonians or sea fans. Polyps are monomorphic for most taxa represent the northernmost in the world. and non-retractile with calyces occurring in many forms They are eurybathic fauna in Alaska occupying depths and arrangements (McFadden et al., 2022). between 6 and 4633 m. All taxa are firmly attached or Primnoids that occur in Alaska waters are colonial fixed and typically grow on bedrock, boulders, cobbles, and ahermatypic. Some species attain very large size, are pebbles, and occasionally on siltstone. Plumarella and locally abundant, and play a major role in construct- Callogorgia spp. have been observed attached to hexac- ing important benthic habitats (Stone, 2006, 2014). tinellid sponge skeleton. Twenty-six species (plus one variant) of primnoids are The current state of taxonomy for the group is ex- known from Alaska waters. Twelve species appear to cellent and there are no taxa with unresolved taxono- be endemic to the Aleutian Islands and four species (all my. Nonetheless, specimens collected in Alaska should Narella) appear to be endemic to seamounts in the Gulf be properly preserved for taxonomic examination and Figure 13-1 A cross-sectional view of a very large Primnoa pacifica specimen (two colonies grown together) collected in Dixon Entrance in the eastern Gulf of Alaska showing the heavily calcified solid axis composed of calcite and aragonite that is characteristic of the family Primnoidae. 234 Professional Paper NMFS 23 Figure 13-2 A map of the North Pacific Ocean showing the distribution of corals in the family Primnoidae () in Alaska waters. zoogeographic study if suspected of being unknown or crab species (Stone, 2014; Stone et al., 2015b). Many a bathymetric or geographic range extension. Conven- species, particularly those in shallower water, host a di- tional terminology is used to describe the various species verse and abundant suite of fauna, particularly brittle (Bayer et al., 1983; Cairns, 2016). stars (Ophiuroidae) that are often cryptically colored. Most taxa are generally not much larger than 30 cm Those species within the current range of fishing activi- in height or width but Primnoa pacifica attains great ties (approximately 1200 m depth) are often observed size: up to 5 m in both height and width in exceptional damaged or detached on the seafloor (Krieger, 2001; specimens. Most taxa are found either in low- or high- Stone, 2006, 2014; Stone et al., 2015b) and are com- density patches, and these patches may provide impor- mon bycatch in fisheries stock assessment surveys (Heif- tant refuge and foraging habitat for juvenile fish and etz, 2002). Chapter 13 235 1. Arthrogorgia kinoshitai Bayer, 1952 Description (Adapted from Cairns, 2011a and Bayer, of Arthrogorgia occur in the Aleutian Islands (Cairns, 1952b) Colonies almost always branch in a uniplanar 2011a). Arthrogorgia kinoshitai is quite similar to and irregularly dichotomous fashion, rarely monopodial Arthrogorgia utinomii Bayer, 1996, but differs in hav- in shallower water; the largest known colony is about ing longer buccal scales that form a cowl around the 35 cm in height. Polyps are arranged in whorls of six to operculum, and in having fewer or no infrabasal scales nine with their mouths directed downward. Polyps are (Cairns, 2011a). 3.0–3.5 mm in length with about 2.5 polyps/cm branch This species hosts a rich suite of associated fauna length. The axis is light brown to golden brown; color including pleustid amphipods and antarcturid isopods of the coenenchyme is very light orange to orange in life (Watling and Stone7). The coenenchymal sclerites and yellowish-orange when dried. are composed of high-magnesium calcite (8.3 mol% Each polyp is protected by eight opercular scales; one MgCO3) whereas the skeletal axis is composed entirely pair of buccal and basal scales; zero to three rows of of aragonite (senior author and S. D. Cairns, unpubl. transverse, curved infrabasal scales; and four to six pairs data). of adaxial scales. The distal edge of the buccal scales are rounded and elongate, forming a cowl around the oper- Distribution Uncommon (Fig. 13-3). In Alaska – en- cular scales; the distal edge of each basal scale bears a demic to the Aleutian Islands (Tanaga Pass to west of long, longitudinally ridged spine. Coenenchymal scales Attu Island). Elsewhere – not reported. are stellate to irregular in shape. Habitat Typically found singly but occasionally in Remarks Arthrogorgia is quite similar to Paracalyp- small patches, attached to cobbles and large pebbles, in trophora but differs in having multiple transverse rows areas of moderate current, and at depths between 150 of infrabasal scales. Three of the four known species and 1309 m. Figure 13-3 A map of the Aleutian Islands showing the distribution of Arthrogorgia kinoshitai () in Alaska waters. 236 Professional Paper NMFS 23 Photos A) Dried branches of an A. kinoshitai colony at a depth of 1307 m. C) A lateral view of the same (USNM 1481912) collected on Amchixtam Chaxsxii, specimen in photo B. D) A close-up view of the same central Aleutian Islands, at a depth of 150 m. B) A specimen in photo B. E) The same colony in photo B dried whole A. kinoshitai colony (USNM 1115594) prior to collection. The distance between the red laser collected in Amchitka Pass, central Aleutian Islands, marks is 10 cm. Chapter 13 237 2. Arthrogorgia otsukai Bayer, 1952 Description (Adapted from Cairns, 2011a and Bayer, mouth of the Aangan River, Kamchatka.” Later Bayer 1952b) Colonies have uniplanar, equal, and dichotomous (1996) indicates that the “exact station data were oblit- branching; the largest known colony is 20 cm in height. erated.” Cairns (2011a) further indicates that the col- Polyps are arranged in whorls of five to seven with their lection lot consists of 16 colonies from which Bayer mouths directed downward. Polyps are 2.30–2.75 mm (1952b) did not specifically designate a holotype, hence in length with 3.5–4.5 polyps/cm branch length. The axis the lot consists of 16 syntypes, but again with no addi- is golden brown; the color of the coenenchyme is light tional station data. orange to orange in life and yellowish-orange when dried. The location description provided is quite confound- Each polyp is protected by eight opercular scales; one ed since the Aangan River is on the west side of the Kam- pair each of buccal and basal scales; four or five trans- chatka Peninsula and drains into the Sea of Okhotsk and verse rows of small, curved infrabasal scales; and two is not even in the same sea as Bowers Bank (eastern Ber- short rows (four to six scales in each row) of adaxial ing Sea, U.S.). Furthermore, we can find no formal ref- scales. The distal edges of the buccal and basal scales are erence to “codfish banks” aside from the one written in lobate. The coenenchymal scales are thin, flat, polygo- the U.S. Fisheries Steamer Albatross expedition notes re- nal, and up to 0.5 mm in width. garding the 1906 collections. The Smithsonian Museum of Natural History database currently lists the specimen Remarks Arthrogorgia otsukai differs from other spe- as being collected on 21 June 1906 at a depth of 46 m, cies in the genus in lacking spines on its basal scales. which corresponds to the date and depth for U.S. Fisher- Much uncertainty underlies the type locality of the ies Steamer Albatross station 4798 near the mouth of the holotype (USNM 49979) described by Bayer (1952b), Aangan River in the Sea of Okhotsk. The depth recorded who provided only that the specimen was collected in at that station is much too shallow for the species howev- 1906 by the U.S. Fisheries Steamer Albatross “between er. Of the seven stations (4793–4799) surveyed “off the Bowers Bank, Bering Sea and the codfish banks off the mouth of the Aangan River,” station 4797 (52°37.30´N, 158°50.00´E) with a depth of 1247 m (compared to 46–126 m for the other six stations) is most likely the actual location of the type locality. We suggest that the specimen was indeed collected at station 4797 but not retrieved from the trawl gear until after sta- tion 4798 was surveyed the next day. This species hosts ophiu- roids. The coenenchymal scales are composed of high-magnesium calcite (7.8 mol% MgCO3; senior au- thor and S. D. Cairns, un- publ. data). Distribution Rare. In Alaska – known only from two locations (south of Tanaga Island and southern Amchitka Pass) in the cen- Figure 13-4 tral Aleutian Islands (Fig. A map of the Aleutian Islands showing the distribution of Arthrogorgia otsukai ( ) in Alaska 13-4) but fairly abundant at  waters. those locations. Elsewhere – Sea of Okhotsk, Russia. 238 Professional Paper NMFS 23 Habitat In Alaska – typically found in small patches coenenchymal webbing between its branches. B) A close- but occasionally singly, on hard rock including cob- up view of the same specimen in photo A. C) A patch of bles and siltstone, in areas of moderate current, and at A. otsukai in southern Amchitka Pass at a depth of 1349 depths between 1332 and 1348 m. Elsewhere – found at m. Colonies (USNM 1115597 and 115595) are within a depth of 1247 m. the white circle. A Verrill’s king crab (Paralomis verrilli) takes shelter among the corals. D) An A. otsukai colony Photos A) A dried whole A. otsukai colony (USNM observed in the same patch in southern Amchitka Pass 1115597) collected in southern Amchitka Pass, central at a depth of 1359 m. The distance between the red laser Aleutian Islands, at a depth of 1348 m, showing a unique marks in photos C and D is 10 cm. Chapter 13 239 3. Arthrogorgia utinomii Bayer, 1996 Description (Adapted from Bayer, 1996 and Cairns, This species hosts ophiuroids. The coenenchymal 2011a) Colonies branch in an equal and dichotomous sclerites are composed principally of aragonite (67%), fashion; the largest known colony is 40 cm in height with the remainder high-magnesium calcite (8.8 mol% and 25 cm in width. Polyps are arranged in whorls of MgCO3; senior author and S. D. Cairns, unpubl. data). 7–10 with their mouths directed downward. Polyps are 3.0–3.5 mm in length with 2.5–3.0 polyps/cm branch Distribution Uncommon (Fig. 13-5). In Alaska – en- length. The axis is golden to dark brown; the coenen- demic to the Aleutian Islands south of Yunaska Island chyme is pinkish orange to light orange in life and yel- to southwest of Attu Island. Elsewhere – not reported. lowish-orange when dried. Each polyp is protected by eight opercular scales; Habitat Typically found in small patches but occa- one pair each of buccal and basal scales; two to four sionally singly, on hard rock including cobbles, in areas rows of transverse, curved infrabasal scales; one pair of of moderate current, and at depths between 163 and relatively large adaxial scales; and smaller adaxial scales 882 m. proximal to those. The distal edge of the buccal scales is rounded, but the distal edge of each basal scale bears a long, longitudinally ridged spine. Coenenchymal scales Photos A) A fresh whole A. utinomii colony (USNM are elongate, irregularly shaped, and up to 0.65 mm in 1115598) collected in northern Amchitka Pass, central length. Aleutian Islands, at a depth of 712 m. B) A close-up view of the same specimen in photo A, dried. C) The Remarks Arthrogorgia utinomii is very similar to A. same colony in photo A just prior to collection. The dis- kinoshitai and is compared to that species (see Remarks tance between the red laser marks is 10 cm. on page 235). 240 Professional Paper NMFS 23 3. Arthrogorgia utinomii Bayer, 1996 (continued) Figure 13-5 A map of the Aleutian Islands showing the distribution of Arthrogorgia utinomii () in Alaska waters. Chapter 13 241 4. Callogorgia compressa (Verrill, 1865) Description (Adapted from Cairns, 2011a) Colonies Remarks Until recently this species was placed in the branch in a uniplanar, equal, and dichotomous fashion; genus Fanellia because of its coarse tuberculate body the largest known colony is 80 cm in height and 50 cm wall scales, but it has since been transferred to Cal- in width. Colonies may have more than 100 branch logorgia based on molecular evidence (Cairns and Wir- tips. Polyps face upward and are arranged in whorls shing, 2018). This species is compared to Callogorgia of 8–12 on distal branches, but up to 25 on larger- fraseri (Hickson, 1915) in the account of that species. diameter branches. Polyps are 1.6–2.0 mm in length Callogorgia compressa colonies are often heavily with 4–6 whorls/cm branch length. The holdfasts are populated with ophiuroids (>100/colony; see photos C covered with coenenchyme and strongly attached to the and D on next page). Other associated fauna include the substrate. The axis is metallic bronze; the coenenchyme bivalve Acesta sp. that attaches to exposed areas of the is orange to light orange in life and a paler yellowish- axial skeleton (Watling and Stone7). orange when dried. Each polyp is protected by eight opercular scales and Distribution Locally common. In Alaska – endemic to eight longitudinal rows of body wall scales; the sclerite the Aleutian Islands from near Umnak Island to west of formula is 7–10: 5–7, 2–3: 1. Body wall and opercular Attu Island (Fig. 13-6). Elsewhere – not reported. scales are covered with coarse, longitudinally aligned tubercles. The coenenchymal sclerites consist of close- Habitat Typically found in small patches, rarely singly, ly fitted polygonal plates and numerous tuberculate on hard rock (cobbles and pebbles), in areas of moderate spheroids. current, and at depths between 82 and 1341 m. Figure 13-6 A map of the Aleutian Islands showing the distribution of Callogorgia compressa () in Alaska waters. 242 Professional Paper NMFS 23 Photos A) A fresh whole (except the holdfast) C. C) A C. compressa colony harboring many ophiuroids, compressa colony (USNM 1123883) collected on the photographed northwest of Adak Island at a depth of Adak Canyon shelf, central Aleutian Islands, at a depth 210 m. D) A close-up view of the same colony in photo of 155 m. B) A close-up view of a C. compressa speci- C. The distance between the red laser marks in photos men (USNM 1411817) collected near the Delarof Is- C and D is 10 cm. lands, central Aleutian Islands, at a depth of 100 m. Chapter 13 243 5. Callogorgia fraseri (Hickson, 1915) Description (Adapted from Cairns, 2011a) Colonies nenchymal sclerites are composed of high-magnesium branch in a uniplanar, alternate pinnate fashion; the calcite (12.4 and 8.7 mol% MgCO3, respectively; senior largest known colony is 55 cm in height and 35 cm in author and S. D. Cairns, unpubl. data). width. Colonies have up to 100 branch tips. Polyps face upward and are arranged in whorls of five to eight on Distribution Locally common. In Alaska – from Al- distal branches, but up to 12 on larger-diameter branch- batross Bank in the western Gulf of Alaska to Amchitka es. Polyps are 1.1–1.5 mm in length with 6–7 whorls/cm Pass in the central Aleutian Islands (Fig. 13-7). Else- branch length. The holdfasts are covered with coenen- where – not reported. chyme and strongly attached to the substrate. The axis is brownish-yellow; the coenenchyme is orange to light Habitat Typically found in small patches, rarely sin- orange in life and a paler yellowish-orange when dried. gly, on hard rock (cobbles and pebbles), in areas of Each polyp is protected by eight opercular scales moderate current, and at depths between 52 and 1341 and eight longitudinal rows of body wall scales; the m. sclerite formula is 7–8: 2–3: 1: 0–1. The opercular and body wall scales are covered with coarse, longitudinally Photos A) A fresh whole (except the holdfast) C. fra- aligned tubercles. The coenenchymal sclerites consist of seri colony (USNM 1123890) collected near Little Tan- tuberculate spheroids. aga Island, central Aleutian Islands, at a depth of 98 m. B) A close-up view of the same specimen in photo A. C) Remarks This species differs from C. compressa by A dried branch of a C. fraseri colony (USNM 1123891) having a pinnate colony, fewer body wall scales, more collected south of Adak Island, central Aleutian Islands, whorls per centimeter, and smaller polyps. at a depth of 100 m. D) A C. fraseri colony (USNM This species is often heavily populated with ophiu- 1123890) prior to collection. The distance between the roids (>100 per specimen). The skeletal axis and coe- red laser marks is 10 cm. 244 Professional Paper NMFS 23 5. Callogorgia fraseri (Hickson, 1915) (continued) Figure 13-7 A map of the North Pacific Ocean showing the distribution of Callogorgia fraseri () in Alaska waters. Chapter 13 245 6. Calyptrophora laevispinosa Cairns, 2007 Description (Adapted from Cairns, 2007, 2011a) cies known worldwide (Cairns, 2018). Some colonies Colonies branch usually in a uniplanar but occasionally observed in situ appear to have been preyed upon by an bushy, even scraggly manner (see photo F on next page). unknown predator (likely a sea star). They are dichotomously branched with a slight tendency Small colonies are morphologically similar (e.g., to be pinnate. The largest known colony is 70 cm in branching pattern) to Keratoisis sp. D with which it co- height and 59 cm in width. Large colonies have up to occurs, but the latter species is quite fragile whereas the 80–100 branch tips and prominent, often irregular hold- former is very difficult to tear upon collection. fasts. The axis is golden brown; the coenenchyme is very light orange to white in life and when dried. Distribution Uncommon. In Alaska – known only The polyps face upwards and are arranged in whorls from seamounts in the Gulf of Alaska Seamount Prov- of three or four. Polyps are 1.5–2.0 mm in length with ince (Welker, Pratt, Giacomini, Patton, and Derickson 3.3–4.0 whorls/cm branch length. Each polyp is protect- Seamounts). Also observed on video footage collected ed by eight opercular scales, one basal and one buccal on Dickens and Murray Seamounts (Fig. 13-8). This scale, and two pairs of slender, curved infrabasal scales. species has not been documented from the continental Each basal scale bears two long and slender, smooth slope or the Aleutian Islands. Elsewhere – only docu- (not ridged) spines; each buccal scale bears four to six mented on Gorda Flow on the continental slope off slender, non-ridged spines. The coenenchymal scales Washington. are elongate (up to 1.3 mm in length), elliptical, and sparsely granular. Habitat In Alaska – typically found singly but occa- sionally in small, scattered patches on bedrock, frac- Remarks Calyptrophora laevispinosa is the only Ca- tured bedrock, boulders and cobbles, in areas of moder- lyptrophora species known from Alaska waters and ate current, and at depths between 1754 and 3531 m. one of the deepest dwelling among the 27 extant spe- Elsewhere – at a depth of 3107 m. Figure 13-8 A map of the North Pacific Ocean showing the distribution of Calyptrophora laevispinosa () in Alaska waters. 246 Professional Paper NMFS 23 Photos A) A dried large branch complex of C. lae- 1075474) collected on Pratt Seamount at a depth of vispinosa (USNM 1081196) collected on Derickson Sea- 2676 m. E) A close-up view of the same colony in photo mount at a depth of 3444 m. B) A close-up view of the D. F) A C. laevispinosa colony (USNM 1082617) col- same specimen in photo A. C) A C. laevispinosa colony lected on Patton Seamount at a depth of 1993 m. The (USNM 1075473) collected on Pratt Seamount at a distance between the red laser marks in photos C, D, depth of 2776 m. D) A C. laevispinosa colony (USNM and F is 10 cm. Chapter 13 247 7. Narella abyssalis Cairns and Baco, 2007 Description (Adapted from Cairns and Baco, 2007) three) pairs of body wall scales. It is the deepest dwell- The holotype colony has uniplanar, sparse, dichoto- ing species in the genus. mous branching and is 33 cm in height and 42 cm in width. The polyps face downward and are arranged in Distribution Rare. In Alaska – known only from the whorls of two to four. Polyps are 1.9–2.4 mm in length holotype collected on Derickson Seamount in the Gulf with 3 whorls/cm branch length. The axis is golden of Alaska Seamount Province (Fig. 13-9). Elsewhere – brown; the coenenchyme is very light orange in life and not reported. in ethanol. Each polyp is protected by eight opercular scales, Habitat The holotype was collected at a depth of four pairs of abaxial body wall scales (basals, two pairs 4594 m but other specimens observed on video foot- of medials, and buccals), and one or two pairs of small, age at depths between 4559 and 4633 m appeared to elliptical adaxial body wall scales. The dorsolateral be this species (the other four species are unknown at edges of all body wall scales are prominently ridged. these depths). Colonies were always observed singly or The coenenchymal scales are fusiform, up to 1.3 mm in small patches on bedrock (basalt) and fractured bed- in length, and each bears a tall medial ridge (i.e., a sail rock. scale). Photos A) Preserved (in ethanol) branches of a N. Remarks A tabular key was published (Cairns and abyssalis holotype (USNM 1080450) collected on De- Baco, 2007) comparing the five Narella species known rickson Seamount at a depth of 4594 m. B) A close-up from Alaska, and tabular and dichotomous keys have view of the same specimen in photo A. C) A suspected recently been published for all 50 known species in the N. abyssalis colony on Derickson Seamount at a depth genus (Cairns and Taylor, 2019). Narella abyssalis dif- of 4633 m. The distance between the red laser marks is fers from all other Alaska species by having four (not 10 cm. 248 Professional Paper NMFS 23 7. Narella abyssalis Cairns and Baco, 2007 (continued) Figure 13-9 A map of the North Pacific Ocean showing the distribution of Narella abyssalis () in Alaska waters. Chapter 13 249 8. Narella alaskensis Cairns and Baco, 2007 Description (Adapted from Cairns and Baco, 2007) fusiform, up to 0.9 mm in length, and bear a tall medial Colonies branch in a uniplanar, sparsely dichotomous ridge (i.e., a sail scale). fashion, with up to 30 branch tips in large specimens. The largest colony has dimensions of 41 cm in height Remarks Narella alaskensis differs from all other and 25 cm in width. Colonies appear to be relatively Alaska species by having basal scale spines (Cairns and flexible and wave gently in the current in situ. All pol- Baco, 2007). Colonies are occasionally overgrown with yps face downward and are arranged in whorls of seven small actiniarians (see photo D on next page) and often to nine on large-diameter branches and whorls of four host large numbers of ophiuroids. or five on small-diameter branches. Polyps are 2.7–3.2 mm in length with 3.5 whorls/cm branch length. Large Distribution Common (Fig. 13-10). In Alaska – specimens have prominent, stout, irregularly shaped grows in low-density patches on seamounts in the holdfasts without coenenchyme. The axis is golden Gulf of Alaska Seamount Province (Dickens, Densen, brown; the coenenchyme is very light orange to white in Welker, Murray, and Chirikof Seamounts). Particu- life and light orange in ethanol. larly abundant on Murray Seamount (up to 5 colonies/ Each polyp is protected by eight opercular scales; a m2) where it is the dominant coral in its depth range. pair of basal, medial, and buccal body wall scales; and This species has not been documented on the conti- one or two pairs of small oval adaxial body wall scales. nental slope or from the Aleutian Islands. Elsewhere The abaxial body wall scales are quite flared. The dorso- – only known from San Marcos Seamount off central lateral edge of basal and medial scales sometimes bears California. a low ridge; the buccal body wall scales have no ridges. The distal edge of each basal scale bears a projecting Habitat In Alaska – observed on video footage singly broad, blunt spine or spur. The coenenchymal scales are or in small patches on bedrock (basalt) and fractured Figure 13-10 A map of the North Pacific Ocean showing the distribution of Narella alaskensis () in Alaska waters. 250 Professional Paper NMFS 23 bedrock (boulders and cobbles) at depths between 2216 A close-up view of the same specimen in photo A. C) and 3204 m. Elsewhere – 2193 m depth. A N. alaskensis colony (USNM 1075471) collected on Welker Seamount at a depth of 2634 m. D) A close-up Photos A) A preserved (in ethanol) branch of a N. view of the same colony in photo C showing attached alaskensis colony (USNM 1080453; paratype) col- actiniarians. The distance between the red laser marks lected on Murray Seamount at a depth of 2680 m. B) in photos C and D is 10 cm. Chapter 13 251 9. Narella arbuscula Cairns and Baco, 2007 Description (Adapted from Cairns and Baco, Narella by having a gently curved dorsolateral edge to 2007) Colonies are slightly bushy and dichotomously its basal scales and in having larger polyps. branched; the terminal branches (30+) are up to 12 cm in length. Colonies measure up to 30 cm in height Distribution Uncommon (Fig. 13-11). In Alaska – and width. The branches typically originate just above known only from Giacomini and Derickson Seamounts the holdfast. Colonies appear to be relatively flexible, in the Gulf of Alaska Seamount Province. This species with long floppy branches that wave gently in the cur- has not been documented on the continental slope or rent. The coenenchyme is very thin on most specimens, from the Aleutian Islands. Elsewhere – not reported. clearly revealing the axis beneath. All polyps face down- ward and are arranged in whorls of six or seven. Polyps Habitat Collected or observed on video footage singly are large (3.4–4.7 mm in length) with 2.0–2.5 whorls/ or in small patches on bedrock (basalt) and fractured cm branch length. The axis is a brilliant gold to golden bedrock (boulders and cobbles) at depths between 2710 brown; the coenenchyme is very light orange to white in and 3432 m. life and in ethanol. Polyps are protected by eight opercular scales, three Photos A) A large portion of a preserved (in ethanol) pairs of abaxial body wall scales, and one pair of small, N. arbuscula colony (USNM 1075466; paratype) col- oval adaxial buccal scales. The dorsolateral edges of lected on Giacomini Seamount at a depth of 2819 m. basal scales are sometimes ridged; other body wall B) A close-up view of the same specimen in photo A. scales are not ridged. Coenenchymal scales are elliptical C) The same colony in photo A prior to collection. D) in shape, up to 0.5 mm in diameter, and bear one or two A close-up view of the same colony in photo C. E) A N. very tall ridges (i.e., a sail scale). arbuscula colony (USNM 1074565) collected on Giaco- mini Seamount at a depth of 2819 m. The distance be- Remarks This species differs from all other Alaska tween the red laser marks in photos C and E is 10 cm. Figure 13-11 A map of the North Pacific Ocean showing the distribution of Narella arbuscula () in Alaska waters. 252 Professional Paper NMFS 23 9. Narella arbuscula Cairns and Baco, 2007 (continued) Chapter 13 253 10. Narella bayeri Cairns and Baco, 2007 Description (Adapted from Cairns and Baco, 2007) (2007) and to all other species worldwide by Cairns Colonies are branched in one plane, with sparse dichot- and Taylor (2019). This species has strongly ridged bas- omous branching. Colonies usually have no more than al and medial scales, crowded whorls, and 5–7 polyps/ ten branches, but the largest colonies are up to 29 cm in whorl. height with 50 branch tips. All polyps face downward The branches of N. bayeri appear to be more rigid and are arranged in whorls of five to seven. Polyps are than those of N. arbuscula, with which it co-occurs. The 2.2–3.4 mm in length with about 3.8 closely spaced skeletal axis is composed entirely of aragonite (senior whorls per centimeter branch length. The axis is golden author and S. D. Cairns, unpubl. data). brown; the coenenchyme is very light orange to white in life and light orange in ethanol. Distribution Uncommon. In Alaska – known from Each polyp is protected by eight opercular scales, only three specimens: one collected on the continental three pairs of abaxial body wall scales, and one pair of shelf south of Davidson Bank (western Gulf of Alaska) small adaxial buccal scales. The dorsolateral edge of and two on nearby Derickson Seamount (Fig. 13-12). both basal and medial scales forms a 90° corner, which This is the only Narella species documented from the bears a prominent thin ridge; the buccal scales are not continental slope in the Gulf of Alaska. Elsewhere – not angled or ridged. The coenenchymal scales are fusiform, reported. up to 2.1 mm in length, and bear a complex series of tall ridges (i.e., a sail scale) that sometimes forms a network Habitat Likely occurs singly or in low-density patches on the outer surface of the scale. (up to 2 colonies/m2) on moderately steep habitat of bedrock (basalt), fractured bedrock, and conglomerate Remarks Narella bayeri is compared to the other (a consolidated mix of sand and cobbles), at depths be- Alaska species in a tabular key by Cairns and Baco tween 3280 and 4092 m. Figure 13-12 A map of the North Pacific Ocean showing the distribution of Narella bayeri () in Alaska waters. 254 Professional Paper NMFS 23 Photos A) Preserved branches of a N. bayeri colony C) A N. bayeri colony observed on Derickson Seamount (USNM 1080448; paratype) collected south of David- at a depth of 3793 m. D) A close-up view of the same son Bank, western Gulf of Alaska, at a depth of 3280 colony in photo C. The distance between the red laser m. B) A close-up view of the same specimen in photo A. marks in photos C and D is 10 cm. Chapter 13 255 11. Narella cristata Cairns and Baco, 2007 Description (Adapted from Cairns and Baco, 2007) has smaller polyps, fewer polyps per whorl, and fewer The holotype colony has uniplanar, sparse, and di- whorls per centimeter (Cairns and Baco, 2007). chotomous branching; the longest branch is 8.5 cm in length. The maximum colony height and width is 28 Distribution Likely rare. In Alaska – known from cm. The coenenchyme is very thin on most specimens, only a single specimen collected on Derickson Seamount clearly revealing the axis beneath. Polyps always face in the Gulf of Alaska Seamount Province (Fig. 13-13). downward and are arranged in whorls of two to four. This species has not been documented on the continen- Polyps are 2.1–3.0 mm in length with 2.5–3.3 whorls/ tal slope or from the Aleutian Islands. Elsewhere – not cm branch length. The axis is light to golden brown; the reported. coenenchyme is very light orange to white in life and Habitat Likely occurs singly or in small patches on pinkish in ethanol. bedrock (basalt) and fractured bedrock (cobbles) at Each polyp is protected by eight opercular scales, depths near 3358 m. Observed on video footage to three pairs of abaxial body wall scales, and one pair depths as shallow as 3022 m. of small, elliptical adaxial buccal scales that bear a small central boss. All three pairs of abaxial body Photos A) The preserved (in ethanol) N. cristata ho- wall scales bear a low ridge on their dorsolateral edg- lotype (USNM 1080449; branch fragments of a colony) es. Coenenchymal scales are fusiform, up to 0.8 mm collected on Derickson Seamount at a depth of 3358 m. in length, and each bears a tall medial ridge (i.e., a B) A close-up view of the same specimen in photo A. sail scale). C) The same colony in photo A (indicated by the white circle) prior to collection. The distance between the red Remarks This species is similar to N. bayeri, but laser marks is 10 cm. 256 Professional Paper NMFS 23 11. Narella cristata Cairns and Baco, 2007 (continued) Figure 13-13 A map of the North Pacific Ocean showing the distribution of Narella cristata () in Alaska waters. Chapter 13 257 12. Parastenella doederleini (Wright and Studer, 1889) Description (Adapted from Cairns, 2011a) Colonies large nematocyst pads on the inner surface of its fluted are bushy, dichotomously branched, and relatively small; marginal scales (Cairns and Bayer, 2009). Parastenella the largest colony known from Alaska is only 6 cm in doederleini is most similar to P. ramosa, but differs height but up to 25 cm in height elsewhere in its geo- from that species by having longer and narrower fluted graphical range. The axis is dark brown and easily seen marginal scales, ornamented coenenchymal scales, and through the thin, translucent coenenchyme. The polyps shorter polyps (Cairns, 2007, 2010, 2011a). are isolated, in pairs, or in whorls of three, and project perpendicular to the branch; polyps are 2.1–2.5 mm in Distribution Likely rare. In Alaska – known from length. The coenenchyme is light orange to very light only three specimens collected near Adak Canyon in the pink in life and yellowish in ethanol. central Aleutian Islands (Fig. 13-14). Elsewhere – the Each polyp is protected by eight opercular scales and holotype was collected off Sagami Bay (Honshu Island, five or six longitudinal rows of body wall scales which Japan) and one additional specimen is known from the encircle the polyp (i.e., the adaxial side of the polyp is Ceram Sea, Indonesia. covered with scales). The eight opercular scales form an Habitat Attaches to siltstone at depths between 1746 asymmetrical rosette of narrowly fluted (almost tubular) and 2539 m. Elsewhere – found at depths between 732 scales; some abaxial submarginal body wall scales also and 3423 m. bear lesser-fluted scales. The coenenchymal scales are elongate, up to 0.55 mm in length, and often bear a boss Photos A) A whole preserved (in ethanol) P. doederle- or low ridge at the center. ini colony (USNM 1115561) collected in Adak Canyon, central Aleutian Islands, at a depth of 1746 m. B) A Remarks A key to seven of the eight species in this P. doederleini colony observed growing on siltstone in genus was published by Cairns (2010). Parastenella are Adak Canyon at a depth of 2539 m. C) A close-up view distinctive from other Primnoidae in having marginal of the same colony in photo B. The distance between scales offset from the opercular scales and in having the red laser marks in photos B and C is 10 cm. 258 Professional Paper NMFS 23 12. Parastenella doederleini (Wright and Studer, 1889) (continued) Figure 13-14 A map of the Aleutian Islands showing the distribution of Parastenella doederleini () in Alaska waters. Chapter 13 259 13. Parastenella gymnogaster Cairns, 2007 Description (Adapted from Cairns, 2011a) Colonies photo D on next page). Some colonies are complete- have uniplanar, irregularly dichotomous branching ly overgrown with epizoic zoantharians (Zibrowius that rarely is slightly bushy. Colonies have a height and sp.) and are occasionally preyed upon by the sea star width up to 100 cm. Polyps are isolated in pairs, ar- Hippasteria. ranged in whorls of three, face downward, and are 2.2– 3.0 mm in length. Colonies are supported by a large, Distribution Uncommon but locally abundant (Fig. prominent, robust holdfast (≥6 cm in diameter) that is 13-15). In Alaska – known only from seamounts in the white and reflective in situ. The axis and main branches Gulf of Alaska Seamount Province (Dickens, Welker, are bronze in color, lightly striated, and faintly visible Marchand, Chirikof, and Derickson Seamounts). Else- through the thin coenenchyme. The coenenchyme is where – the Emperor Seamounts in the central-west Pa- pink, sometimes with a purplish hue, or pale orange in cific Ocean (Jingu, Nintoku, Godaigo, Yomei, and Suiko life, and paler shades in ethanol. Seamounts), Juan de Fuca Ridge and Vance Seamount in Each polyp is protected by eight opercular and eight Oregon, and Monterey Bay Canyon in California. marginal scales, but proximal to these are only six lon- gitudinal rows of body wall scales; the scales of the ab- Habitat In Alaska – occurs singly or in small but axial rows are broad and linearly arranged as though sometimes dense patches (>5 colonies/m2 on March- stacked. The eight marginal scales form an asymmetri- and Seamount) on bedrock (basalt), fractured bedrock, cal rosette of short and broad distal flutes; the abaxial boulders, and cobbles including in crevices and un- submarginal body wall scales also bear flutes. The coe- der overhangs, at depths between 2193 and 3074 m. nenchymal scales are irregularly shaped, concave above, Patches of larger colonies resemble the thickets of Prim- and bear a complex series of ridges. noa pacifica in the eastern Gulf of Alaska. Elsewhere – depths between 1100 m (Monterey Bay Canyon) and Remarks Parastenella gymnogaster differs from other 2077 m (Godaigo Seamount). Alaska Parastenella by having only six rows of body wall scales, broad and short fluted scales, and ornately ornamented coe- nenchymal scales (Cairns, 2007, 2011a). Parastenella gymno- gaster, along with Prim- noa wingi, are the only two corals found both in Alaska waters and on the Emperor Seamounts in the central-west Pacific Ocean. The Emperor Seamounts are a series of very old and heavily eroded guyots (flat- topped) and seamounts stretching approximately 6700 km from the Aleu- tian Trench to the North- western Hawaiian Islands (Cairns et al., 2018). They provide an interesting model on the study of the biogeography of North Pa- Figure 13-15 cific Ocean corals. Colonies host several A map of the North Pacific Ocean showing the distribution of Parastenella gymnogaster () in Alaska waters. species of ophiuroids (see 260 Professional Paper NMFS 23 Photos A) The preserved (in ethanol) P. gymnogaster base of the colony (with associated ophiuroids) is clearly holotype (USNM 1075463; many branches of a colony) visible. E) A linear row of six large P. gymnogaster colo- collected on Welker Seamount at a depth of 2635 m. B) nies growing on bedrock on Marchand Seamount at a A close-up view of the same specimen in photo A. C) A depth of 2323 m. F) Two large P. gymnogaster colonies P. gymnogaster colony (USNM 1075359) growing on observed on Marchand Seamount at a depth of 2316 m. basalt on Dickens Seamount at a depth of 2789 m. D) The colony at the upper left is completely overgrown A P. gymnogaster colony (USNM 1075361) growing on with zoantharians. The distance between the red laser basalt on Dickens Seamount at a depth of 2704 m. The marks in photos C, E, and F is 10 cm. Chapter 13 261 14. Parastenella ramosa (Studer, 1894) Description (Adapted from Cairns, 2011a) Colonies why seamount specimens are often completely covered usually branch in a uniplanar fashion, with irregular, with zoantharians (i.e., the skeleton remains intact and dichotomous branching; the largest colonies measure up attached to the substrate after death, providing clean, to 55 cm in height and 45 cm in width. The axis is dark elevated substrate for other fauna). bronze and usually visible beneath the coenenchyme in This species hosts a rich suite of associated fauna live specimens. Polyps occur singly, in pairs and whorls including amphipods (families Caprellidae, Ischyrocer- of three, and usually directed basally; polyps are 2.5– idae, and Pleustidae), the isopod Munna sp. (Watling 3.3 mm in length. The coenenchyme is light orange to and Stone7), ophiuroids including Asteronyx sp., and white in live specimens and when dried or preserved in chirostylid crab species. Colonies on Gulf of Alaska ethanol. seamounts are often overgrown with zoantharian corals Each polyp is protected by eight opercular scales, (Zibrowius sp.). It is preyed upon by the sea star Hippas- eight marginal scales, and five rows of body wall scales. teria sp. (see photo E on next page). The coenenchymal The eight marginal scales form an asymmetrical rosette sclerites are composed principally of aragonite (69.2%) of narrow, almost tubular, fluted scales; abaxial sub- and lesser amounts (30.8%) of high-magnesium calcite marginal scales also bear small flutes. The coenenchy- (8.2 mol% MgCO3; senior author and S. D. Cairns, un- mal scales are irregular to elliptical in shape, concave publ. data). above, up to 0.6 mm in diameter, and not ridged. Distribution Common and broadly distributed (Fig. Remarks This species is similar to P. doederleini (see 13-16). In Alaska – eastern Gulf of Alaska continen- Remarks of that species) and is compared to others in tal shelf, Gulf of Alaska Seamount Province (Dickens, the genus by Cairns (2007, 2011a). Preserved specimens Welker, Pratt, Giacomini, Murray, and Patton Sea- are fragile but live specimens have a strong holdfast and mounts), the central Aleutian Islands (Adak Canyon), are difficult to dislodge in situ. This may partly explain and the eastern Bering Sea. Elsewhere – extremely wide- Figure 13-16 A map of the North Pacific Ocean showing the distribution of Parastenella ramosa () in Alaska waters. 262 Professional Paper NMFS 23 ranging geographically including the western Bering Sea 1075364) collected on Dickens Seamount at a depth of (Commander Islands, Russia), British Columbia, Wash- 850 m. B) A close-up view of the same specimen in pho- ington, California, Panama, and Chile. to A. C) A P. ramosa colony (USNM 1481813) growing on bedrock in Adak Canyon, central Aleutian Islands, Habitat In Alaska – occurs singly or in small patches at a depth of 1429 m. D) A close-up view of a P. ra- on bedrock including basalt lava flows, fractured bed- mosa colony (USNM 1075377) growing on basalt on rock, boulders, cobbles, and occasionally siltstone, at Pratt Seamount at a depth of 960 m. E) A P. ramosa depths between 667 and 1937 m. Observed growing colony (USNM 1075379) being preyed upon by a sea on hexactinellid (Chonelasma oreia) skeleton on Welk- star (Hippasteria sp.) on Pratt Seamount at a depth of er Seamount (Gulf of Alaska Seamount Province) at a 918 m. F) A band of P. ramosa colonies, including spec- depth of 1123 m. Elsewhere – depths between 619 m imen USNM 1075367 (at left), on Welker Seamount (Channel Islands, California) and 1805 m (off Vancou- at a depth of 795 m. The colonies host numerous crab ver Island, British Columbia). (Sternostylus sp.), ophiuroids (Asternonyx sp.), and zo- antharians (Zibrowius sp.). The distance between the Photos A) A dried whole P. ramosa colony (USNM red laser marks in photos C, E, and F is 10 cm. Chapter 13 263 15. Plumarella aleutiana Cairns, 2011 Description (Adapted from Cairns, 2011a) Colonies characteristic were placed in the now defunct subgenus branch in a uniplanar, regular, alternate-pinnate man- Dicholaphis. Of those seven taxa, Plumarella aleutiana ner; the largest known colony is 56 cm in height and is most similar to P. superba (Nutting, 1912) but dif- 42 cm in width. The axis is brown to black and some- fers in having pinnate (not bottlebrush) branching and what stiff. Polyps occur on all sides of the branches in a coarsely serrate body wall scales (Cairns, 2011a). A key crowded fashion with 20–35 polyps/cm branch length. to the Alaska species of this now defunct subgenus (Di- Polyps are small (only 0.9–1.2 mm in length) and cy- cholaphis) is given by Cairns (2011a), including a list of lindrical to slightly flared. The coenenchyme is bright all Plumarella species known at that time. orange to pink in life and orange when dried. This species hosts a rich suite of associated fauna Each polyp is covered with eight opercular scales and including amphipods (families Caprellidae, Ischyroceri- eight longitudinal rows of body wall scales; the sclerite dae, and Pleustidae), ophiuroids, polynoid polychaetes, formula is 5–9: 4–6: 2–5: 2–4. The tip of the marginal and antarcturid isopods (Watling and Stone7). Bivalves scales is sometimes spinose or triangular in shape; the (Paranoidae, Aricidea [Acmira] sp.) occasionally attach margin of the other body wall scales is coarsely serrate. to exposed branch axes. The skeletal axis is composed The coenenchymal scales are irregular in shape, usually of high-magnesium calcite (14.3 mol% MgCO3; senior elongate, up to 0.6 mm in length, and finely granular; author and S. D. Cairns, unpubl. data). smaller (0.05–0.09 mm in diameter) spiny spheroids are also present. Distribution Common, locally abundant. In Alaska – Aleutian Islands near Akutan Island to west of Attu Remarks Seven of the eight Alaska Plumarella spe- Island including Petrel and Bowers Banks, the eastern cies have polyps that occur on all sides of their branches Bering Sea to the Pribilof Islands, and Patton Seamount (versus only on alternating sides) and based on that in the Gulf of Alaska Seamount Province (Fig. 13-17). Figure 13-17 A map of the North Pacific Ocean showing the distribution of Plumarella aleutiana () in Alas- ka waters. 264 Professional Paper NMFS 23 This is the only species of Plumarella reported from the view of a fresh P. aleutiana specimen (USNM 1201880) Gulf of Alaska Seamount Province. Elsewhere – not re- collected in Pribilof Canyon, eastern Bering Sea, at ported. a depth of 300 m. C) A P. aleutiana colony (USNM 1116845) collected in Pribilof Canyon at a depth of Habitat Occurs in small patches and large fields on 310 m. D) A close-up view of the same specimen (fresh) bedrock, boulders, cobbles, pebbles, and occasionally in photo C. E) A P. aleutiana colony observed in Pribi- gastropod shell. Extremely eurybathic; found at depths lof Canyon at a depth of 239 m. A Pacific ocean perch between 79 and 2828 m. (Sebastes alutus) takes refuge beside the colony. F) A cluster of three P. aleutiana colonies observed in Adak Photos A) A fresh whole P. aleutiana colony (USNM Canyon, central Aleutian Islands, at a depth of 2196 m. 1134755) collected southwest of Atka Island, central The distance between the red laser marks in photos C, Aleutian Islands, at a depth of 119 m. B) A close-up E, and F is 10 cm. Chapter 13 265 16. Plumarella echinata Cairns, 2011 Description (Adapted from Cairns, 2011a) Colonies Remarks This species has elongate, spiny marginal branch in a uniplanar, regular, alternate-pinnate man- scales and relatively few body wall scales; these charac- ner; the largest known colony is 28 cm in height and teristics distinguish it from all other Alaska Plumarella width. Larger colonies may have relatively large, robust that have polyps on all sides of the branches (i.e., those holdfasts (up to 3 cm in diameter) that are white and re- species formerly placed in the subgenus Dicholaphis). flective in situ (see photo D on next page). Polyps occur Associated fauna include nemertean polychaetes on all sides of the branches in a crowded fashion, with (Watling and Stone7). Small actiniarians occasionally at- 20–30 polyps/cm of branch length; polyps are slightly tach to exposed branch axes. The coenenchyme is com- flared distally and 1.1–1.4 mm in length. The axis is posed of high-magnesium calcite (14.2 mol% MgCO3; bronze and fairly flexible; the coenenchyme is light or- senior author and S. D. Cairns, unpubl. data). ange to pink in life and a pale yellowish orange when dried. Distribution Common. In Alaska – Aleutian Islands, Each polyp is covered with eight opercular scales and south of Amukta Pass to Buldir Reef, including Pe- eight longitudinal rows of body wall scales; the scler- trel and Bowers Banks (Fig. 13-18). Elsewhere – not ite formula is 3–5: 3–4: 2–3: 2–4. Most marginal scales reported. are prominently spinose; the spines themselves are spiny and not ridged or smooth. The submarginal scales also Habitat Occurs in small patches on bedrock, boul- bear shorter spines, and the remainder of the body wall ders, cobbles, and pebbles at depths between 150 and scales have a triangular margin. Coenenchymal scales 1692 m. are elongate, coarsely granular, and up to 0.55 mm in length. Figure 13-18 A map of the Aleutian Islands showing the distribution of Plumarella echinata () in Alaska waters. 266 Professional Paper NMFS 23 Photos A) A dried whole (except the holdfast) P. echi- Two large P. echinata colonies (holdfasts indicated by nata colony (USNM 1135993) collected in Adak Can- the white circles) collected southwest of Amlia Island, yon, central Aleutian Islands, at a depth of 1692 m. B) central Aleutian Islands, at a depth of 844 m. Colony A dried whole (except the holdfast) P. echinata colo- USNM 1134500 is at left. E) A close-up view of the ny (USNM 1131995) collected in northern Amchitka same colonies in photo D. The distance between the red Pass, central Aleutian Islands, at a depth of 711 m. C) laser marks in photos D and E is 10 cm. A close-up view of the same specimen in photo B. D) Chapter 13 267 17. Plumarella hapala Cairns, 2011 Description (Adapted from Cairns, 2011a) Colo- Associated fauna include amphipods (family Ischyro- nies branch in a uniplanar, coarsely alternate-pinnate ceridae), the isopod Arctura sp. (Watling and Stone7), manner; the largest known colony is 26 cm in height and ophiuroids. The axial holdfast is composed of ara- and 22 cm in width. Polyps occur on all sides of the gonite (senior author and S. D. Cairns, unpubl. data). branches in a crowded manner, with 18–25 polyps/cm of branch length. The polyps are distally flared, some- Distribution Common. In Alaska – Aleutian Islands what fragile (often damaged during collection), and from the Islands of Four Mountains to southwest of large (2.0–2.4 mm in length). The axis is yellowish- Buldir Island including Petrel Bank (Fig. 13-19). Else- brown and relatively stiff. The coenenchyme is light where – not reported. orange to pink in life and light orange to cream when dried. Habitat In Alaska – occurs in small patches on bed- Each polyp is protected by eight opercular scales and rock, boulders, cobbles, and pebbles at depths between eight rows of body wall scales; the sclerite formula is 120 and 407 m. 6–8: 5–7: 3–6: 5–8. The tips of most marginal scales are pointed (not spinose); the submarginal scales are pro- Photos A) A dried whole (except the holdfast) P. gressively less pointed. The body wall scales are thin. hapala colony (USNM 1134212) collected in Amchitka The coenenchymal scales are elliptical to rectangular, Pass, central Aleutian Islands, at a depth of 120 m. B) A with a smooth outer surface. close-up view of the same specimen in photo A. C) A P. hapala colony (USNM 1134094) collected in southern Remarks This species is most similar to Plumarella Amchitka Pass, central Aleutian Islands, at a depth of aleutiana, but differs in having larger polyps that are 192 m. The distance between the red laser marks is 10 more fragile due to the thin body wall scales. cm. 268 Professional Paper NMFS 23 17. Plumarella hapala Cairns, 2011 (continued) Figure 13-19 A map of the Aleutian Islands showing the distribution of Plumarella hapala () in Alaska waters. Chapter 13 269 18. Plumarella nuttingi Cairns, 2011 Description (Adapted from Cairns, 2011a) Colonies Distribution Rare (Fig. 13-20). In Alaska – known consist of relatively few main branches arranged in a from only nine specimens collected from three locations plane from which smaller branchlets diverge in a bot- in the central Aleutian Islands, northeast of Great Sit- tlebrush fashion; the largest known colony is only 19 kin Island to southern Amchitka Pass. Elsewhere – not cm in height. Polyps occur on all sides of the branch- reported. lets with only 9–13 polyps/cm of branchlet length. The polyps are 3.0–3.6 mm in length and are slightly flared Habitat Occurs singly or in small patches on bedrock, and fleshy. The basal region (6–8 mm) of each branch- boulders, and cobbles at depths between 350 and 888 m. let is immersed in a thin coenenchymal membrane that contains scales. The axis is light brown to pale yellow; Photos A) A large portion of a dried P. nuttingi the coenenchyme is light orange to pink in life and light colony (USNM 1006331; paratype) collected north- orange when dried. east of Great Sitkin Island, central Aleutian Islands, at Each polyp is protected by eight opercular scales and a depth of 350 m. B) A close-up view of the same eight rows of body wall scales; the sclerite formula is specimen in photo A. C) A small P. nuttingi colony 7–8: 7–8: 6–7: 6–7. The tips of the marginal scales are (USNM 1134464; paratype; between the red laser pointed to spinose; the remaining body wall scales have marks) collected in southern Amchitka Pass, central a serrate distal edge. Coenenchymal scales are elliptical Aleutian Islands, at a depth of 888 m. A shortspine and flat with a granular outer surface. thornyhead (Sebastolobus alascanus) rests nearby. D) A close-up view of the same colony in photo C. The Remarks This species is distinctive in having large distance between the red laser marks in photos C and polyps and consequently a low number of polyps per D is 10 cm. unit area and a coenenchymal membrane around the bases of branchlets. It shares the bottlebrush branching pattern only with P. superba. 270 Professional Paper NMFS 23 18. Plumarella nuttingi Cairns, 2011 (continued) Figure 13-20 A map of the Aleutian Islands showing the distribution of Plumarella nuttingi () in Alaska waters. Chapter 13 271 19. Plumarella profunda Cairns, 2011 Description (Adapted from Cairns, 2011a) Colonies est dwelling species in the subgenus Dicholaphis (a sub- branch in a uniplanar, loose, alternate-pinnate man- genus no longer accepted) when it was first described, ner; colonies are delicate but quite flexible. The largest hence the species name. Only P. aleutiana has been doc- known specimen is only 21 cm in height and 14 cm in umented at greater depths. width but with 24 branch tips and 4th order branching. The axis is a light brown to pale yellow and often visi- Distribution Rare. In Alaska – known from only two ble under the thin coenenchyme. Polyps are 1.5–1.9 mm specimens collected at the same location in Adak Can- in length and occur on all sides of the branches with yon, central Aleutian Islands (Fig. 13-21). Elsewhere – 10–14 polyps/cm of branch length. The coenenchyme not reported. is light orange to pink in live specimens and in ethanol. Each polyp is protected by eight opercular scales and Habitat Occurs singly or in small but dense patches eight rows of body wall scales; the sclerite formula is (up to 10 colonies/m2) on siltstone at depths around 5: 5: 4: x (variable). All body wall scales are roughly 2514 m. rectangular in shape, with a straight, finely serrate distal margin; the outer surface of the body wall scales are smooth to finely granular. Opercular scales are elongate Photos A) The preserved (in ethanol) P. profunda ho- (0.47–0.86 mm in length), and somewhat triangular and lotype (USNM 1134074; whole colony except holdfast) pointed. The coenenchymal scales are elliptical to circu- collected in Adak Canyon, central Aleutian Islands, at a lar, 0.1–0.3 mm in diameter, and bear low granules on depth of 2514 m. B) A close-up view of the same speci- their outer surface. men in photo A showing the details and arrangement of the polyps. C) The same colony in photo A (at center), Remarks This species is unique among the Alaska in situ. D) A close-up view of the same colony in photo Plumarella in having rectangular body wall scales and C. The distance between the red laser marks in photos elongate opercular scales. It was thought to be the deep- C and D is 10 cm. 272 Professional Paper NMFS 23 19. Plumarella profunda Cairns, 2011 (continued) Figure 13-21 A map of the Aleutian Islands showing the distribution of Plumarella profunda () in Alaska waters. Chapter 13 273 20. Plumarella robusta Cairns, 2011 Description (Adapted from Cairns, 2011a) Colo- (Watling and Stone7) and ophiuroids. Small actiniarians nies consist of one main branch or several (up to four), and hydroids occasionally attach to bare branch axes. each branching as a uniplanar, alternate-pinnate frond; The coenenchyme is composed of high-magnesium cal- the largest colonies are 36 cm in height and 32 cm in cite (8.2 mol% MgCO3; senior author and S. D. Cairns, width. Branchlets are up to 10 cm in length. The axis unpubl. data). is fairly flexible and light brown in color. Polyps occur on all sides of the branchlets in a crowded manner with Distribution Locally common. In Alaska – known about 20 polyps/cm of branchlet length. Polyps measure from only seven specimens collected in the central Aleu- 1.4–1.8 mm in length and are cylindrical with a flared tian Islands (Amchitka Pass and south of Tanaga Is- tip. Colonies are light pink to orange in life, but light land) but video observations indicate that it is common orange to yellow when dried. where present (Fig. 13-22). Elsewhere – not reported. Each polyp is protected by eight opercular scales and eight rows of thick body wall scales; the sclerite formula Habitat Occurs in small and large patches on boul- is 4–5: 3–4: 3–4: several. Marginal scales are promi- ders and cobbles at depths between 115 and 1061 m. nently spinose and the distal spines bear seven or eight spiny, longitudinal ridges. Remaining body wall scales Photos A) A dried whole (except the holdfast) P. are also spinose or have a pointed distal margin. The robusta colony (USNM 1481801) collected in south- coenenchymal scales are elongate, thick, up to 0.6 mm ern Amchitka Pass, central Aleutian Islands, at a in length, and bear granular radiating ridges on their depth of 115 m. B) A close-up view of the same spec- outer surface. imen in photo A. C) The dried P. robusta holotype (USNM 1135992; whole colony) collected in northern Remarks The marginal spines in this species are simi- Amchitka Pass, central Aleutian Islands, at a depth of lar to those in P. echinata but differ in having ridges 711 m. D) The same colony in photo C prior to col- and thicker body wall scales. lection. The distance between the red laser marks is Associated fauna include caprellid amphipods 10 cm. 274 Professional Paper NMFS 23 20. Plumarella robusta Cairns, 2011 (continued) Figure 13-22 A map of the Aleutian Islands showing the distribution of Plumarella robusta () in Alaska waters. Chapter 13 275 21. Plumarella spicata Nutting, 1912 Description (Adapted from Cairns, 2011a) Colonies from all other Alaska Plumarella by the arrangement of branch in a uniplanar, quasi-dichotomous fashion or its polyps in an alternating biserial manner versus on all loose alternate-pinnate arrangement; branches are deli- sides of the branchlets. This arrangement was customar- cate and straggly; the largest known specimen is 27 cm ily used to distinguish the subgenus Plumarella whereas in height and 16 cm in width (USNM 1134073). The those species with crowded, randomly arranged polyps branch axes are a golden brown and visible through were placed in the subgenus Dicholaphis, although this the thin coenenchyme in live specimens. Polyps occur nomenclatural distinction is no longer observed. Based in an alternating biserial arrangement, usually with on molecular evidence, the two subgenera were recently about 12–14 polyps/cm of branch length. Polyps are synonymized (Cairns and Wirshing, 2018). slightly flared and 1.4–1.6 mm in length. The coen- enchyme is light orange in life and yellowish orange Distribution Uncommon. In Alaska – central and when dried. western Aleutian Islands (Adak Canyon to southeast Each polyp is protected by eight opercular scales and of Agattu Island) including Bowers Bank (Fig. 13-23). eight rows of thick body wall scales; the sclerite formula Elsewhere – not reported. is 5–6: 5–6: 4–6: 4–6. Opercular scales are elongate and pointed; marginal scales are prominently spinose with Habitat Occurs singly or in small patches on bed- the narrow spine constituting up to 70% of the length rock, siltstone, boulders, and cobbles in areas of mod- of the sclerite. Body wall scales have serrate distal mar- erate current, and at depths between 712 and 1913 gins. The coenenchymal scales are elliptical in outline m. This species co-occurs with P. echinata and Acan- and coarsely granular. thogorgia spissa in dense linear rows on bedrock and siltstone ridges in Adak Canyon (see photo E on next Remarks Plumarella spicata is easily distinguished page). Figure 13-23 A map of the Aleutian Islands showing the distribution of Plumarella spicata () in Alaska waters. 276 Professional Paper NMFS 23 Photos A) A fresh whole P. spicata colony (USNM the time of collection. D) A close-up view of the same 1134073) collected in northern Amchitka Pass, cen- specimen in photo C. E) A patch of P. spicata colonies tral Aleutian Islands, at a depth of 712 m. B) A dried on a siltstone ledge in Adak Canyon at a depth of 1745 branch of a P. spicata colony (USNM 1481803) collect- m. The distance between the red laser marks in photos ed in Adak Canyon, central Aleutian Islands, at a depth C and E is 10 and 20 cm, respectively. of 1746 m. C) A P. spicata colony (USNM 1134073) at Chapter 13 277 22. Plumarella superba (Nutting, 1912) Description (Adapted from Cairns, 2011a) Colonies Distribution Locally common and relatively wide- branch in a plane, consisting of relatively few main spread. In Alaska – Aleutian Islands north of Umnak Is- branches that bear numerous rigid and wiry branchlets land to southeast of Agattu Island including Petrel Bank in a characteristic bottlebrush arrangement. The largest (Fig. 13-24). There is a single record from deep water collected specimen is 26 cm in height and 32 cm in width, (1258 m) in Pribilof Canyon, eastern Bering Sea, col- but colonies up to about 50 cm in height and width have lected with a beam trawl by the U.S. Fisheries Steamer been observed in situ. The axis is woody in texture and Albatross in 1893. Given the geographical and bathy- black in color. Small polyps (only 1.0–1.3 mm in length) metric separation from all other records, we highlight occur on all sides of the branchlets in a crowded manner this record as suspicious, possibly a translocation by with 30–35 polyps/cm of branch length. The color of the sampling gear. Elsewhere – this is the only Alaska the coenenchyme is light to bright orange, occasionally Plumarella species found outside of Alaska. There is a creamy white in shallower-water specimens. single record (Institute of Marine Biology Institute FEB Each polyp is protected by eight opercular scales and RAS; Vladivostok, Russia; MIMB 16534) from the Sea eight rows of thick body wall scales; the sclerite formula of Okhotsk, Russia (Dautova, 2007). is 5–7: 4–6: 3–4: 1–2. Opercular scales are lanceolate with a serrate margin. Marginal scales are triangular Habitat In Alaska – typically occurs in patches but to prominently spinose with smooth (not keeled) inner occasionally singly on bedrock, boulders, cobbles, and surfaces. The remaining body wall scales are triangular pebbles at depths between 40 and 715 m. Elsewhere – with serrate margins; their outer face is either smooth or found at a depth of 29 m. longitudinally ridged. Coenenchymal scales are irregu- lar in shape but elongate and bear coarse granules. Photos A) A fresh whole P. superba colony (USNM 1481810) collected on the Adak Canyon shelf, central Remarks This species was originally described in Aleutian Islands, at a depth of 155 m. B) Fresh branch- the genus Primnodendron and later transferred to es of a P. superba colony (USNM 1011276) collected Thouarella, probably because of its bottlebrush-ar- near Little Tanaga Island, central Aleutian Islands, at a ranged branchlets. Cairns (2011a) ultimately trans- depth of 79 m. C) A fresh, large portion of a P. superba ferred it to Plumarella based on the lack of keels on colony (USNM 1011277) collected on Petrel Bank, cen- its marginal scales. It is one of only two species in the tral Aleutian Islands, at a depth of 40 m. D) A close-up genus that has bottlebrush-arranged branchlets (see Re- view of the same specimen in photo C. E) A P. superba marks of Plumarella nuttingi). It is also compared to colony (USNM 1135287) collected near Little Tanaga P. aleutiana in that account, and is keyed by Cairns Island at a depth of 100 m. F) A P. superba colony (2011a). (USNM 1134749) collected near Little Tanaga Island The microbial ecology of this species has been studied at a depth of 138 m. G) A P. superba colony (USNM from specimens collected in the central Aleutian Islands 1011277; at center) in a patch of P. superba on Petrel (Gray et al., 2011). Colonies often host ophiuroid as- Bank at a depth of 40 m. The distance between the red sociates. laser marks in photos E–G is 10 cm. 278 Professional Paper NMFS 23 22. Plumarella superba (Nutting, 1912) (continued) Chapter 13 279 Figure 13-24 A map of the Aleutian Islands showing the distribution of Plumarella superba () in Alaska waters. 280 Professional Paper NMFS 23 23. Primnoa pacifica Kinoshita, 1907 Description (Adapted from Cairns and Bayer, 2005) gence, a phenomenon whereby organisms that typically Colonies branch in a plane or are slightly bushy and dwell in the deep sea are able to exist in shallow-water dichotomously branched. The axis is rather stiff and ex- areas because of unusual oceanographic conditions tremely stout in large colonies. The height and width there. Emergence has been documented in glacial fjords are typically to 1–2 m but 5 m or more in exceptional to depths of 6 m in Holkham Bay and 9 m in Glacier specimens. Polyps are stout and large (5.0–6.5 mm in Bay, Southeast Alaska (Stone and Mondragon, 2018), length) and occur on all sides of the branches. The color and 12 m in Knight Inlet, northern Vancouver Island, of the axis is typically golden to dark brown but is of- British Columbia (McDaniel6). ten heavily calcified in larger colonies. The color of the Their tolerant temperature range is approximately coenenchyme is light pink or orange to bright orange 0.2–7.5°C (Matsumoto, 2007; Stone and Mondragon, and occasionally creamy white in the eastern Gulf of 2018) but they may tolerate temperatures near 10°C for Alaska (Bowie Seamount, British Columbia, to the Fair- several days (Stone and Mondragon, 2018). They are weather Ground; less common in the northern part of fairly euryhaline and able to tolerate salinities as low that range). as 26.9 for several days or more. They are preyed upon Each polyp is protected by eight opercular scales; one by calliostomatid snails (Akoya platinum), nudibranchs pair of basal scales; a variable number of unpaired, fu- (Tritonia tetraquetra), and the spiny red sea star (Hip- siform medial scales; and eight marginal scales. The ad- pasteria phrygiana) (Stone et al., 2015b). Associated axial side of the polyp is naked. Basal scales are massive, fauna of colonies residing in shallow water include the each bearing a robust horn-like distal spine. The medial shrimps Lebbeus groenlandicus, Eualus butleri, E. suck- scales occur on the abaxial and lateral sides of the pol- leyi, and E. townsendi. Several fish species, including the yps and are usually much smaller than the basal and goldeneye snailfish (Allocareproctus unangas), lay their buccal scales. There are five large marginal scales in the egg masses on red tree coral colonies (Busby et al., 2006). abaxial and lateral positions, and three much smaller The coenenchyme including the sclerites is composed marginals in the adaxial position. Opercular scales are of high-magnesium calcite (8.1–8.3 mol% MgCO3), lanceolate to tongue-shaped and bear multiple ridges on whereas the carbonate mineralogy of the skeletal axis their inner face. Coenenchymal scales are elongate and and holdfast are more variable, ranging from principally round in cross section and up to 1.5 mm in length. high-magnesium calcite (10.7–14.4 mol% MgCO3) to principally aragonite (senior author and S. D. Cairns, Remarks This robust species was described and illus- unpubl. data). trated by Cairns and Bayer (2005), who also provide a key to all species in the genus. Distribution Common, very abundant locally, and Primnoa pacifica, also known as “red tree corals,” widespread (Fig. 13-25). In Alaska – eastern GOA in- are the most ecologically important cold-water corals cluding the inside waters of Southeast Alaska, western in the North Pacific Ocean. Red tree corals are the larg- GOA including the inside waters of Prince William est and most geographically widespread corals in the re- Sound, Aleutian Islands (Samalga Pass to Buldir Reef), gion and provide important habitat for myriad species and a single record from Zhemchug Canyon in the east- including commercially important fish and crabs (Stone ern Bering Sea. Conspicuously absent from the Gulf of et al., 2015b). They are consequently the most studied Alaska Seamount Province but likely present on Dick- cold-water coral in the region and quite possibly the ens Seamount, the easternmost seamount in the Prov- world. Studies have been done on their ecology (Krieg- ince; see Primnoa pacifica var. willeyi (Hickson, 1915) er, 2001; Krieger and Wing, 2002; Du Preez and Tun- species account. Elsewhere – this species appears to be nicliffe, 2011; Stone et al., 2015b; Masuda and Stone, relatively common in northern British Columbia (Alas- 2015; Hartill et al., 2020), reproduction (Waller et al., ka border to west of Queen Charlotte Sound) where it 2014, 2019), growth (Andrews et al., 2002; Matsu- forms thickets and then uncommon south to northern moto, 2007; Williams et al., 2007; Aranha et al., 2014; Washington. In 2018, small patches of P. pacifica were Choy et al., 2020), genetics (Morrison et al., 2015), discovered just south of the Oregon–California border microbial ecology (Goldsmith et al., 2018), and the ef- and off Punta Gorda at depths between 362 and 582 m fects of ocean warming (Stone and Mondragon, 2018; (Everett5). There is a single confirmed record (USNM Johnstone et al., 2021) and ocean acidification (Rossin 57557) from off Point La Jolla in southern California et al., 2019). collected by the U.S. Fisheries Steamer Albatross in Red tree corals are one of only a few octocoral spe- 1904. The record is so geographically disjunct and in an cies worldwide documented to display deep-sea emer- area of atypical habitat for the species that we suspect Chapter 13 281 Figure 13-25 A map of the North Pacific Ocean showing the distribution of Primnoa pacifica () in Alaska waters. the specimen was translocated by a fishing vessel. In Photos A) Three large P. pacifica specimens collected the western Pacific Ocean the species ranges to the Sea in the eastern GOA. Left: a fresh major branch from of Japan including the Bay of Peter the Great, Russia, an irregular colony collected in Dixon Entrance at a and west of Honshu and Hokkaido Islands including depth of 320 m. The branch was damaged at the base the Shiribeshi Seamount in Japan (Matsumoto, 2005). by fishing gear. Center: a fresh bushy colony collected Kinoshita (1907) described the type specimen from in Dixon Entrance at a depth of 335 m. The colony was Sagami Bay (600 m depth) near Tokyo on the Pacific damaged at the base by fishing gear. Right: a fresh uni- Ocean side of Honshu Island, and Broch (1935) later planar but multi-fanned colony collected on the Fair- reported a specimen from the Sea of Okhotsk, Russia. weather Ground at a depth of 163 m. B) A large dried branch complex from a P. pacifica specimen (USNM Habitat In Alaska – generally occurs in thickets that 1481839) collected on Portlock Bank, western GOA, at are often extensive in the eastern GOA. Occurs singly a depth of 326 m. C) A close-up view of a fresh branch on glacial erratics also known as drop stones (i.e., boul- from a P. pacifica specimen (USNM 1116849) collected ders dropped as glaciers or icebergs melt away in the in Zhemchug Canyon, eastern Bering Sea, at a depth ocean). Grows on bedrock, boulders, and large cobbles of 520 m. D) A school of yellowtail rockfish (Sebastes in areas with moderate current. Also known to grow flavidus) feeding in a thicket of P. pacifica in Dixon En- on derelict fishing gear including steel trawl cables; such trance at a depth of 165 m. E) A yelloweye rockfish (S. occurrences may provide a timeline of settlement from ruberrimus) taking shelter near large P. pacifica colonies which age and growth can be estimated (Krieger, 2001). on the Shutter Ridge, eastern GOA, at a depth of 160 Found at depths between 6 and 640 m but video obser- m. F) A silvergray rockfish (S. brevispinis) resting on a vations from the central Aleutian Islands indicate that large P. pacifica colony on the Fairweather Ground at a this species may occur as deep as 899 m there (Stone, depth of 170 m. G) The white color-phase of P. pacifica 2014). Elsewhere – reported from bedrock and boulder on the Shutter Ridge, eastern GOA, at a depth of 203 habitats at depths between 12 m in Knight Inlet, north- m. H) A blue king crab (Paralithodes platypus) resting ern Vancouver Island, British Columbia, and 1250 m in on a P. pacifica colony in Tracy Arm in Holkham Bay, the Sea of Japan, Russia. Southeast Alaska, at a depth of 18 m. I) A close-up view 282 Professional Paper NMFS 23 of a Primnoa pacifica colony on the Shutter Ridge at juvenile (one-year old) walleye pollock feeding on eu- a depth of 192 m. J) A young-of-the-year walleye pol- phausiids (likely Thysanoessa raschii) that are swarming lock (Gadus chalcogrammus) (seen within the black around P. pacifica in Glacier Bay, Southeast Alaska, at circle) shelters in a Primnoa pacifica colony in Tracy a depth of 156 m. The distance between the red laser Arm, Holkham Bay, at a depth of 20 m. K) A school of marks in photos D–H, J, and K is 10 cm. Chapter 13 283 284 Professional Paper NMFS 23 24. Primnoa pacifica var. willeyi (Hickson, 1915) Description See Remarks. tions from Dickens Seamount in the Gulf of Alaska Sea- mount Province (Fig. 13-26). Elsewhere – there are two Remarks Variety willeyi, considered to be the same records from Chatham Sound, British Columbia, imme- species as P. pacifica, differs from typical colonies diately south of Southeast Alaska. in having more slender and often twisted polyps that are somewhat constricted at mid-height, thus seeming Habitat In Alaska – occurs singly or in small to support an overly large distal portion of the polyp. thickets with P. pacifica on bedrock and boulders at Also, they tend not to have medial scales on the lateral depths between 27 and 863 m. Elsewhere – no infor- sides of their polyps. The variety is described and illus- mation available. trated by Cairns and Bayer (2005). Associated fauna include pleustid and stenothoidid Photos A) A dried branch of a P. pacifica var. wil- amphipods (Watling and Stone7) and ophiuroids. The leyi colony (USNM 1115515) collected on Amchixtam skeletal axis is composed of high-magnesium calcite Chaxsxii in Amchitka Pass, central Aleutian Islands, (14.8 mol% MgCO3; senior author and S. D. Cairns, at a depth of 863 m. B) A close-up view of the same unpubl. data). specimen in photo A. C) The same colony in photo A just prior to collection. D) A close-up view of the same Distribution Uncommon but widespread. In Alaska colony in photo C. E) A P. pacifica var. willeyi colony – eastern Gulf of Alaska including the inside waters of (USNM1075478) collected on Dickens Seamount at a Southeast Alaska, western Gulf of Alaska, central and depth of 758 m. F) A close-up view of the same colony western Aleutian Islands (Amchitka Pass), and a single in photo E. The distance between the red laser marks in collected specimen along with several video observa- photos C–E is 10 cm. Figure 13-26 A map of the North Pacific Ocean showing the distribution of Primnoa pacifica var. willeyi () in Alaska waters. Chapter 13 285 286 Professional Paper NMFS 23 25. Primnoa wingi Cairns and Bayer, 2005 Description (Adapted from Cairns and Bayer, 2005) is often ridged. The coenenchymal scales are elongate, Colonies branch in a uniplanar or sometimes bushy and slender, and similar to the body wall scales in shape. dichotomous manner. The largest colonies approach 1 m in height and width. The color of the axial skeleton Remarks Primnoa wingi differs from other species in is golden brown, occasionally with thin black streaks. the genus by having inconspicuous body wall scales that Polyps are densely arranged on branches and most are are immersed in the fleshy polyp tissue, spatulate oper- directed downward, but some curve or corkscrew in dif- cular scales, and highly concave marginal scales. All ferent directions. Polyps are elongate, slender but fleshy, species are keyed in Cairns and Bayer (2005). large (up to 12 mm in length, twice the size of P. paci- Primnoa wingi is typically found in complex habi- fica polyps), and support a bulbous apical tip. The coe- tats and is an excellent indicator species of coral garden nenchyme is bright orange to reddish-orange in life and habitat. Primnoa wingi, along with Parastenella gym- a lighter orange when dried. nogaster, are the only two corals found both in Alaska Each polyp is protected by eight opercular scales; a waters and on the Emperor Seamounts in the central- variable number of unpaired, fusiform medial scales; west Pacific Ocean. and eight large marginal scales. The adaxial side of the Associated fauna include stenothoidid amphipods polyp is naked. Medial body wall sclerites are elongate (Watling and Stone7). The axial holdfast is composed and slender, occurring in a thin strip on the abaxial of high-magnesium calcite (8.5 mol% MgCO3; senior polypar side, and sunken into the polyp tissue; no defin- author and S. D. Cairns, unpubl. data). able basal scales are present. Marginal scales are quite large and convex, forming a prominent collar that enve- Distribution Uncommon. In Alaska – Amukta Pass lopes the lower part of the opercular scales. The oper- to southwest of Attu Island in the Aleutian Islands and cular scales are unique in shape, being elongate, medi- Zhemchug Canyon in the eastern Bering Sea (Fig. 13- ally constricted, and spatulate distally; their outer face 27). Elsewhere – there is a single record from Nintoku Figure 13-27 A map of the North Pacific Ocean showing the distribution of Primnoa wingi () in Alaska waters. Chapter 13 287 Seamount in the Emperor Seamounts, central-west Pa- 1115519) collected in northern Amchitka Pass, cen- cific Ocean. tral Aleutian Islands, at a depth of 711 m. B) A close- up view of a fresh large branch of a P. wingi colony Habitat Occurs singly or in small patches on bedrock (USNM 1116847) collected in Zhemchug Canyon, east- and boulders at depths between 217 and 914 m. Video ern Bering Sea, at a depth of 914 m. C) A close-up view observations from the central Aleutian Islands indicate of the same specimen in photo B. D) A bushy P. wingi that this species may occur as deep as 1280 m there colony observed on Amchixtam Chaxsxii in Amchitka (Stone, 2014). Elsewhere – found at a depth of 1156 m. Pass, central Aleutian Islands, at a depth of 1280 m. The distance between the red laser marks is 10 cm. E) A Photos A) A fresh whole P. wingi colony (USNM close-up view of the same colony in photo D. 288 Professional Paper NMFS 23 26. Thouarella cristata Cairns, 2011 Description (Adapted from Cairns, 2011a) Colonies Remarks Twenty-five species of Thouarella were branch primarily in a plane, consisting of a few main monographed and keyed by Taylor et al. (2013) but branches from which branchlets originate in a loose unfortunately T. cristata was overlooked in their ac- pinnate manner. The presence of an unidentified com- count. This species is however included in a recent mensal polychaete that forms a tube along the axis of account of all 35 known species in the genus (Cairns, the main branches may occasionally induce a third set 2021). Thouarella cristata is distinctive in having thick, of branchlets to occur, forming a bushy or even bottle- highly ridged body wall scales and a quasi-bottlebrush brush arrangement. The largest known colony is 50 cm arrangement of branchlets. in height. Polyps occur on all sides of the branchlets in In order to observe the ridging on the inner face of a crowded manner with 24–26 polyps/cm. Polyps are the marginal scales, which is diagnostic of the genus, the 1.7–2.5 mm in length and brooding polyps are com- polyps must first be bleached to disarticulate the scler- mon. The axis is pale yellow to dark brown; the coe- ites, and then the sclerites can be examined at 50–100× nenchyme is light orange to bright orange in life and power under a microscope. Dyeing the dry sclerites with fading to lighter orange in ethanol and when dried. a colored marker elucidates the feature. Each polyp is protected by eight opercular scales and Associated fauna include polynoid polychaetes eight rows of thick body wall scales; the sclerite formula (Watling and Stone7) and ophiuroids. The skeletal axis is 6–9: 6–7: 4–6: 3–4. Additional small elliptical scales is composed of high-magnesium calcite (12.7 mol% are also present on the adaxial surface. The marginal MgCO3; senior author and S. D. Cairns, unpubl. data). scales are pointed and both the outer and inner surfaces are prominently ridged (the innermost surface is keeled). Distribution Uncommon but locally abundant. In The coenenchymal scales are irregular in shape, usual- Alaska – Aleutian Islands (Adak Island to Kiska Island) ly elongate, and up to 0.9 mm in length. All scales are including Petrel Bank (Fig. 13-28). Elsewhere – not re- thick and robust. ported. Figure 13-28 A map of the Aleutian Islands showing the distribution of Thouarella cristata () in Alaska waters. Chapter 13 289 Habitat Occurs singly or in patches with the congener Bank, central Aleutian Islands, at a depth of 154 m. B) T. trilineata on boulders, cobbles, pebbles, and occa- A close-up view of the same specimen in photo A when sionally bedrock at depths between 91 and 744 m. it was fresh. C) A view (from above) of the holotype in situ. D) A view of the holotype during collection. The Photos A) The dried T. cristata holotype (USNM distance between the red laser marks in photos C and 1115600; large branch complex) collected on Petrel D is 10 cm. 290 Professional Paper NMFS 23 27. Thouarella trilineata Cairns, 2011 Description (Adapted from Cairns, 2011a) Colonies cristata, but differs in lacking the polychaete commen- branch primarily in a plane, consisting of a few main sal, and in having trilinear rows of branchlets, smaller branches from which branchlets originate in three rows, polyps, and non-spinose adaxial marginal scales. giving the colony a bushy to bottlebrush appearance; Ophiuroids are common associates of this coral, and commensal polychaetes are absent. The largest collected they are often the same color as the coenenchyme. colony is 23 cm in height. Polyps occur on all sides of the branchlets in a crowded manner with 35–40 pol- Distribution Uncommon but locally abundant. In yps/cm. Polyps are 1.4–1.7 mm in length and brooding Alaska – Aleutian Islands from Unalaska Island to polyps are absent. The axis is pale yellow; the coenen- Amchixtam Chaxsxii (Fig. 13-29). Elsewhere – not chyme is light orange to bright orange in life, fading to reported. lighter orange in ethanol and when dried. Each polyp is protected by eight opercular scales and Habitat Occurs singly or in patches with the congener eight rows of thick body wall scales; the sclerite formula T. cristata on boulders, cobbles, pebbles, and occasion- is 5–6: 4–6: 3–5: 1. Additional (6–10) small, randomly ally bedrock at depths between 97 and 1266 m. arranged elliptical scales are also present on the adaxial surface. The marginal scales are pointed and both their Photos A) A preserved (in ethanol) whole T. trilinea- outer and inner surfaces are prominently ridged (the in- ta colony (USNM 1009943; paratype) with associated nermost surface is keeled), however the adaxial margin- ophiuroids collected near Amlia Island, central Aleutian als are not spinose. Coenenchymal scales are irregularly Islands, at a depth of 198 m. B) A close-up view of the shaped, elongate, and up to 0.6 mm in length. same specimen in photo A with associated ophiuroid (indicated by the black circle). C) The preserved (in Remarks Thouarella trilineata is quite similar to T. ethanol) T. trilineata holotype (USNM 1010175; whole Figure 13-29 A map of the Aleutian Islands showing the distribution of Thouarella trilineata () in Alaska waters. Chapter 13 291 colony) collected near the Islands of Four Mountains, la trilineata colonies photographed near Great Sitkin eastern Aleutian Islands, at a depth of 202 m. D) A T. Island, central Aleutian Islands, at a depth of 175 m. trilineata colony photographed near Bobrof Island, cen- The distance between the red laser marks in photos D tral Aleutian Islands, at a depth of 160 m. E) Thouarel- and E is 10 cm. 292 CHAPTER 14 Family Sarcodictyonidae (Adapted from McFadden et al., 2022) Sarcodictyonids species (S. incrustans) (Broch, 1935) and one other spe- are octocorals without a skeletal axis and with polyps cies yet described. Sarcodictyon was formerly included connected basally by ribbon-like stolons that often fuse in the now defunct suborder Stolonifera. Madsen (1944) to form membranous expansions or mats. Polyps are referred to the Stolonifera as the most heterogeneous monomorphic and the anthocodiae are retractile into group within the Octocorallia given their membranous prominent calyces that are typically evenly distributed growth form. Sarcodictyon are uncommon in Alaska over the mat. Lateral budding of primary polyps often waters and consequently have a limited distribution in produces daughter polyps. The entire colony is cov- the eastern Gulf of Alaska and on a few seamounts in ered with a thin coenenchyme. Polyp sclerites are small the Gulf of Alaska Seamount Province (Fig. 14-1). The blunt rods with ornamented tubercles and are sparse group is represented by only two taxa: one found in or often arranged in longitudinal rows but not in the shallow water (15–238 m) and the other in deep water arrangement of collaret and points. Sclerites through- (2665–2881 m). Colonies encrust bedrock, and the bare out the rest of the colony are mostly stellate plates and skeletons of octocorals (Primnoa pacifica), barnacles, crosses. brachiopods, and hexactinellid sponges. Of the two known genera of the recently erected fam- All Sarcodictyonidae specimens collected in Alaska ily Sarcodictyonidae (McFadden et al., 2022), only Sar- waters should be retained and properly preserved for codictyon is found in Alaska waters with one known morphological and molecular studies. Figure 14-1 A map of the North Pacific Ocean showing the distribution of corals in the family Sarcodicty- onidae () in Alaska waters. Chapter 14 293 1. Sarcodictyon incrustans (Broch, 1935) Description (Adapted from Broch, 1935) Colonies 0.23 mm in length) derelict clubs. Sclerites are densely form crust-like coatings on substrates, including one packed everywhere but especially in the stolon where specimen that had completely overgrown an attached they are the largest (0.17–0.70 mm in length) and hydroid (Tubularia sp.) stalk and resembled a gorgo- consist of slender but quite strong, heavily thorned nian similar to Anthothela. The stolon plate forms a spindles (0.50–0.60 mm in length) and slender clubs continuous crust from which polyps are densely packed. (0.25–0.30 mm in length) with strong thorns on one The polyps are fully retractable into the coenenchyme side and weak on the other. and are 6 mm in height (without tentacles) and 3 mm in The contraction of the polyps begins with the place- width. The arrangement of the polyp sclerites are simi- ment of the tentacles over the mouth disc. Accordingly, lar to those of Clavularia armata but are smaller and the polyp wall folds in the middle, and the distal part have well-defined double rows with the middle scler- of the polyp is drawn into the lower half by inserting ites arranged en chevron and converging upwards. The the polyp wall. One could believe that a calyx was pres- double rows are just above the polyp base and form the ent; however, the indentation continues until the whole backs of the eight slightly raised longitudinal elevations polyp is retracted (or rather “indented”) into the coe- of the polyp wall. nosarc and can only be perceived as a swelling in the The aboral side of the tentacle trunk contains a stolon plate. platoon of longitudinally arranged sclerites (0.24–0.42 Alaska colonies form thin mats moderately populat- mm in length) that are irregularly rod-shaped with ed by scattered polyps or clusters of up to five polyps. low warts or thorns. In the upper part of the polyps, Polyps appear to be mostly non-retractile and up to 6 the sclerites are the same size as those in the tentacles, mm in length. Mats up to 0.5 m2 have been observed in but the warts are much more strongly developed. The situ that are contagiously distributed, giving the appear- polyp wall sclerites are 0.17–0.37 mm in length and ance of an infestation, and are strongly associated with the lower polyp sclerites are slightly s horter (0.17– red tree corals (Primnoa pacifica) with which they co- occur in shallow water of recently deglaciated fjords in Southeast Alaska (Stone and Mondragon, 2018). Color in life is white to light pink; color in ethanol is light yellow to creamy white. Remarks This species was originally described as Eva- gora incrustans by Broch (1935). Sarcodictyon in- crustans is a pioneer spe- cies in recently deglaciated fjords of Glacier Bay Na- tional Park and Preserve in Southeast Alaska (Stone and Mondragon, 2018). Distribution Uncommon. In Alaska – Muir and Johns Hopkins Inlets in Glacier Bay, Southeast Alaska (Fig. 14-2). Elsewhere – the type Figure 14-2 specimen was collected in A map of the North Pacific Ocean showing the distribution of Sarcodictyon incrustans () and the Sea of Okhotsk, Russia. Sarcodictyon sp. A (♦) in Alaska waters. A few specimens (USNM 57986 and 57987) were 294 Professional Paper NMFS 23 collected in Barkley Sound, southwestern Vancouver Photos A) Preserved (in ethanol) whole S. incrustans Island, British Columbia, at a depth of 15 m and were colonies (USNM 1407056) collected in Muir Inlet, Gla- identified as Sarcodictyon sp. but may likely be S. cier Bay, Southeast Alaska, at a depth of 21 m. The incrustans. colonies are growing on deepwater giant barnacles. B) A close-up view of the specimen at right in photo A. C) Habitat In Alaska – grows on hard rock, skeletons of Sarcodictyon incrustans colonies growing in Muir Inlet both live and dead deepwater giant barnacles (Chirona at a depth of 17 m. D) Sarcodictyon incrustans colonies evermanni) and brachiopods (Laqueus californicus), growing in Muir Inlet at a depth of 24 m. E) Sarcodicty- and the bare skeleton of red tree corals at depths be- on incrustans colonies growing in Muir Inlet at a depth tween 15 and 238 m. Elsewhere – the type specimen of 23 m. Note that some colonies have polyps that are was collected at a depth of 335 m. in various stages of retraction. Chapter 14 295 2. Sarcodictyon sp. A Description This species is known only from a sin- polyps were sparse and arranged mostly in rows that gle collected specimen that had completely overgrown were oriented in various directions. The color of the live the skeleton, including the holdfast of a hexactinellid colonies was light orange. sponge (Crateromorpha sp.). The sponge was attached to fractured bedrock and was a rare form in that the Distribution Rare. In Alaska – known from a single stalk was bifurcated approximately two-thirds from the collected specimen and video observations on Giaco- base. The large distal caps of the sponge were missing mini and Welker Seamounts in the Gulf of Alaska Sea- at the time of collection, and the stolon had completely mount Province (Fig. 14-2). Elsewhere – unknown. overgrown the tip of each stalk. The main stalk was 35 cm in length, the secondary stalk was 14 cm in length, Habitat In Alaska – attaches to the bare skeleton of and the mean circumference of the stalk was 5 cm. The the hexactinellid sponge Crateromorpha sp. and on entire colony had a surface area of 245 cm2. boulders at depths between 2665 and 2811 m. Calyces (polyps) are obviously conical and range up to 9 mm in diameter at the base and 6 mm in height, Photos A) A preserved (in ethanol) whole Sarcodic- are relatively sparse, but are distributed throughout tyon sp. A colony (USNM 1076622), broken into three the entire colony in clusters of up to seven individuals. pieces when collected on Giacomini Seamount at a The coenenchyme is quite thin and the dark brown depth of 2811 m. B) A close-up view of the same speci- color of the underlying sponge is apparent. The color men in photo A. C) The same colony in photo A with in life is a uniform peach-orange; the color in ethanol polyps fully retracted, photographed just prior to collec- is yellowish-orange. tion. D) A close-up view of the same colony in photo C We made observations of an additional colony (not with polyps fully retracted. E) Two Sarcodictyon sp. A collected) growing on a Crateromorpha sp. skeleton colonies photographed on Welker Seamount at a depth on Welker Seamount and two colonies (not collected) of 2665 m. F) A Sarcodictyon sp. A colony with polyps growing approximately 3 cm apart on the side of a boul- extended, growing on the bare skeleton of the hexac- der on Welker Seamount at 2665 m depth. These latter tinellid sponge, Crateromorpha sp., on Giacomini Sea- two colonies were irregularly shaped, very thin encrus- mount at a depth of 2808 m. The distance between the tations with dimensions of 11×5 cm and 8×2 cm. The red laser marks in photos C, E, and F is 10 cm. 296 Professional Paper NMFS 23 2. Sarcodictyon sp. A (continued) 297 CHAPTER 15 Superfamily Pennatuloidea Corals in the superfamily Pennatuloidea are known as of a single large polyp called the oozooid that through sea pens and sometimes referred to as sea whips. Pen- lateral budding gives rise to secondary polyps, most no- natuloideans that occur in Alaska waters are colonial, tably large autozooids and siphonozooids. The autozo- ahermatypic, gonochoric, and presumably broadcast oids are feeding polyps with eight well-developed feath- spawners. As in other cnidarians, sea pens are modular er-like tentacles (Fig. 15-1A), and the siphonozooids are organisms that can be viewed either as individuals with smaller polyps with highly reduced or no tentacles and many standardized and repeating feeding structures, or have a water circulation function (Fig. 15-1D). Together as colonies composed of several to numerous individu- with two other kinds of polyps found in a few species als. We will use the latter term here. of sea pens (mesozooids and acrozooids), pennatuloi- Conventional terminology is used to describe sea pen dean polyps are pentamorphic— that is, there are a total morphology (Fig. 15-1). A sea pen colony is composed of five kinds of polyps found throughout the range of Figure 15-1 Illustrations of four species of coral in the superfamily Pennatuloidea showing distinct morphological features: (A) Anthoptilum gowlettholmesae, (B) Acanthoptilim longifolium, (C) Ptilosarcus gurneyi, and (D) Veretillum ma- nillense. Illustrations were provided by Stephanie King (A and B), Laura Garrison (C), and Jessica Machnicki (D), California Academy of Sciences. 298 Professional Paper NMFS 23 pennatuloidean diversity (Williams et al., 2012). The lies currently recognized worldwide (Williams, 1995) body of a sea pen colony is composed of the rachis (the represented. We provisionally list 20 distinct taxa but distal portion of the colony that contains the polyps) caution that this group of corals is in desperate need of and the peduncle (the muscular proximal portion that taxonomic study. The current state of taxonomy for the anchors the colony into soft benthic sediments) (Fig. 15- group is poor and there are several taxa with unresolved 1, C and D). taxonomy (Anthoptilum spp., Balticina sp. A, Kopho- In a few species known as rock-inhabiting sea pens, belemnon sp., Protoptilum sp., Virgularia cf. bromleyi, the end bulb or basal portion of the peduncle forms and Umbellula sp.). Furthermore, there are several spe- a plunger-like expansion (Fig. 15-1A) that is capable cies with identifications based on only a few specimens, of adhering the colony to hard rocky substrate (Wil- including Anthoptilum murrayi Kölliker, 1880; Baltici- liams and Alderslade, 2011). Rock-inhabiting sea pens na californica (Moroff, 1902); Kophobelemnon sp.; Sta- are known to inhabit a depth range of 368 to 1969 m chyptilum superbum Studer, 1894; Virgularia bromleyi (Williams14) and have only recently been discovered in Kölliker, 1880; and Virgularia cf. glacialis. One taxon Alaska waters. Two of the 14 sea pen families possess (Pennatula sp.) is known only from video records. All well-developed lateral extensions called polyp leaves specimens collected in Alaska, except the obvious Bal- emanating from the sides of the rachis that can each ticina willemoesi (Kölliker, 1880) and Ptilosarcus gur- contain numerous autozooids, thus greatly increasing neyi (Gray, 1860), should be properly preserved for the number of feeding polyps on a single colony (Fig. taxonomic examination and zoogeographic study. 15-1, B and C). Pennatuloideans are found in all regions of Alaska Pennatuloideans in Alaska are not particularly speci- except the Arctic; none are found further north than ose but appear to be relatively diverse with 9 of 14 fami- Navarin Canyon in the eastern Bering Sea (Fig. 15-2). The records for most taxa represent the northernmost 14Williams, G. C. 2019. The rockpens - sea pens that attach to in the world. They are common throughout the Gulf rocky surfaces. Octocoral Research Center. Last revised Septem- of Alaska and Bering Sea, less common in the Aleutian ber 2019. [Available from https://researcharchive.calacademy. Islands, and rare in the Gulf of Alaska Seamount Prov- org/research/izg/Rockpen%20Forest%20and%20Detail.html, ac- cessed April 2021.] ince (Fig. 15-2) where they are known from only four Figure 15-2 A map of the North Pacific Ocean showing the distribution of corals in the superfamily Pen- natuloidea () in Alaska waters. Chapter 15 299 species (Anthoptilum murrayi, Anthoptilum sp. A, An- for formal description and a second one (Anthoptilum thoptilum sp. B, and Umbellula sp.) on five seamounts sp. B), observed on video footage only, is listed here to (Welker, Pratt, Quinn, Giacomini, and Der ickson Sea- highlight it for future collection. mounts). They are extremely eurybathic fauna in Alaska Many species are small, less than 50 cm in length and occupying depths between 3 m (Ptilosarcus gurneyi) and 5 cm in width, but a few attain lengths of 100 cm or 4656 m (Umbellula sp.). more, and B. willemoesi attains lengths of 3 m—mak- In Alaska and worldwide, most pennatuloideans ing it the largest sea pen in the world. Some species are inhabit soft-sediment areas of low-relief seafloor with locally abundant and are found in low- or high-density low to moderate water current. Colonies are anchored patches or “groves” and play a major role in construct- in place using a combination of hydrostatic pressure ing important benthic habitats in areas of otherwise and peristalsis to bury the peduncle into the sediment featureless seafloor (Brodeur, 2001; Stone et al., 2005). (Williams, 1995). However, unlike most other corals, The groves provide important refuge habitat for juve- they are sessile rather than sedentary. Several species nile fish and crabs and intercept and concentrate passing dislodged from the sediment, notably B. willemoesi and detritus, offal, and dislodged macroalgae (Stone et al., Ptilosarcus gurneyi, are able to rebury their peduncles 2005). Pennatuloideans appear to have few faunal as- and recover to an erect position (Malecha and Stone, sociates compared to other octocorals, but often serve 2009), and long-term observations of shallow-water as elevated feeding platforms for numerous species of populations indicate that individuals may move season- ophiuroids. Some nudibranch species from genera such ally especially with regard to depth distribution (senior as Tritonia and Armina are known to prey on pennatu- author, personal observ.). loideans. Species within the current range of fishing ac- Recently, a few species of pennatuloideans, known tivities in Alaska (approximately 1200 m depth), espe- as rock-inhabiting sea pens, have been discovered that cially those not capable of retracting partially or wholly attach directly to hard substrate via a peduncle that is into the sediment (e.g., Balticina spp.), are particularly modified to function like a suction cup (Williams and vulnerable to physical disturbance (Stone et al., 2005) Alderslade, 2011). The first rock-inhabiting sea pen and are common bycatch in trawl fisheries and stock from Alaska (Anthoptilum sp. A) is currently queued assessment surveys (Heifetz, 2002). 300 Professional Paper NMFS 23 Family Anthoptilidae (Adapted from Nutting, 1909 and Williams, 1995) Cor- oids that are numerous, elongate, and arranged biserial- als in this family have elongate, whip-like colonies with ly along either side of the rachis or in oblique rows situ- free polyps and without bifurcated calyces. The rachis is ated in two series along the rachis. Zooids are present without sclerites. The autozooids are disposed on raised on all surfaces; sclerites are absent except in the interior ridges or pads that are obliquely arranged along the ra- of the stalk; and the anthocodiae are non-retractile. Ad- chis. The proximal portions of adjacent autozooids are jacent autozooids may be united at their bases. Siphono- fused to some degree, forming small raised ridges, or zooids are minute but numerous on the rachis between joined only at the base. Polyp leaves are absent. the rows of autozooids. Species in the genus Anthoptilum have large autozo- 1. Anthoptilum grandiflorum (Verrill, 1879) Description Colonies are elongate and whip-like. Pol- lantic Ocean (off the Falkland Islands), Gulf of Mexi- yps are arranged in oblique rows and adjacent polyps co, and the North Atlantic Ocean (off the east coast of are fused at the base. Alaska specimens typically display North America). a strongly curved posture. The largest colonies are to about 50 cm in length. The colonies have a very large Habitat In Alaska – typically found in groves, some- peduncle which appears as an exposed disk at the sedi- times densely, rarely singly in soft sediment (silt and ment surface and is capable of partial retraction into sand) at depths between 600 and 2146 m. Video ob- the sediment. The color is purple to crimson except the servations indicate that the species ranges to depths of tissue along the axis and the peduncle is a paler orange at least 2511 m in the central Aleutian Islands (Stone, or crimson. 2014). Often found in mixed groves with the sea pen Anthoptilum sp. A and the chrysogorgiid Radicipes Remarks Anthoptilum grandiflorum occasionally stonei. Elsewhere – eurybathic, found in soft-sediment serves as an elevated perch for the deep-sea ophiuroid areas at depths between 188 and 3651 m. Asteronyx sp., presumably to suspension feed (Fujita and Ohta, 1988). The skeletal axis is composed of high-magnesium Photos A) Fresh whole A. grandiflorum colonies calcite (10.3 mol% MgCO ) with very small amounts (CAS 222962) collected in Navarin Canyon in the 3 of aragonite and low-magnesium calcite (senior author eastern Bering Sea at a depth of 1116 m. B) A fresh and S. D. Cairns, unpubl. data). whole A. grandiflorum colony (CAS 222949) collected north of Atka Island, central Aleutian Islands, at a Distribution Uncommon but locally abundant. In depth of 2146 m. C) An A. grandiflorum colony pho- Alaska – eastern Gulf of Alaska, central Aleutian Is- tographed north of Atka Island at a depth of 2220 m. lands including Bowers Bank that extends northward The distance between the red laser marks is 20 cm. into the eastern Bering Sea, and the canyons of the east- D) A grove of A. grandiflorum colonies north of Atka ern Bering Sea (Fig. 15-3). Elsewhere – cosmopolitan. Island at a depth of 2145 m. Also pictured is an An- North Pacific Ocean (off Washington, Oregon, Califor- thoptilum sp. A colony with an associated ophiuroid nia, and Hawaii), South Pacific Ocean (off Chile and (Asteronyx sp.). The distance between the red laser Palau), the Straits of Magellan, Antarctica, South At- marks is 40 cm. Chapter 15 301 Figure 15-3 A map of the North Pacific Ocean showing the distribution of Anthoptilum grandiflorum () in Alaska waters. 302 Professional Paper NMFS 23 2. Anthoptilum murrayi Kölliker, 1880 Description (Adapted from Kölliker, 1880) Colonies The third specimen (CAS 9459) was collected during are elongate, whip-like, and appear to be quite flexible the same expedition on Bowers Bank further west at in situ (see photo E). Alaska specimens range to about a depth of 1068 m. The identifier of that specimen is 90 cm in length. One specimen collected on Giacomini unknown. Eight additional specimens were observed in Seamount in the Gulf of Alaska Seamount Province at a situ on Giacomini Seamount. Elsewhere – there are pur- depth of 756 m (see photo B) has a total length of 48 cm ported records in the North Pacific Ocean from Hawaii, divided over three regions as follows: 1) the peduncle is off California, and near Honshu, Japan, but this is prin- 33 mm in length and 6 mm in diameter (contracted), 2) cipally a North Atlantic Ocean species where it is com- the naked stalk is 57 mm in length and 4 mm in diam- mon along the east coast of the United States, the Ba- eter, and 3) the polypoid section of the stalk is 39 cm in hamas, Caribbean Sea, and off the west coast of Africa. length with the diameter tapering to 2.7 mm. In larger specimens the tip of the rachis is flimsy and often tipped Habitat In Alaska – occurs singly or in low-density in situ, similar to that in Balticina sp. A. The rachis is patches in low-relief, soft sediment (sand, silt, and green sparsely populated with polyps arranged in oblique mud) habitat with light currents at depths between rows and adjacent polyps are separate and not fused at 1068 and 2266 m (Aleutian Islands). Seamount speci- the base. Colonies are not able to retract any part of mens occur on moderate slopes of sand, pebbles, and the rachis aside from the lower peduncle into the sedi- scattered cobbles and at depths between 744 and 762 ment. Color in situ is crimson to cranberry red, except m. Elsewhere – offshore banks and canyons at depths the tissue along the axis is a paler crimson to orange. between 558 and 2491 m. Specimens preserved in ethanol are pale orange, except in fresher specimens the polyps retain their natural color. Photos A) Preserved (in ethanol) whole Anthopti- lum murrayi colonies (USNM 16843) collected north Remarks The specimen collected on Giacomini Sea- of Amukta Pass, eastern Aleutian Islands, at a depth of mount is the first specimen documented from a sea- mount anywhere for the species. The reproductive biology of this species has been studied from speci- mens collected off the coast of Brazil in the southwest- ern Atlantic Ocean (Pires et al., 2009). Atlantic Ocean specimens are known to host the large ophiuroid As- teronyx sp. Distribution Rare (Fig. 15-4). In Alaska – known from only four specimens collected in the Aleutian Islands and on Giacomini Seamount in the Gulf of Alaska Seamount Province. The Aleutian Island speci- mens (USNM 16843) were collected north of Amukta Pass in the eastern Aleutian Islands at a depth of about Figure 15-4 2266 m by the U.S. Fisher- A map of the North Pacific Ocean showing the distribution of Anthoptilum murrayi () in ies Steamer Albatross and Alaska waters. identified by C. C. Nutting. Chapter 15 303 2266 m. B) A preserved (in ethanol) whole A. murrayi showing the detail and arrangement of the polyps on colony (CAS 233658) collected on Giacomini Seamount the tip of the rachis. E) The same colony in photo B pri- at a depth of 756 m. C) A close-up view of the same or to collection showing the flexibility of the rachis. F) specimen in photo B showing the detail and arrange- The opposite view of the same colony in photo B prior ment of the polyps on the central region of the rachis. to collection. The distance between the red laser marks D) A close-up view of the same specimen in photo B in photos E and F is 10 cm. 304 Professional Paper NMFS 23 3. Anthoptilum sp. A Description Colonies are elongate and have a bushy Remarks The two collected specimens represent a appearance in situ. They have a slight bend but are new species of rock-inhabiting sea pen, a recently dis- quite flexible and not rigid. The symmetry of the colony covered and unique group of pennatuloideans that at- is not bilateral but rather trilateral or circular. The larg- tach directly to rock rather than bury in soft sediment. est colony (USNM 1075802) measures 9.5 cm in height This new species is currently queued for formal descrip- and 7.6 cm in width (with the polyps fully extended). tion. It is morphologically similar to A. lithophilum The peduncle is short (< 1 cm) in preserved specimens Williams and Alderslade, 2011 known from off Cali- but considerably larger in situ when expanded. The pe- fornia including Rodriguez Seamount at depths of 669 duncle terminus is modified into a large (25 mm diam- to 700 m. eter) suction cup for attachment to rock and is light or- ange in color. The rachis consists of a short naked stalk Distribution Rare. In Alaska – known from only two (only 8% of the total length); the remaining rachis is specimens (USNM 1075802 and CAS 233656) collected sparsely populated with up to 28 large polyps (autozo- on Pratt and Quinn Seamounts in the Gulf of Alaska oids) that originate directly from the rachis. The polyp Seamount Province but observed on video footage else- stalk is long and tubular (up to 2.8 cm in length and 0.5 where on Pratt Seamount and on nearby Giacomini Sea- cm in width); the tentacles are up to 15 mm in length. mount (Fig. 15-5). Occurs singly, rarely two together. Compared to Anthoptilum sp. B, this species has far Elsewhere – unknown. fewer polyps that appear to be more circularly arranged than in a single plane. Polyp color in life is crimson Habitat Attaches to fractured bedrock and cobbles in and the coenenchyme elsewhere is a paler crimson to moderate- to high-relief habitat with moderate currents, light orange (almost translucent). The color in ethanol at depths between 898 and 1095 m. is brownish purple and the peduncle and axis are pale orange. Photos A) A fresh whole Anthoptilum sp. A colony Figure 15-5 A map of the North Pacific Ocean showing the distribution of Anthoptilum sp. A () in Alaska waters. Chapter 15 305 (CAS 233656) collected on Quinn Seamount at a depth A colony (CAS 233656) prior to collection. E) A lateral of 898 m. B) A close-up view of the same specimen in view of an Anthoptilum sp. A colony (USNM 1075802) photo A. C) A preserved (in ethanol) whole Anthopti- prior to collection. F) The view from atop the same lum sp. A colony (USNM 1075802) collected on Pratt colony in photo E. The distance between the red laser Seamount at a depth of 1095 m. D) An Anthoptilum sp. marks in photos D–F is 10 cm. 306 Professional Paper NMFS 23 4. Anthoptilum sp. B Description Colonies are elongate and whip-like but lum sp. A and while the two species are found on the with a bushy appearance in situ. They characteristically same seamounts, they are not found in close proximity have a slight bend. The largest colonies measure to about to each other. 22 cm in height and about 9 cm in width (with the polyps fully extended). The polyps start directly above the na- Distribution Uncommon. In Alaska – observed on ked, cup-shaped peduncle and there are approximately three seamounts (Welker, Pratt, and Giacomini) in the 20 polyp pairs opposite of each side of the axis with Gulf of Alaska Seamount Province (Fig. 15-6). Mostly the length of the polyps decreasing distally. Very large occurs singly but occasionally two or three together. polyps (the tubular polyp stalk is up to 3.8 cm in length) Elsewhere – unknown. originate directly from the rachis. The color in life is crimson; the peduncle is a paler crimson; and the axis is Habitat Attaches to bedrock, fractured bedrock, boul- light orange. ders, and cobbles in moderate-relief, moderate-current habitat, at depths between 797 and 1784 m (Welker Remarks This taxon also represents a new species of Seamount, 797–1126 m; Pratt Seamount, 917–1193 m; rock-inhabiting sea pen. The species is known from 53 Giacomini Seamount, 819–1784 m). observations made from video footage of the seafloor on Welker, Pratt, and Giacomini Seamounts in the Gulf Photos A) An Anthoptilum sp. B colony observed on of Alaska Seamount Province. The species cannot be Pratt Seamount at a depth of 1053 m. B) An Anthopti- formally described until a specimen is collected but until lum sp. B colony observed on Giacomini Seamount at that time we list it here with an informal description. a depth of 991 m. C) An Anthoptilum sp. B colony ob- It is clearly different from Anthoptilum sp. A and not served on Welker Seamount at a depth of 1032 m. The similar to any congeners in the northern North Pacific distance between the red laser marks in photos A–C is Ocean. It appears to be more common than Anthopti- 10 cm. Chapter 15 307 Figure 15-6 A map of the North Pacific Ocean showing the distribution of Anthoptilum sp. B () in Alaska waters. 308 Professional Paper NMFS 23 5. Anthoptilum sp. C Description Colonies are elongate, whip-like, and for the deep-sea ophiuroid Asteronyx sp., presumably very slender. The largest colonies measure to at least 95 to suspension feed (Fujita and Ohta, 1988). Anthopti- cm in height and to only 1.5 cm in width. The lower lum sp. C is preyed upon by an unknown sea star, ap- 20–27% of the rachis is without autozooids. The pe- parently a species of Solaster (Stone, 2014). The skeletal duncle is moderately enlarged (approximately twice the axis is composed principally of high-magnesium calcite width of the lower rachis), is typically 13–17% of the (10.4 mol% MgCO3) with very small amounts of ara- total colony length, and rarely has a slight bend at the gonite and low-magnesium calcite (senior author and S. base. The autozooids are arranged in two discrete rows D. Cairns, unpubl. data). on the same side of the rachis and appear to be nearly opposite but may be randomly arranged. The peduncle Distribution Uncommon but locally abundant. In is the only part of the colony that is retractable into the Alaska – north of Atka Island to Amchitka Pass in the sediment. Colonies bend in the current, particularly at central Aleutian Islands (Fig. 15-7). Elsewhere – un- the tip where there is little and often no flesh. The color known. of the polyps is dark purple to crimson, lighter orange along the axis, and the peduncle is a slightly darker or- Habitat In Alaska – typically found in groves, rarely ange. singly in soft sediment (silt and sand), at depths be- The taxonomy of this species is unresolved so spec- tween 1667 and 2707 m (Stone, 2014). imens should be carefully re-examined. It may be the same species as Anthoptilum sp. D but here we list them Photos A) Fresh whole Anthoptilum sp. C colonies separately due to the apparent difference in zoogeogra- (CAS 222973) collected northwest of Tanaga Island, phy, depth distribution, habitat, and gross morphology. central Aleutian Islands, at a depth of 2308 m. B) A grove of Anthoptilum sp. C and associated ophiuroids Remarks The species often serves as an elevated perch (Asteronyx sp.) northwest of Tanaga Island at a depth Figure 15-7 A map of the Aleutian Islands showing the distribution of Anthoptilum sp. C () in Alaska waters. Chapter 15 309 of 2250 m. The distance between the red laser marks land at a depth of 2329 m. D) A close-up view of an is 30 cm. C) A close-up view of an Anthoptilum sp. Anthoptilum sp. C colony tip and associated ophiuroid C colony (J2096-2-1; CAS # pending) with associated (Asteronyx sp.) in a grove northwest of Tanaga Island ophiuroid (Asteronyx sp.) in a grove north of Atka Is- at a depth of 2308 m. 310 Professional Paper NMFS 23 6. Anthoptilum sp. D Description Colonies are elongate, whip-like, and for the deep-sea ophiuroid Asteronyx sp., presumably very slender. The largest colonies are up to at least 66 to suspension feed (Fujita and Ohta, 1988). cm in height but only 1 cm in width. The lower 26– 32% of the rachis is without autozooids. The peduncle Distribution Uncommon but locally abundant. In is moderately enlarged (approximately twice the width Alaska – the continental slope of the eastern Bering Sea of the lower rachis), is typically 11–17% of the total including the large submarine canyons that incise the colony length, and often has a slight bend near the base. slope (Fig. 15-8). Elsewhere – unknown. The autozooids are arranged somewhat haphazardly in two rows on the same side of the rachis and appear to be nearly opposite but may be randomly arranged. The Habitat In Alaska – typically found in groves, rarely peduncle is the only part of the colony that is retract- singly in soft sediment (silt and sand), at depths be- able into the sediment. The color of the polyps is dark tween 230 and 1116 m. purple to crimson, lighter orange along the axis, and the peduncle is a slightly darker orange. Photos A) Fresh whole Anthoptilum sp. D colonies The taxonomy of this species is unresolved so spec- (CAS 222961) collected in Navarin Canyon, eastern imens should be carefully re-examined. It may be the Bering Sea, at a depth of 1116 m. B) A fresh whole An- same species as Anthoptilum sp. C but here we list them thoptilum sp. D colony (CAS 223765) collected south separately due to the apparent difference in zoogeogra- of Pribilof Canyon, eastern Bering Sea, at a depth of phy, depth distribution, habitat, and gross morphology. 244 m. C) A close-up view of a fresh Anthoptilum sp. D specimen (CAS 223756) collected south of Pervenets Remarks The species often serves as an elevated perch Canyon, eastern Bering Sea, at a depth of 465 m. Chapter 15 311 Figure 15-8 A map of the North Pacific Ocean showing the distribution of Anthoptilum sp. D () in Alaska waters. 312 Professional Paper NMFS 23 Family Balticinidae (Adapted from Nutting, 1909 and Williams, 1995) Cor- rachis between the oblique rows of autozooids. Species als in this family have elongate, whip-like colonies with in this family differ from those in the family Anthoptili- polyps that are retractile into calyces bearing two teeth. dae by having a rachis with sclerites, polyps with bifur- The rachis is bilateral throughout. The autozooids are cated calyces as opposed to none, and the presence of lateral only and disposed on raised ridges or pads that polyp leaves. are obliquely arranged along the rachis. The proximal After a very recent and extensive literature review of portions of adjacent autozooids are fused to some de- this group of sea pens, they were placed in the family gree, forming polyp leaves or raised ridges, or joined Balticinidae (= Halipteridae) and the genus Balticina (= only at the base. Spiculated calyces are present and are Halipteris) by Pérez et al. (2021). bifurcated. Siphonozooids are sparsely scattered on the 7. Balticina californica (Moroff, 1902) Description (Adapted from Kükenthal, 1915b) Col- Distribution Uncommon. In Alaska – eastern Gulf of onies are elongate and whip-like; the largest colonies Alaska including the inside waters of Southeast Alas- measure up to 82 cm in height and up to 2 cm in width. ka, central and western Gulf of Alaska, and the eastern The lower third of the colony is typically without pol- Aleutian Islands (Fig. 15-9). Elsewhere – found south yps. There are less than five polyps per row on each along the west coast of North America to northern Cal- polyp leaf compared to more than five polyps per row ifornia. in Balticina willemoesi. The lower peduncle is the only part of the colony that is retractable into the sediment. Habitat In Alaska – typically found singly but occa- The color of the polyps is orange to light orange with sionally in small patches or groves of up to 10–20 colo- purple/crimson highlights; the coenenchyme along the nies in soft sediment (sand and silt) at depths between rachis is a lighter orange. All colors fade when the colo- 35 and 823 m. Elsewhere – found in soft-sediment areas ny is preserved in ethanol. at depths between 60 and 2189 m. The taxonomy of this species is unresolved so speci- mens should be carefully re-examined. It is represented Photos A) A preserved (in ethanol) whole B. califor- in Alaska waters by only five specimens that may be the nica colony (USNM 1212106) collected in Monterey same species as Balticina sp. A. We list them separately Bay, northern California, at a depth of 1567 m. B) A until the specimens have been expertly re-examined. close-up view of the lower rachis and tip of a preserved (in ethanol) B. californica specimen (USNM 1206259) Remarks This species often serves as an elevated feed- collected in Monterey Bay at a depth of 60 m. ing platform for the common basket star (Gorgono- cephalus eucnemis) in shallower water. Chapter 15 313 Figure 15-9 A map of the North Pacific Ocean showing the distribution of Balticina californica () in Alaska waters. 314 Professional Paper NMFS 23 8. Balticina willemoesi (Kölliker, 1880) Description (Adapted from Kükenthal, 1915b) Col- Distribution Common, widespread and locally abun- onies are elongate and whip-like; the largest colonies dant. In Alaska – eastern Gulf of Alaska including the typically measure up to 150 cm in height but up to 3 m inside waters of Southeast Alaska, the central and west- in exceptional specimens (north of Amlia Island, central ern Gulf of Alaska, the Aleutian Islands west to at least Aleutian Islands) and up to 3 cm in width. There are Adak Island, and the eastern Bering Sea continental more than five polyps per row on each polyp leaf com- slope including the canyons (Fig. 15-10). Absent from pared to less than five polyps per row in Balticina cali- the seamounts in the Gulf of Alaska Seamount Province fornica. The lower peduncle is the only part of the colo- likely because all summits are deeper than the range of ny that is retractable into the sediment. The color of the this species. Elsewhere – found south along the west flesh is orange to light pink and lighter along the axis. coast of North America to southwest of the Columbia All colors fade when the colony is preserved in ethanol. River Basin, Oregon; west to the Sea of Okhotsk, Rus- sia; and the east side of Hokkaido, Japan. Five speci- Remarks The skeletal axis is composed principally of mens collected by the U.S. Fisheries Steamer Albatross high-magnesium calcite (9.8 mol% MgCO3) with very in 1906 from the East China Sea (west of the Osumi small amounts of aragonite and low-magnesium calcite Islands, Japan) at a depth of 660 m were identified by (senior author and S. D. Cairns, unpubl. data). The age G. Williams as B. cf. willemoesi. and axial growth rates have been estimated for this species in the Bering Sea and range up to 44 years and Habitat In Alaska – typically found in groves, some- 3.6–6.1 cm/year depending on age (Wilson et al., 2002). times quite dense, rarely singly, in soft sediment (sand This species is preyed upon by the nudibranch Tritonia and silt) at depths between 18 and 310 m. Elsewhere – diomedea (Malecha and Stone, 2009), and colonies are found in soft-sediment areas at depths between 69 m in capable of producing a bluish-green luminescence when the Sea of Okhotsk and 1204 m off Washington. This physically disturbed in situ. species has been observed in dense groves as shallow as Figure 15-10 A map of the North Pacific Ocean showing the distribution of Balticina willemoesi () in Alaska waters. Chapter 15 315 15 m depth in Howe Sound, southern British Columbia Canyon, eastern Bering Sea, at a depth of 236 m. C) (McDaniel6). A close-up lateral view (central region of the rachis) of the same specimen in photo B. D) A grove of B. wil- Photos A) A preserved (in ethanol) whole B. wil- lemoesi in Auke Bay, Southeast Alaska, at a depth of lemoesi colony (USNM 1206340) collected southwest 24 m. The distance between the red laser marks is 40 of the Columbia River Basin, Oregon, at a depth of cm. E) A close-up view of a B. willemoesi colony in that 529 m. B) A close-up lateral view (tip of the rachis) same grove showing the details and arrangement of the of a fresh B. willemoesi specimen collected in Pribilof polyps near the tip. 316 Professional Paper NMFS 23 9. Balticina sp. A Description Colonies are elongate, slender, and whip- Distribution Uncommon but widespread. In Alaska – like; the largest colonies measure up to 93 cm in height scattered throughout the Gulf of Alaska, the Aleutian but only up to 2 cm in width. The lower sixth of the Islands to Amchitka Pass, and the canyons of the east- colony is typically without polyps. Leaves are alternate ern Bering Sea (Fig. 15-11). Elsewhere – unknown. or random; one side of the rachis is bare. The lower peduncle is the only part of the colony that is retract- Habitat In Alaska – typically found in groves, some- able into the sediment. Colonies bend in the current, times densely (up to 5 colonies per m 2), but occasionally particularly at the tip where there is often little flesh, so singly in soft sediment (silt and sand) at depths between they are a very good indicator of current strength and 253 and 1154 m. Unlike other species of Balticina this direction. The color of the polyps is crimson to dark one is occasionally found in rockier, rougher habitats pink; the color of the coenenchyme along the axis and including silt-laden ledges on bedrock slopes or scarps. peduncle is a lighter pink or orange. Video observations indicate that the species ranges to depths of at least 1391 m in the central Aleutian Islands (Stone, 2014). Elsewhere – unknown. Remarks The taxonomy of this species is unresolved so specimens should be carefully re-examined. It may be the Photos A) A fresh whole Balticina sp. A colony (CAS same species as Balticina californica but here we list them 223760) collected in Dixon Entrance, eastern Gulf of separately due to apparent differences in zoogeography, Alaska, at a depth of 341 m. B) Balticina sp. A colo- depth distribution, habitat, and gross morphology. nies photographed on Portlock Bank, western Gulf of This species is used as an elevated perch, presumably Alaska, at a depth of about 300 m. C) The same colony to suspension feed, by the deep-sea ophiuroid Astero- in photo A just before collection. D) A close-up view of nyx sp. (Fujita and Ohta, 1988) in the deeper part of the same colony in photo C. The distance between the its range. red laser marks in photos B and C is 10 cm. Figure 15-11 A map of the North Pacific Ocean showing the distribution of Balticina sp. A () in Alaska waters. Chapter 15 317 318 Professional Paper NMFS 23 Family Kophobelemnidae (Adapted from Williams, 2005) Colonies are elongate rachis and not in oblique rows, or disposed on three sides and cylindrical or short, stout, and distinctively clavate of the rachis with a naked dorsal tract along the entire with the distal portion typically somewhat wider than the length of the rachis. Polyp leaves are absent. Anthocodiae rest of the colony. The distal end is either rounded and are mostly non-retractile and calyces are absent. Small knob-like or distinctively pointed. Autozooids number siphonozooids are numerous and arranged all along the between 2 and 50 and are arranged biserially along the rachis. Sclerites are three-flanged needles and spindles. 10. Kophobelemnon sp. Description This species is represented by only two Remarks Kophobelemnidae are typically Atlantic, specimens (USNM 57272) collected by the U.S. Fisher- Antarctic, and South Pacific species. There are several ies Steamer Albatross with a beam trawl in 1888 and records of Kophobelemnon hispidum Nutting, 1912 no- identified by F. M. Bayer. These two specimens should men dubium from near Hawaii and Japan and a single be re-examined to determine if they represent an un- specimen identified as Kophobelemnon sp. from British known species. Columbia at a depth of 706 m. The dimensions (not including autozooids) of the two colonies are 7.2 cm in height by 4.3 cm in width Distribution Rare. In Alaska – the two specimens were and 8.4 cm in height by 5.0 cm in width. The peduncles collected at the same location west of Prince of Wales appear to be the entire lower half of the colonies and Island in the eastern Gulf of Alaska (Fig. 15-12). This reach close to the first polyp. The large peduncle may species may be more common in the region than real- indicate that colonies bury deeply into the soft sediment ized since seafloor habitats below 1000 m have been they occupy with the first polyps close to the sediment/ seldom sampled or visually explored. Not to mention, water interface. The rachis is noticeably enlarged (2.6 that colonies are small and may bury deeply in the sedi- to 3.3 times larger than the peduncle) and apparently ment so would be difficult to collect with non-selective wrinkled, giving the appearance that it may become sig- sampling gear. Elsewhere – unknown. nificantly inflated in life. The distal section of the rachis Habitat The two known specimens were collected at is blunt with an abrupt pointed tip. a depth of 2869 m where the habitat likely consisted of One colony has two large polyps near the top that low-relief, soft-bottom abyssal plain. are obliquely positioned. The larger colony has three polyps—a single unpaired polyp proximally and a pair Photos A) A preserved (in ethanol) whole Kopho- of polyps distally. The latter specimen was broken in half belemnon sp. colony (USNM 57272) collected in the upon collection so it is not possible to determine if the eastern Gulf of Alaska (west of Prince of Wales Island) three polyps are possibly on three sides of the rachis. The at a depth of 2869 m. B) Another preserved (in ethanol) autozooids are quite large, between 80 and 100 mm in whole Kophobelemnon sp. colony (USNM 57272) col- total length, with long tentacles. Siphonozooids are nu- lected at the same location and depth as the colony in merous and densely packed on the rachis but not the pe- photo A. C) A close-up view of the top half of the speci- duncle or autozooids. The color of specimens preserved men in photo B. The distal end of the colony is on the in ethanol is a uniform light orange; presumably colonies right. D) A close-up view of the same specimen in photo are a darker orange color in life. B showing the detail of one of the polyps. Chapter 15 319 Figure 15-12 A map of the eastern Gulf of Alaska showing the distribution of Kophobelemnon sp. () in Alaska waters. 320 Professional Paper NMFS 23 Family Pennatulidae (Adapted from Nutting, 1909 and Williams, 2005) Colonies in the genus Ptilosarcus are particularly Colonies in the genus Pennatula are feather-shaped stout with kidney-shaped polyp leaves bearing sinuous with conspicuous large polyp leaves. Autozooids are ar- margins. Autozooids are crowded on the distal margins ranged in one or more rows along the margins of the of the polyp leaves. Anthocodiae are retractile into ca- polyp leaves. The anthocodiae are retractile into perma- lyces that are mostly spiculiferous, usually with one or nent spiculiferous calyces. The calyces are tubular and two indistinct to very conspicuous terminal teeth. Sipho- typically bear eight terminal teeth. Siphonozooids are nozooids are numerous in two longitudinal tracts along confined to the rachis between the polyp leaves. Meso- the rachis but not on the polyp leaves. Sclerites are scat- zooids may be present on the rachis or on the margin tered over the entire surface and not confined to the bor- of the polyp leaves opposite the autozooids. Sclerites ders of the leaves. Sclerites are three-flanged needles and are three-flanged needles of the calyces, inconspicuous spindles or longitudinally grooved oval-shaped plates three-flanged rods on the peduncle surface, and small and rods of the calyces and polyp leaves, and smooth, ovals in the interior of the peduncle. relatively large ovals of the peduncle. 11. Pennatula aculeata Danielssen, 1860 Description Colonies are bushy and similar to Pennat- the U.S. Fisheries Steamer Albatross (in 1888) in very ula phosphorea Linnaeus, 1758. The Alaska specimens deep water (2869 m). This species may be more com- measure up to about 9.5 cm in height and up to about mon than realized due to the lack of collection efforts in 3.5 cm in width but reportedly up to 18 cm in height very deep water in the Alaska region. Elsewhere – there elsewhere. Branches are opposite, largest at the center of are only a few records in the North Pacific Ocean (the the rachis, and start very close to the base of the rachis Channel Islands off central California). This is princi- near the sediment interface. The branches are up to 2 cm pally a North Atlantic Ocean species, ranging along the in length. Approximately the lower half of the colony east coast of North America from Newfoundland to consists of a moderately enlarged peduncle and a short North Carolina. section of the rachis without branches; thus the polyp leaves are almost in contact with the sediment in situ Habitat Several records consist of very large collec- (see photo D). The color in life is unknown but crimson tions indicating that this species likely forms groves. (see photo D) to light orange elsewhere in its geographic In Alaska – habitat consists of low-relief, soft sediment range and a uniform light orange in ethanol. (silt and sand) in areas of low current and at depths be- tween 2869 and 2972 m. Elsewhere – an extremely eu- Remarks Study of the behavior and distribution of rybathic species, occurring at depths between 110 and this species in the Gulf of Maine indicates that they 5710 m. have a contagious distribution with densities up to 8 colonies/m2 (Langton et al., 1990), and the gonadal Photos A) Preserved (in ethanol) whole P. aculeata morphology and gametogenesis have also been stud- colonies (USNM 43431) collected in Pribilof Canyon, ied from specimens collected there (Eckelbarger et al., eastern Bering Sea, at a depth of 2972 m. B) A close-up 1998). view of the same specimens in photo A. C) A zoomed-in view of the same specimens in photo A. D) A P. acu- Distribution Rare (Fig. 15-13). In Alaska – known leata colony observed in the Gulf of Maine, Northwest from only two locations: the eastern Bering Sea, sam- Atlantic Ocean, at a depth of about 65 m. The distance pled by the U.S. Fisheries Steamer Albatross (in 1890) between the red laser marks is about 10 cm. Photo D in very deep water (2972 m), and the eastern Gulf of is courtesy of R. Langton, National Marine Fisheries Alaska, west of Prince of Wales Island, also sampled by Service. Chapter 15 321 Figure 15-13 A map of the North Pacific Ocean showing the distribution of Pennatula aculeata () in Alaska waters. 322 Professional Paper NMFS 23 12. Pennatula sp. Description This taxon is known only from multiple been collected near Honshu Island (Japan) at a depth observations made from video footage. Colonies are uni- of 1097 m. planar, similar to Pennatula aculeata, but more delicate This taxon differs from all the species listed above and and with very different coloration. Colonies measure up all other pennatuloideans known to occur in Alaska and to about 8 cm in height and up to about 3 cm in width. thus likely represents an undescribed species. Specimens Most specimens have 16–18 pairs of branches that ap- are small and difficult to collect but future attempts of pear to be alternately arranged, with the largest at the collection should be a high priority. center of the rachis. A short section (approximately 1 cm) of the bare rachis is exposed above the sediment, keep- Distribution Uncommon. In Alaska – observed in ing the lowest branches in the water column. The color deep water north of Atka Island, central Aleutian Is- is uniform pure white which is very unusual for a genus lands (Fig. 15-14). Elsewhere – unknown. known for colorful and even phosphorescent species. Habitat Always found singly but in large, low-density patches. Habitat consists of low-relief, soft sediment Remarks The WoRMS Editorial Board (2022) lists (silt and sand) in areas of low current and at depths be- only 11 valid species of Pennatula worldwide, prin- tween 2239 and 2930 m. cipally in the North Atlantic, South Pacific and In- dian Oceans. Aside from our records of P. aculeata Photos A) A Pennatula sp. colony (lower right) with (see above), few Pennatula specimens have been col- polyps extended observed north of Atka Island, central lected in deep-water areas of the North Pacific Ocean. Aleutian Islands, at a depth of 2821 m. An unknown Specimens identified as Pennatula cf. phosphorea cali- brisingid star is at left. B) A Pennatula sp. colony, with fornica have been collected in British Columbia (2023 polyps retracted, observed north of Atka Island at a depth m depth) and in Monterey Bay and near the Channel of 2930 m. An unknown holothurian is burrowed above Islands, northern California (1997–2710 m depth). Ad- the sea pen. The distance between the red laser marks in ditionally, specimens of P. naresi Kölliker, 1880 have both photos is 10 cm. Chapter 15 323 Figure 15-14 A map of the Aleutian Islands showing the distribution of Pennatula sp. () in Alaska waters. 324 Professional Paper NMFS 23 13. Ptilosarcus gurneyi (Gray, 1860) Description Colonies are bushy and resemble a quill Distribution Common and locally abundant. In Alas- pen. Colonies measure to 50 cm or more in height and ka – eastern Gulf of Alaska including the inside waters 25 cm or more in width when fully inflated (i.e., ex- of Southeast Alaska, the central and western Gulf of panded). The anthocodiae are retractile into calyces Alaska, and the Aleutian Islands west to at least Bul- with two conspicuous terminal teeth. Siphonozooids dir Island (Fig. 15-15). Conspicuously absent from the are numerous but not arranged in clumps. Colonies are eastern Bering Sea. Also absent from the seamounts in capable of slow but complete withdrawal into the sedi- the Gulf of Alaska Seamount Province, likely because ment such that only the tip of the retracted colony is all summits are deeper than the range of this species. visible in a depression. The color of colonies in life and Elsewhere – found south along the west coast of North in ethanol is a uniform bright to medium orange. America to southern California. There are six, highly geographically disjunct records from the Philippines Remarks The skeletal axis is composed almost entire- (NMNH 43427) that should be re-examined. ly of high-magnesium calcite (12.0 mol% MgCO3) with very small amounts of aragonite and low-magnesium Habitat In Alaska – found in groves, sometimes calcite (senior author and S. D. Cairns, unpubl. data). densely, rarely singly in soft sediment (sand and silt) This species is preyed upon by the nudibranch Tritonia at depths between 3 and 274 m. Juveniles (<6 cm in diomedea that often spawns its egg masses (long rib- height) are often observed in shallower water (<10 m in bon-like strings that are the same color as the sea pens) depth) upslope from groves of adult colonies. Elsewhere around the base of the colony or immediately near it. – found in soft-sediment areas at depths of 16 to 293 m. Figure 15-15 A map of the North Pacific Ocean showing the distribution of Ptilosarcus gurneyi () in Alaska waters. Chapter 15 325 Photos A) Juvenile and adult Ptilosarcus gurneyi at a depth of 16 m. Photo credit: P. Malecha, National specimens collected in Glacier Bay, Southeast Alaska, at Marine Fisheries Service. D) Ptilosarcus gurneyi colo- a depth of 15 m. B) A P. gurneyi colony photographed nies held in a laboratory aquarium for study. E) A close- in Glacier Bay at a depth of 18 m. Several small Baltici- up view of the largest P. gurneyi colony in photo D. The na willemoesi are visible in the background. C) A grove distance between the red laser marks in photos B and C of P. gurneyi colonies near Auke Bay, Southeast Alaska, is 10 cm. 326 Professional Paper NMFS 23 Family Protoptilidae (Adapted from Nutting, 1909 and Williams, 2005) spiculated calyces which usually have three to eight Colonies are slender and elongate, never clavate, with terminal teeth. Siphonozooids are two to three times autozooids in one to three longitudinal rows along op- more abundant than autozooids and are mostly con- posite sides of the rachis. Individual autozooids may centrated near them. Sclerites of the tentacles, calyces, be scattered on the rachis between the lateral rows. rachis, and peduncle are three-flanged needles, spin- Polyp leaves are absent. Anthocodiae are retractile into dles, rods, and ovals. 14. Protoptilum sp. Description Colonies are elongate, whip-like, and Larger colonies are often used as an elevated feeding measure up to 44 cm in height. The rachis is quite slen- platform by the common basket star (Gorgonocephalus der, measuring to only about 6 mm in width, but rela- eucnemis) and also occasionally host smaller ophiuroids. tively stiff and flexible. Larger colonies bend slightly in the current. The lower 30–40% of the axis is without Distribution Uncommon but locally abundant. In polyps. Only the small, slightly bulbous peduncle is re- Alaska – Southeast Alaska (Glacier Bay) and the west- tractable into the sediment. The color of the flesh is very ern Gulf of Alaska from south of Prince William Sound light orange to creamy orange and slightly lighter along to south of Kodiak Island (Fig. 15-16). Elsewhere – un- the axis. known. Remarks The taxonomy of this species is unresolved Habitat In Alaska – typically found in small patches or and this taxon as we present it may represent more groves, sometimes densely, rarely singly in soft sediment than one species. Museum specimens should be care- (sand and silt) and at depths between 18 and 142 m. fully re-examined and all specimens collected during surveys and research expe- ditions should be retained for study. There are only seven valid species of Protopti- lum worldwide (WoRMS Editorial Board, 2022). They are principally North Atlantic Ocean taxa rang- ing along the east coast of North America from the Labrador Sea (Greenland) to Florida. Aside from our records of Protoptilum from relatively shallow-wa- ter areas of Alaska, North Pacific Ocean records in- clude: 1) P. orientale Nut- ting, 1912 from the Pacific Ocean side of Honshu, Ja- pan, at depths between 472 and 515 m, and 2) Protop- tilum sp. from Japan (148 Figure 15-16 m depth), British Columbia A map of the North Pacific Ocean showing the distribution of Protoptilum sp. () in Alaska (940 m depth), and Oregon waters. (137–1946 m depth). Chapter 15 327 Photos A) Large preserved (in ethanol) whole Prot- in Glacier Bay, Southeast Alaska, at a depth of 18 m. optilum sp. colonies (CAS 222956) collected in Chin- D) A grove of large and small Protoptilum sp. colo- iak Gully, western Gulf of Alaska, at a depth of 142 nies observed in Chiniak Gully at a depth of 140 m. m. B) Small preserved (in ethanol) whole Protoptilum E) Several large Protoptilum sp. colonies observed in sp. colonies (CAS 222955) collected in Two-headed Chiniak Gully at a depth of 130 m. The distance be- Gully, western Gulf of Alaska, at a depth of 127 m. tween the red laser marks in photos D and E is 10 C) Fresh small Protoptilum sp. specimens collected cm. 328 Professional Paper NMFS 23 Family Stachyptilidae (Adapted from Williams, 2005) Colonies are stout and lyces. Calyces are conspicuously toothed, generally with clavate or slender; firm or spongy in texture. Polyp two or three long terminal teeth. Siphonozooids are leaves are absent and the autozooids are arranged in mostly in oblique rows between the rows of autozooids oblique rows in two longitudinal series along the ra- and often have calyces formed by fan-shaped arrays of chis. Each row is situated alternately to a corresponding sclerites. Sclerites are three-flanged needles and spindles row on the other side of the rachis. Anthocodiae (about of the polyp calyces and rachis and oval- or rod-shaped three per row) are retractile into densely spiculated ca- plates in the peduncle. 15. Stachyptilum superbum Studer, 1894 Description (Original description from Studer, 1894) The single known Alaska specimen (USNM 1206296) The type specimen has the appearance of a wheat is much smaller, about 59 mm in height and about 5 mm “cob” and is 200 mm in height with the lower 58–60 in width at its widest point. The specimen appears to be mm of the stalk (the stem) lacking polyps. The colony complete but the intact axis has apparently protruded is cylindrical, a little swollen towards its base, and ter- through the base of the peduncle as a result of contrac- minating in an obtuse tip. The rachis is thicker than the tion of the flesh during long-term preservation in etha- stem; the dorsal part is very tightly covered with ca- nol. The color in ethanol is a uniform pale orange. lyces that are cylindrical and long in shape (5–6 mm) with openings that cover the bases of the superimposed Remarks Four of the five genera in the family Sta- calyces. The calyces are arranged in 34 (from base to chyptilidae are found in the Pacific Ocean. Stachyptilum tip) slanted rows of four or five on each side. The ca- is a small, obscure genus of only five species, including lyces are armed with long, spiniform sclerites, which two of uncertain status. project a little beyond the opening and form two or Distribution Rare. In Alaska – known from only a three prominent teeth. The ventral part of the rachis single specimen (USNM 1206296) collected on Petrel contains a narrow section bare of polyps and a deep Bank in the central Aleutian Islands (Fig. 15-17). Else- furrow runs along its entire length. On both sides of where – occurs along the Pacific Coast of the Americas this furrow the ventral part is covered with small zo- (northern Oregon south of the Columbia River Basin, oids, which are tubular and project a little on the sur- Channel Islands and Monterey Bay in California, and face; they have at their base a crown of sclerites. The central Mexico to Chile). The type specimen was col- axis is cylindrical, calcitic, elastic in consistency, and lected off the coast of Panama at a depth of 384 m. ends in a thin net towards the base. The color of the polyps is purple-brown with a white rachis. Habitat In Alaska – likely soft-bottom (sand and (Adapted from Nutting, 1909) Colonies are to up to silt) habitat at a depth of 388 m. Elsewhere – found at 170 mm in height with the lower 81 mm of the stalk depths between 48 and 1238 m; a single specimen was without polyps. There is a relatively prominent terminal collected from an area of green mud in Monterey Bay bulb and a swelling above the bulb. The ventral surface at a depth of 69 m and Nutting (1909) reports another of the rachis has a deep, even, undulating grove. Polyps specimen collected in Monterey Bay at a depth of 48 almost completely surround the rachis. Calyces are te- m. Several of the deeper-water collections with trawls rete, rather slender (3.5–5.0 mm in length by 1.25 mm consisted of many specimens, indicating that this species in width) and in four rows forming an oblique series. has a patchy distribution and possibly even forms small The autozooids are dark and well defined and present groves. in a row on either side of the ventral groove. They are particularly conspicuous distally, and siphonozooids ap- Photos A) A preserved (in ethanol) whole Stachyp- pear in small rows lateral from the main rows and in tilum superbum specimen (USNM 1206296) collected between the calyces on the dorsal side of the rachis. The on Petrel Bank, central Aleutian Islands, at a depth of sclerites are large, needle-like, and placed longitudinally 388 m. Note that the axis has protruded through the on the walls of the calyces. The ventral groove is devoid peduncle. B) A close-up view of the same specimen in of sclerites, in marked contrast to the rest of the rachis. photo A. Chapter 15 329 Figure 15-17 A map of the Aleutian Islands showing the distribution of Stachyptilum superbum () in Alaska waters. 330 Professional Paper NMFS 23 Family Umbellulidae (Adapted from Williams, 2005) Colonies have very large peduncle. Sclerites are three-flanged or round spindles, polyps radiating from a cluster (or crown) at the end rods, ovoid rods, and needles. of a long, slender stalk. Polyp leaves are absent and the Umbellula is the only genus in the family but there autozooids are restricted to the crown of 1–40 polyps. are 24 valid species recognized worldwide (WoRMS Anthocodiae are non-retractile and there are no calyces. Editorial Board, 2022). Umbellula appear to be wide- Siphonozooids are present at the base of the autozo- ly distributed in the northern North Pacific Ocean but oids and below the crown on the upper part of the stalk. only a few specimens have been identified to species and Sclerites are present in only a few species and if so are seemingly inaccurately. distributed in the tentacles, wall of the autozooids, and 16. Umbellula sp. Description Colonies are elongate, whip-like, and America), Gulf of Mexico, Caribbean Sea, South At- have a very narrow but rigid axis. The peduncle is lantic Ocean (off the east coast of South America), the slightly enlarged and is only about 15–20% of the col- Southern Ocean, and South Pacific Ocean (New Zea- ony length. Colonies may have between 3 and 20 large land). The species is also reported in the North Pacific polyps terminally on the crown, and there seems to be Ocean off California and Oregon. Umbellula ioma Nut- little relationship between the number of crowns and ting, 1909 (an unaccepted species according to WoRMS colony height. The anthocodiae with extended polyps Editorial Board [2022]) is reported from the Sea of Ok- may reach 5 cm in length. The largest colonies are up to hotsk, Russia. 35 cm in height and the crown is up to 10 cm in width. The color in life is light orange to white with purplish Habitat In Alaska – occurs singly in soft sediment (silt polyps and often crimson tentacles; the color in ethanol and sand) at depths between 840 and 4656 m (the deep- is dark golden brown to light brown. est record in the North Pacific Ocean). Video observa- tions indicate that the species ranges at depths between Remarks The taxonomy of this species is unresolved 2546 and 2947 m in the central Aleutian Islands (Stone, and it may represent one or more undescribed species or 2014). This species is often found in mixed groves with may be the cosmopolitan Umbellula lindahli Kölliker, the sea pens Anthoptilum grandiflorum, Anthoptilum sp. 1875. Further examination of all specimens from the re- C, and the chrysogorgiid Radicipes stonei. Elsewhere – gion is warranted. extremely eurybathic. Umbellula species reported from Associates include the large amphipod Amathillopsis the northern North Pacific Ocean range at depths be- annectens (Watling and Stone7) that clings to the stalk tween 352 and 4062 m. (see photos F and G). A large, solitary hydroid, possibly Branchiocerianthus imperator reported from very deep Photos A) A preserved (in ethanol) whole Umbellula water (3422 m) in the region of Japan, was observed at sp. colony (CAS 222947) collected south of the Trinity several locations on Derickson Seamount and the sur- Islands, western Gulf of Alaska, at a depth of 840 m. B) A rounding abyssal plain in the western Gulf of Alaska close-up view of the crown on the same specimen in pho- and can easily be confused with Umbellula (see photo to A. C) A preserved (in ethanol) whole Umbellula sp. I). colony (USNM 1081174) collected on Derickson Sea- mount at a depth of 4656 m. D) A close-up view of the Distribution Uncommon but widespread (Fig. 15-18). colony terminus on the same specimen in photo C. The In Alaska – eastern and western Gulf of Alaska, central colony has three large polyps; numerous siphonozooids Aleutian Islands including Bowers Ridge that extends are visible at the base of the crown. E) An Umbellula sp. northward into the eastern Bering Sea, and Giaco- colony observed north of Atka Island, central Aleutian mini and Derickson Seamounts in the Gulf of Alaska Islands, at a depth of 2840 m. The distance between the Seamount Province. Elsewhere – North Pacific Ocean red laser marks is 20 cm. F) A close-up view of the same records of Umbellula range from British Columbia to colony in photo E. Note the two large amphipods, Am- Oregon. Umbellula lindahli appears to be cosmopolitan athillopsis annectens, clinging to the stalk. G) A zoomed- with specimens reported principally from the North At- in view of the same colony in photo E. H) An Umbellula lantic Ocean (Baffin Bay and off the east coast of North sp. colony observed north of Atka Island at a depth of Chapter 15 331 about 2830 m. The distance between the red laser marks tor, at a depth of 4568 m on Derickson Seamount. The is 10 cm. I) A large, solitary hydroid, possibly B. impera- distance between the red laser marks is 20 cm. 332 Professional Paper NMFS 23 16. Umbellula sp. (continued) Figure 15-18 A map of the North Pacific Ocean showing the distribution of Umbellula sp. () in Alaska waters. Chapter 15 333 Family Veretillidae (Adapted from Williams, 1995) Colonies are cylindrical, chis has a single longitudinal furrow or V-shaped region capitate, clavate, or elongate with an erect rachis not devoid of polyps. Anthocodiae are retractile directly lying on the substrate. Polyp leaves are absent. The au- into the rachis; there are no calyces. Siphonozooids are tozooids are without calyces and distributed evenly on densely positioned between the autozooids. Sclerites are all sides of the rachis or the proximal portion of the ra- present but may be absent in the polyps. 17. Cavernularia vansyoci Williams, 2005 Description (Adapted from Williams, 2005) Colonies Remarks Cavernularia is a relatively large genus with are clavate and the largest colonies measure up to 57 19 species currently recognized worldwide (WoRMS mm in height and up to 37 mm in width. The stalk or Editorial Board, 2022). Most species are found in the peduncle widens noticeably about halfway up the col- Indo-Pacific region and Atlantic Ocean including the ony. The color of the preserved holotype is pale orange Mediterranean Sea. Only two other species are known to creamy orange throughout, but live specimens are from the North Pacific Ocean: Cavernularia habereri likely light orange. Moroff, 1902 and Cavernularia glans Kölliker, 1872, The polyps of the preserved holotype are all complete- both reported along the east coast of Asia. ly retracted. They are flush with the surface of the rachis This species is known from only two collected speci- and nowhere do they extend past the surface. Calyces are mens. Colonies are quite small and inconspicuous so absent. The polyps and polyp walls contain minute scler- they might be easily overlooked when observed in situ ites, similar to the sclerites of the superficial coenenchyme and rarely retained when encountered by bottom trawls, and peduncular interior. As in the coenenchymal tissues, a major source of collected specimens in Alaska. these sclerites can only ac- curately be detected at mi- croscope magnifications at 400× or higher. The sclerites are smooth spindles, rods or ovals, and are all colorless. The sclerite complement is composed entirely of very small ovals (0.003–0.007 mm in length). The shape of these sclerites is only ac- curately discernable at mi- croscopic magnifications of 400× or more. They vary in shape from rounded-recti- linear to oval, more-or-less round, or elliptical. Some sclerites are irregular in shape. These minute scler- ites are relatively dense in all parts of the colony exam- ined, including the coenen- chyme of the surface of the rachis and peduncle, as well as the interior of the colony, but seem to be less densely Figure 15-19 distributed in the anthoco- A map of the Aleutian Islands showing the distribution of Cavernularia vansyoci () in Alaska diae and polyp walls. waters. 334 Professional Paper NMFS 23 Preserved Cavernularia vansyoci specimens can eas- Photos A) The preserved (in ethanol) C. vansyoci ho- ily be mistaken for the soft coral Alcyonium sp. with- lotype (CAS 168894; whole colony) collected north of out careful examination to confirm the presence of si- Atka Island, central Aleutian Islands, at a depth of 94 phonozooids between the autozooids and to verify that m. The whole colony was cut longitudinally into two the lower part of the colony is a peduncle and not a halves. External and internal views of both halves are holdfast. at left and right, respectively. The figure is reprinted with permission from Williams (2005). B) A preserved Distribution Rare (Fig. 15-19). In Alaska – known (in ethanol) whole C. vansyoci colony (USNM 100721) only from the Aleutian Islands, north of Yunaska Island collected north of Yunaska Island, eastern Aleutian Is- to off the northwestern tip of Atka Island (the type lo- lands, at a depth of 126 m. The widened top of this col- cality). Elsewhere – not reported. ony was likely compressed somewhat laterally during preservation. Habitat In Alaska – soft-sediment (likely coarse sand and fine pebbles) habitat at depths between 94 and 126 m. Chapter 15 335 Family Virgulariidae (Adapted from Williams, 2005) Colonies in the genus Anthocodiae are retractile into the bulbous fleshy base Virgularia are long, slender, and often vermiform or of the polyps, thus forming calyx-like mounds. Siphono- more stout, robust, and rigid. The calcareous axis ex- zooids are sparsely distributed on the polyp leaves be- tends beyond the tip of the rachis in some species. Polyp low the free parts of the autozooids or more commonly leaves are relatively short, and often congested or with on the rachis between polyp leaves. Sclerites in Virgu- sections of bare rachis between adjacent leaves. Tubular laria are virtually absent except for minute oval bodies autozooids are fused for most of their length to form in the interior of the peduncle. relatively thin polyp leaves, 3–100 autozooids per leaf. 18. Virgularia bromleyi Kölliker, 1880 Description The original description by Kölliker Remarks Known from five specimens (USNM 43778) (1880) is rudimentary and based on a single fragment- collected by the U.S. Fisheries Steamer Albatross with ed colony. The polyps were described as nearly sessile a beam trawl in 1890 and later identified by E. Deich- with very small pinnules. The pinnules are nearly oppo- mann. The specimens were collected at a single location site with each about 4 mm apart. The rachis has lateral in deep water (1889 m) in the eastern Bering Sea, north zooids disposed at the base of the leaves in one single of Yunaska Island in the eastern Aleutian Islands. row of three individuals. The sclerites are very small or- dinary needles and are scantily distributed in the stalk, Distribution Rare. In Alaska – the eastern Bering Sea rachis, and the polyp tentacles. off the island arc slope of the Aleutian Islands (Fig. 15- The original description of 13 specimens of Halis- 20). Elsewhere – largely unknown. The holotype was ceptrum cystiferum Nutting, 1909 (now synonymized collected southeast of Honshu, Japan, at Challenger sta- with Virgularia bromleyi) indicates the largest colonies tion 235, and Nutting’s (1909) specimens were collected are to 120 mm in length with the lower 65 mm of the at two locations (U.S. Fisheries Steamer Albatross sta- stalk without leaves. The terminal bulb is a prominent tions 4541 and 4514) near Monterey Bay, California. bladder-like expansion and the ventral side of the rachis has a distinct groove. The pinnae are very short, and so Habitat In Alaska – low-relief, low-current habitat closely set that the polyps appear to be attached directly consisting of silt and sand in deep water (1889 m). Else- to the stem. There are approximately 32 pairs of pin- where – the holotype was collected in an area of mud at nae including the rudimentary ones. There are four or a depth of 1033 m and the California specimens were five large (compared to the pinnae themselves) polyps collected at depths of 839 and 1114 m. per pinna and they appear to be only partially retract- able. Zooids were noted as absent. Sclerites were not ob- Photos A) Preserved (in ethanol) mostly whole V. served by Nutting, but he noted that if present they were bromleyi colonies (USNM 43778) collected north of rare and very small—an observation that prompted him Yunaska Island (central Aleutian Islands), eastern Ber- to place his “new species” in the genus Halisceptrum. ing Sea, at a depth of 1889 m. B) A close-up view of the None of the five known Alaska specimens (USNM same specimens in photo A. 43778) appear to be intact but the largest colony is likely about 150 mm in length. The color in ethanol is uniform light orange. 336 Professional Paper NMFS 23 18. Virgularia bromleyi Kölliker, 1880 (continued) Figure 15-20 A map of the North Pacific Ocean showing the distribution of Virgularia bromleyi (), V. cf. bromleyi (♦), and V. cf. glacialis (▲) in Alaska waters. Chapter 15 337 19. Virgularia cf. bromleyi Background We synonymize this taxon with Virgu- should be carefully re-examined along with other re- laria bromleyi tuberculata reported from the Pacific cords of Virgularia sp. (especially the deeper records) Ocean coast of Honshu, Japan, at 300 m depth (iden- from northern Washington, northern California (13– tified by E. Deichmann), and from British Columbia 839 m depth), and southern California (25–1038 m at depths of 20 to 25 m (identified by F. M. Bayer in depth), and compared to the records of V. bromleyi and 1978). These four records are listed in the Smithsonian V. cf. glacialis from Alaska. National Museum of Natural History database. Subspe- cies tuberculata is an invalid designation according to Distribution Uncommon. In Alaska – reported from the WoRMS Editorial Board (2022) and V. tuberculata the inside waters of Southeast Alaska and through the Marshall, 1883 is strictly a North Atlantic Ocean spe- Gulf of Alaska to the eastern Aleutian Islands (Fig. 15- cies. We suspect that the four specimens from Japan 20). Elsewhere – British Columbia and possibly south to and British Columbia are the same species as four speci- southern California. mens from Alaska identified as Virgularia but otherwise with unresolved taxonomy. These specimens appear to Habitat In Alaska – areas of sand, silt, and pebbles have similar morphology to V. bromleyi although they at depths between 20 and 472 m. Elsewhere – areas of are considerably larger and occupy a much shallower rock and grey sand, green mud, and soft-brownish shale bathymetric range. at depths between 20 and 1038 m. Description Colonies are whiplike and measure up Photos A) A preserved (in ethanol) whole Virgularia to 54 cm in length. Colonies are slender but with long cf. bromleyi colony (USNM 57267) collected southeast branches when extended in situ; width in preserved of Sitkalidak Island, western Gulf of Alaska, at a depth specimens is up to 15 mm but up to 35 mm when ex- of 110 m. B) A close-up view of the rachis and polyp tended in situ. There are up to 66 polyp leaves that are leaves in the same specimen in photo A. C) A close-up obliquely set or nearly opposite and tapered both proxi- view of the distal section of the rachis and the peduncle mally and distally. Proximal polyp leaves closest to the (right) in the same specimen in photo A. D) A Virgularia sediment interface are rudimentary and without polyps. cf. bromleyi colony (USNM 57976) photographed in Fully developed leaves have five polyps that are gen- Indian Arm, Burrard Inlet, British Columbia, at a depth erally directed laterally. There is a prominent, distinct of 25 m. E) A close-up view of the same colony in photo groove on the ventral side of the axis. The color in life D. F) A Virgularia cf. bromleyi colony (CAS 163735) is a creamy or translucent white, almost iridescent; the photographed in Sea Otter Sound, Southeast Alaska, at color in ethanol is a uniform light orange. a depth of 22 m. Reprinted with permission from Barr and Barr, 1983. G) A Virgularia cf. bromleyi colony Remarks This taxon includes four shallow-water re- (USNM 57976) photographed in Indian Arm, Burrard cords from Alaska with unresolved taxonomy. These Inlet, British Columbia, at a depth of 25 m. Photos D, E, specimens likely represent an undescribed species but and G are courtesy of Neil McDaniel. 338 Professional Paper NMFS 23 19. Virgularia cf. bromleyi (continued) Chapter 15 339 340 Professional Paper NMFS 23 20. Virgularia cf. glacialis Description The original description of Virgularia identified by F. M. Bayer and collected from the same lo- glacialis by Sars (year unknown) is based on a small cation north of Amlia Island, central Aleutian Islands, at fragmented specimen and is rather rudimentary (i.e., it a depth of 122 m. Unfortunately the six specimens are does not even include location data). The subsequent severely deteriorated and of no descriptive value. Since description by Kölliker in 1870 (as Virgularia steenstru- these are the only specimen records of the species from pii) was based on an almost intact specimen. That speci- the Pacific Ocean and we cannot re-verify their identity men is 29.9 cm in length but only 2 mm in width and due to their poor condition, we list them here as V. cf. 18.7 cm (70%) of the specimen contains rather thick glacialis. They may be a different species of Virgularia, polyp leaves (2.2 mm in length) with 10 leaves per 3.2 perhaps V. cf. bromleyi or an undescribed species. cm of rachis length. The leaves are nearly triangular or sickle-shaped, attached along their whole length, but Distribution Rare. In Alaska – reported from a single with the keel lying such that the dorsal end is much location north of Amlia Island, central Aleutian Islands higher than the ventral end. (Fig. 15-20). Elsewhere – only known from the north- Polyps number 6–10 with scarcely separated goblets. west Atlantic Ocean, principally the fjords of Norway The back of the keel is completely covered by the leaves where the type specimen (V. steenstrupii) was described and the ventral side is completely exposed except for from Varangerfjord. Virgularia steenstrupii is now syn- two notches of the keel. Zooids are numerous with each onymized with V. glacialis as is another unaccepted filling the space between two leaves. species V. affinis Koren and Danielssen, 1877 also de- The axis is rather strong and reinforced with short scribed from Norwegian waters. radial fibers. Sclerites are not present, at least in the coenenchyme. The color of the colony is yellow to light Habitat In Alaska – found at a depth of 122 m. Else- brown; the axial skeleton is white. where – unknown. Remarks Known from six specimens (USNM 1010251) Photos None available. 341 CHAPTER 16 Class Hydrozoa Order Anthoathecata Family Stylasteridae Corals in the order Anthoathecata, family Stylasteridae of taxonomy for the group is very good (Cairns and are sometimes referred to incorrectly as “hydrocorals” Lindner, 2011). Nonetheless, all specimens collected in and are actually hydrozoans rather than anthozoans. Alaska should be properly preserved for taxonomic ex- The more proper common name for stylasterids is “lace amination and zoogeographic study. corals,” as the more general term hydrocoral is poly- Six genera of stylasterids are found in Alaska waters: phyletic, including corals from two different families: 1) Crypthelia – one species, 2) Cyclohelia – one spe- Stylasteridae and Milleporidae. Free medusae are not cies, 3) Distichopora – one species, 4) Errinopora – six produced in stylasterids like they are in many other species, 5) Stylantheca – one species, and 6) Stylaster hydrozoans, such as the milleporids. Stylasterids occur – 13 species (including one subspecies). The 10 species from the Arctic to the Antarctic at depths of 0 to 2789 in the genus Errinopora, six of which occur in Alaska m, but are most common at depths between 200 and waters, are compared in tabular and dichotomous keys 1200 m. Currently there are about 316 Recent species, by Cairns and Lindner (2011). The 14 species and sub- making it the second largest family within the Hydro- species in the genus Stylaster known from the Northeast zoa (Cairns, 2015). Pacific Ocean, 13 of which occur in Alaska waters, are Stylasterids that occur in Alaska waters are colonial also compared in a tabular key by Cairns and Lindner but ahermatypic; that is, they do not construct true (2011). reefs. Some species of stylasterids, however, are locally Conventional terminology used to describe stylaster- abundant and play a major role in constructing impor- ids, including an illustrated glossary, was published by tant benthic habitats (Stone, 2006, 2014). Twenty-two Cairns (2011a), and a key to all the stylasterid genera species (plus one subspecies) of stylasterids from six can be found in Cairns (2015). Morphology of the cal- genera are known from Alaska waters. The current state careous skeleton is used almost exclusively for identi- fication at the species and genus levels (Figs. 16-1, 16-2, 16-3, 16-4, and 16- 5). Examination of the col- ony with a scanning elec- tron microscope is often helpful, but not essential, to identify most Alaska species. Applying dye from a marker to the dry coenos- teum will help to determine the coenosteal texture. The stylasterid colony is polymorphic, mean- ing that every colony is composed of three types of polyps that are con- nected within the branch by a reticulate canal sys- tem: the gastrozooid (the feeding polyp), the dac- tylozooid (the defensive Figure 16-1 polyp), and the gonophore A photograph of the cyclosystem (center) of a Stylaster brochi colony showing the gastropore (the reproductive polyp). and surrounding structures. Each of these polyps has a corresponding calcare- 342 Professional Paper NMFS 23 Figure 16-2 A photograph of the distichoporine pore row (center) of a female Distichopora borealis colony showing the gastropore row, accompanying dactylopore rows, and ampullae. Figure 16-3 A close-up view of the fixed lid covering the cyclosystem of a Crypthelia trophostega colony and the associated structures. Chapter 16 343 Figure 16-4 A close-up view of the gastropore tube of a Stylaster brochi colony and the associated structures. Figure 16-5 A close-up view of a Distichopora borealis colony showing an adcauline dactylopore spine. 344 Professional Paper NMFS 23 ous structure associated with it: the gastrozooid has a continental shelf; however, several species (Crypthelia gastropore and gastropore tube, the dactylozooid has a trophostega Fisher, 1938; Cyclohelia lamellata Cairns, dactylopore and dactylopore spine, and the gonophore 1991; and Distichopora borealis Fisher, 1938) occur on has an ampulla. In some genera, several dactylopores Petrel and Bowers Banks that extend northward from encircle a single gastropore in a structure called a cy- the central Aleutian Islands into the Bering Sea. The closystem. Each gastrozooid has a mouth encircled by Aleutian Islands are far and away the hotspot of diver- a ring of tentacles, and each dactylozooid consists of sity in Alaska waters; 15 of 23 taxa appear to be en- one functional tentacle. Most stylasterids, including all demic to the region and half of the remaining eight taxa from Alaska, are gonochoric, meaning that each colony occur there. The records for most genera represent the is either male or female, which can usually be deter- northernmost in the world. They are eurybathic fauna mined by the size and shape of their ampullae. in Alaska, occupying depths between 11 and 2124 m. The reproductive ecology of seven species from the All taxa are firmly attached and typically grow on bed- Aleutian Islands was studied and all were brooders, rock, boulders, cobbles, pebbles, and occasionally on with the majority of gonophores containing mature em- siltstone and hexactinellid sponge skeleton. bryos or planulae (Brooke and Stone, 2007). The de- Most taxa are quite small, generally less than 10 cm velopmental stage of gametes within individual colonies in height and width, but a few species —e.g., Stylaster was not highly synchronized; females contained eggs as campylecus (Fisher, 1938) and Stylaster alaskanus well as planulae, and males exhibited a range of gamete Fisher, 1938—attain sizes up to 60 cm in height and development. These reproductive traits indicate that hy- width. Most taxa are found either in low- or high- drocorals have limited potential to recolonize disturbed density patches; these patches may provide important areas in the Aleutian Islands (Brooke and Stone, 2007). refuge and foraging habitat for juvenile fish and crabs Stylasterids are found in only three regions of Alaska (Stone and Shotwell, 2007). They are a fairly common (eastern Gulf of Alaska, western Gulf of Alaska, and bycatch item in fisheries and stock assessment surveys the Aleutian Islands) and are noticeably absent from but are extremely fragile, often fragmented, and seldom the Gulf of Alaska Seamount Province and the Arctic retained for formal identification and reference collec- (Fig. 16-6). They are also unknown from the Bering Sea tions. Figure 16-6 A map of the North Pacific Ocean showing the distribution of corals in the family Stylasteridae () in Alaska waters. Chapter 16 345 1. Crypthelia trophostega Fisher, 1938 Description (Adapted from Cairns and Lindner, Remarks Crypthelia is a species-rich genus (46 spe- 2011) Colony branching is usually uniplanar, but oc- cies) worldwide; however, only a single species is casionally multiplanar or bushy, and dichotomous. known from Alaska waters. They are easily diagnosed Branch anastomosis is common. The largest colony by having a cyclosystem lid. The reproductive ecology known is about 20 cm in height and width. The coe- of this species has been studied from colonies collected nosteum is linear-imbricate in texture. Nematopores in the Aleutian Islands (Brooke and Stone, 2007). The are common and occur on the coenosteum, cyclosystem coenosteum is composed mostly of aragonite with small lid, and pseudosepta. The coenosteum and tissue are amounts of low-magnesium calcite (Cairns and Mac- creamy white to white. intyre, 1992; senior author and S. D. Cairns, unpubl. The gastropores and dactylopores are arranged in data). discrete cyclosystems that occur unilinearly and unifa- cially or bifacially on branches. The cyclosystems are Distribution Common. In Alaska – endemic to the 2.2–2.6 mm in diameter, each having 13–23 dactylo- Aleutian Islands (Amukta Pass to southeast of Agattu pores, but averaging about 19. Every cyclosystem is cov- Island) including Bowers Bank (Fig. 16-7). There is a ered with a large horizontal fixed lid that covers most single record (USNM 76781) collected by a fishing ves- of the gastropore (Fig. 16-3). The gastropore tube is sel near the Pribilof Islands, eastern Bering Sea, at a double-chambered. depth of 550 m. We discount this record as one that The female ampullae are massive hemispherical swell- was almost certainly translocated by fishing gear. Else- ings, one or two of which occur on the cyclosystem lid, where – not reported. and their efferent pores open beneath the lid. The male ampullae are smaller, up to 11 also occur on the lid, and Habitat Typically found singly but occasionally in their efferent pores also open beneath the lid. small patches, on hard rock including cobbles and large Figure 16-7 A map of the Aleutian Islands showing the distribution of Crypthelia trophostega () in Alaska waters. 346 Professional Paper NMFS 23 pebbles, in areas of moderate current, and at depths be- A. C) A C. trophostega colony photographed in Adak tween 138 and 1913 m. Canyon, central Aleutian Islands, at a depth of 1335 m. D) A C. trophostega colony photographed northwest Photos A) Most of a whole dried C. trophostega col- of Tanaga Island, central Aleutian Islands, at a depth ony (USNM 1482073) collected east of Semisopochnoi of 1247 m. An unknown liparid (snailfish) takes refuge Island, in the central Aleutian Islands, at a depth of 457 near the coral. The distance between the red laser marks m. B) A close-up view of the same specimen in photo in photos C and D is 10 cm. Chapter 16 347 2. Cyclohelia lamellata Cairns, 1991 Description (Adapted from Cairns, 1991 and Cairns Remarks Cyclohelia is a monotypic genus closely re- and Lindner, 2011) Colonies are firmly attached by a lated to Distichopora but differs in having non-linear robust cylindrical stem that is up to 2.5 cm in diam- gastro- and dactylopores and internal ampullae. The eter and bifurcates into two lamellae or plates, each reproductive ecology of this species has been studied of which increases its surface area by folding and un- from colonies collected in the Aleutian Islands (Brooke dulating its surface into a complex three-dimensional and Stone, 2007). The coenosteum is composed entirely structure. Some colonies are coarsely digitiform. Large of aragonite (senior author and S. D. Cairns, unpubl. colonies are up to 8 cm in height and 10 cm in width. data). The coenosteum is reticulate-granular in texture; nema- topores are absent. The color of the coenosteum is Distribution Uncommon. In Alaska – endemic to the typically reddish-orange and rarely yellow in lamellate Aleutian Islands (Islands of Four Mountains to west of forms, but is typically pink in digitate forms. Buldir Island) including Petrel Bank (Fig. 16-8). There The gastro- and dactylopores are abundant and occur is a single record (USNM 85007) collected by a fishing uniformly but without coordination on all corallum fac- vessel near the Pribilof Islands, eastern Bering Sea, at es. The gastropores are circular and flush with coenos- a depth of 550 m. We discount this record as one that teum; the gastrostyles are elongate and needle-shaped. was almost certainly translocated by fishing gear. Else- The dactylopore spines are elliptical in shape and the where – not reported. lateral edges are higher than the edges at the vertices. The female ampullae are primarily internal and up Habitat Typically found singly but occasionally in to 1.1 mm in internal diameter. The male ampullae are small patches, on hard rock including bedrock and cob- also internal but smaller, only up to about 0.6 mm in bles, in areas of high to moderate current, and at depths internal diameter, and communicate to the surface via a between 27 and 405 m (video observations to 691 m; narrow efferent duct. Stone, 2014). Figure 16-8 A map of the Aleutian Islands showing the distribution of Cyclohelia lamellata () in Alaska waters. 348 Professional Paper NMFS 23 Photos A) Most of a dried whole C. lamellata colony central Aleutian Islands, at a depth of 115 m. D) A la- (USNM 1482043) in the common lamellar form. This mellar form of C. lamellata (at the bottom-center of the specimen was collected northwest of Kiska Island, west- image) photographed northwest of Tanaga Island, cen- ern Aleutian Islands, at a depth of 366 m. B) A close-up tral Aleutian Islands, at a depth of 170 m. E) A lamellar view of the same specimen in photo A. C) A dried branch form of C. lamellata (center) photographed in the central of a C. lamellata colony in the less common digitate form Aleutian Islands at a depth of 498 m. (USNM 1482046) collected in southern Amchitka Pass, Chapter 16 349 3. Distichopora borealis Fisher, 1938 Description (Adapted from Cairns and Lindner, 2011) guished from all other stylasterids in Alaska waters by Colonies are usually uniplanar in branching but occa- the unique coordination of its gastro- and dactylopores sionally multiplanar or arborescent. Branching is irregu- in pore rows. Otherwise, Distichopora is a widely dis- larly dichotomous and sometimes anastomotic; branch tributed genus at depths of 0 to 1361 m, containing 27 tips are rounded to blunt. Large colonies are up to 11 cm described species (Cairns, 2015). in height and 20 cm in width, with a basal branch diam- The coenosteum is composed principally (92– eter up to 3 cm. The coenosteum is reticulate-granular 99%) of high-magnesium calcite (9.0–9.7 mol% MgCO3) in texture and often longitudinally ridged; nematopores with small amounts of aragonite and low-magnesium cal- are absent. The coenosteum is white to light orange, and cite (senior author and S. D. Cairns, unpubl. data). the latter color morph has a white core. The gastro- and dactylopores are arranged in pore Distribution Common. In Alaska – this species has a rows; the gastropores are unilinearly arranged on branch highly disjunct distribution. It is known mostly from the edges and flanked on either side by a row of dactylopore Aleutian Islands (southwest of Umnak Island to west of spines. The gastropores are circular, 12–16 per centime- Attu Island) including Petrel and Bowers Banks and also ter; the gastrostyles are needle-shaped. The dactylopore from Shutter Ridge in the eastern Gulf of Alaska (Fig. spines are U-shaped and occur 18–40 per centimeter; 16-9). Only a single specimen has been collected in the dactylostyles are absent. eastern Gulf of Alaska, but video observations (Stone The female ampullae are prominent superficial mounds et al., 2015a) indicate that the species is relatively com- up to 1.5 mm in diameter, each having a lateral efferent mon there at depths between 137 and 244 m. Elsewhere pore. The male ampullae are primarily internal, irregular – not reported. in shape, and rarely more than 0.4 mm in diameter. Habitat Typically found singly but occasionally in Remarks This species (and genus) is easily distin- small patches, on hard rock including bedrock, cobbles, Figure 16-9 A map of the North Pacific Ocean showing the distribution of Distichopora borealis () in Alaska waters. 350 Professional Paper NMFS 23 and pebbles, in areas of moderate current, and at depths in the western Aleutian Islands at a depth of 256 m. between 137 and 1361 m. The latter depth is the great- C) A close-up view of the same specimen in photo B. est for any species in the genus. In the eastern Gulf of D) Dried branches of a D. borealis colony (USNM Alaska this species forms patches with Stylaster para- 1482068) collected in the central Aleutian Islands at geus parageus (Fisher, 1938) where it constitutes about a depth of 1266 m. The tip of the lower branch is 10% of the total number of hydrocorals present. overgrown by a demosponge. E) Several D. borealis colonies in situ in the central Aleutian Islands at a Photos A) Most of a whole dried D. borealis colo- depth of 681 m. F) A D. borealis colony in situ in ny (USNM 1482063) collected in the western Aleu- the central Aleutian Islands at a depth of 720 m. The tian Islands at a depth of 292 m. B) Most of a whole distance between the red lasers marks in photos E and dried D. borealis colony (USNM 1482065) collected F is 10 cm. Chapter 16 351 4. Errinopora dichotoma Lindner and Cairns, 2011 Description (Adapted from Cairns and Lindner, Remarks The coenosteum is composed principal- 2011) Colonies are robust, with sparse, dichotomous, ly (98.7%) of high-magnesium calcite (10.9 mol% three-dimensional branching. Colonies are up to 11.5 MgCO3) with very small amounts of aragonite and low- cm in height with basal branch diameters up to 2 cm; magnesium calcite (senior author and S. D. Cairns, un- the branch tips are blunt. The coenosteal texture is re- publ. data). ticulate-spinose. The coenosteum is light orange in color and the branch core is white. Distribution Rare (known from only four specimens). The gastropores are circular, 0.3–0.5 mm in diame- In Alaska – endemic to the eastern Aleutian Islands ter, and arranged randomly or in rows; the gastrostyles from the Islands of Four Mountains to near Seguam Is- are lanceolate. Smaller gastropores (0.22–0.30 mm in land (Fig. 16-10). Elsewhere – not reported. diameter) are also present. The dactylopore spines are quite variable in orientation, sometimes aligned in lon- Habitat Largely unknown but attaches to hard rock gitudinal or transverse rows flanking the gastropores, in areas of moderate to high current and at depths be- or isolated in random orientation. The dactylopore tween 175 and 405 m. spines are thick-walled, up to 1.1 mm in height, and often laterally fused to one another. The dactylostyles Photos A) Most of a whole dried E. dichotoma colony are robust. (USNM 1123507; paratype) collected near the Islands The female ampullae are known only from spent de- of Four Mountains, eastern Aleutian Islands, at a depth pressions (1.0–1.1 mm in diameter) in the coenosteum. of 405 m. B) A close-up view of a dried E. dichotoma The male ampullae are small (0.5–0.6 mm in diameter), specimen (USNM 1137600; paratype) collected near the porous superficial hemispheres. Islands of Four Mountains at a depth of 176 m. 352 Professional Paper NMFS 23 4. Errinopora dichotoma Lindner and Cairns, 2011 (continued) Figure 16-10 A map of the Aleutian Islands showing the distribution of Errinopora dichotoma () in Alaska waters. Chapter 16 353 5. Errinopora disticha Lindner and Cairns, 2011 Description (Adapted from Cairns and Lindner, Remarks Errinopora disticha is the only species of 2011) Colonies are robust with dichotomous branching Errinopora in Alaska that has a distichoporine arrange- that forms uniplanar to multiplanar coralla up to 13.5 ment of its gastro- and dactylopores (Fig. 16-2). cm in height. Branches are circular in cross section, but The coenosteum is composed principally (92%) of most often strongly compressed. The coenosteal texture high-magnesium calcite (10.4–11.6 mol% MgCO3) is reticulate-spinose. The coenosteum is light orange in with small amounts of aragonite and low-magnesium color with a white branch core. Parasitic spionid poly- calcite (senior author and S. D. Cairns, unpubl. data). chaete tubes are found in branch axes of some colonies. The gastropores are circular in shape, 0.3–0.7 mm Distribution Rare. In Alaska – endemic to the Aleu- in diameter, and often arranged unilinearly in a shal- tian Islands from the Islands of Four Mountains to low sulcus similar to Distichopora; the gastrostyles are northwest of Semisopochnoi Island (Fig. 16-11). Else- lanceolate. A smaller class of gastropores measures only where – not reported. 0.2–0.3 mm in diameter. The dactylopore spines are ar- ranged in long meandering rows flanking both sides of Habitat Largely unknown but attaches to hard rock a gastropore row (thus bilaterally arranged) and their in areas of moderate to high current and at depths be- lateral edges are often fused. The dactylopore spines are tween 176 and 536 m. thick-walled and short (only 0.5–0.9 mm in length); the dactylostyles are robust. Photos A) The dried E. disticha holotype (USNM The female ampullae are superficial mounds, 1.3–1.5 1123524; most of a whole colony) collected near Semi- mm in diameter, and sometimes with a ridged surface. sopochnoi Island, western Aleutian Islands, at a depth The male ampullae are also superficial blisters, 0.4–0.6 of 530 m. B) Most of a whole dried E. disticha colony mm in diameter, and often occur in dense concentra- (USNM 1123523; paratype) collected near Semisopoch- tions. noi Island at a depth of 475 m. 354 Professional Paper NMFS 23 5. Errinopora disticha Lindner and Cairns, 2011 (continued) Figure 16-11 A map of the Aleutian Islands showing the distribution of Errinopora disticha () in Alaska waters. Chapter 16 355 6. Errinopora fisheri Lindner and Cairns, 2011 Description (Adapted from Cairns and Lindner, ka species in having gastro- and dactylopores arranged 2011) Colonies branch in a uniplanar fashion, with fine in pseudocyclosystems. irregular dichotomous branching; the branch tips are The coenosteum is composed principally (95%) of slender. Colonies are up to 9 cm in height and 6.5 cm in high-magnesium calcite (10.6 mol% MgCO3) with very width. The coenosteal texture is reticulate-granular and small amounts of aragonite and low-magnesium calcite the coenosteum is light orange. (senior author and S. D. Cairns, unpubl. data). The gastropores are circular, 0.3–0.5 mm in diam- eter, and arranged uniformly on the branch surfaces; Distribution Extremely rare (known only from the the gastrostyles are squat. The dactylopore spines are two type specimens). In Alaska – endemic to the west- arranged in abcauline crescents below a gastropore or ern Aleutian Islands (west of Attu Island; Fig. 16-12). even encircling a gastropore, similar to the way they are Elsewhere – not reported. in Stylaster. The dactylopore spines are up to 1.5 mm in height and laterally fused; the dactylotomes are ellipti- Habitat Largely unknown but attaches to hard rock cal in shape (not slit-shaped); and the dactylostyles are in areas of moderate to high current and at depths be- robust. tween 455 and 458 m. The female ampullae are superficial hemispheres (1.0–1.2 mm in diameter); efferent pores appear to be Photos A) The dried E. fisheri holotype (USNM lacking. Male ampullae are unknown. 1123526; whole colony) collected near Attu Island, western Aleutian Islands, at a depth of 455 m. B) A Remarks Errinopora fisheri is unique among the Alas- close-up view of the same specimen in photo A. 356 Professional Paper NMFS 23 6. Errinopora fisheri Lindner and Cairns, 2011 (continued) Figure 16-12 A map of the Aleutian Islands showing the distribution of Errinopora fisheri () in Alaska waters. Chapter 16 357 7. Errinopora nanneca Fisher, 1938 Description (Adapted from Cairns and Lindner, 2011) tian Islands (Brooke and Stone, 2007). The skeleton Colonies are quite variable in shape; branching is usu- (coenosteum) is composed principally (97%) of high- ally uniplanar and irregularly dichotomous but occasion- magnesium calcite (9.9–10.9 mol% MgCO3) with very ally multiplanar and multilobate. Parasitic spionid poly- small amounts of aragonite and low-magnesium calcite chaetes often form tubes along the branch axes. Colonies (Cairns and Macintyre, 1992; senior author and S. D. are large, up to 21 cm in height and width. The coe- Cairns, unpubl. data). nosteal texture is reticulate-spinose. The coenosteum is typically orange but occasionally pink (in deeper water). Distribution Common and locally abundant. In Alas- The gastropores are circular, flush with the coenos- ka – endemic to the Aleutian Islands (Islands of Four teum, and 0.15–0.44 mm in diameter; the gastrostyles Mountains to Tahoma Reef southwest of Buldir Island), are lanceolate. The dactylopore spines are isolated or including Petrel Bank (Fig. 16-13). Elsewhere – not re- arranged in transverse to oblique rows with the dactylo- ported. tomes facing upward (abcauline) and their lateral edges fusing. Dactylopore spines are relatively small (only 0.4 Habitat Found singly but occasionally in small mm in height); the dactylostyles are robust. patches on hard rock, including bedrock, cobbles, and The female ampullae are large superficial hemispheres pebbles, in areas of moderate current, and at depths be- (1.1–1.8 mm in diameter) and each has a lateral efferent tween 22 and 594 m. pore. The male ampullae are smaller mounds (0.4–0.7 mm in diameter) and are often clustered. Both types of Photos A) A dried whole E. nanneca colony (USNM ampullae are porous. 1481973) collected near Bobrof Island, central Aleutian Islands, at a depth of 27 m. B) A dried whole E. nan- Remarks The reproductive ecology of this species neca colony (USNM 1481966) collected near Semiso- has been studied from colonies collected in the Aleu- pochnoi Island, central Aleutian Islands, at a depth of Figure 16-13 A map of the Aleutian Islands showing the distribution of Errinopora nanneca () in Alaska waters. 358 Professional Paper NMFS 23 22 m. C) A dried whole E. nanneca colony (USNM chitka Pass, central Aleutian Islands, at a depth of 125 1223510) collected near Cape Moffet, Adak Island, m. E) An E. nanneca colony (top, center) photographed central Aleutian Islands, at a depth of 150 m. D) A near Semisopochnoi Island at a depth of 27 m. F) An close-up view of a dried E. nanneca specimen (USNM E. nanneca colony photographed near Semisopochnoi 1123450) collected on Amchixtam Chaxsxii in Am- Island at a depth of 27 m. Chapter 16 359 8. Errinopora undulata Lindner and Cairns, 2011 Description (Adapted from Cairns and Lindner, ner, 2011 or Cyclohelia lamellata, but these two spe- 2011) Colonies are lamellate, with thin sinusoidal dis- cies are in other discrete genera with other diagnostic tal edges, and a massive basal branch. Colonies are up characteristics. to 13 cm in height and 15 cm in width. Parasitic spio- The coenosteum is composed principally (97–98%) nid polychaete tubes are found in some colonies. The of high-magnesium calcite (10.3–10.7 mol% MgCO3) coenosteal texture is reticulate-spinose; the coenosteal with very small amounts of aragonite and low-magne- strips are separated by wide slits. The coenosteum is sium calcite (senior author and S. D. Cairns, unpubl. light orange. data). The gastropores are circular, flush with the coenos- teum, and measure 0.3–0.45 mm in diameter; the gas- Distribution Rare (known from only three speci- trostyles are lanceolate. The dactylopore spines are clus- mens). In Alaska – endemic to the Aleutian Islands tered around the gastropores in pseudocyclosystems. (Amukta Pass to west of Semisopochnoi Island; Fig. 16- The dactylopore spines are small (0.25 mm in height) 14). Elsewhere – not reported. and thin-walled; the dactylostyles are robust. The female ampullae are irregularly shaped, superfi- Habitat Largely unknown but attaches to hard rock cial hemispheres and measure up to 1.6 mm in diameter. in areas of moderate to high current and at depths be- Male ampullae are unknown. tween 366 and 425 m. Remarks Only five species of stylasterids have a la- Photos A) Most of a whole dried E. undulata colony mellate corallum and four of those occur in Alaska wa- (USNM 1123528; paratype) collected in Amukta Pass, ters. The gross morphology of Errinopora undulata can eastern Aleutian Islands, at a depth of 425 m. B) A be confused with Stylaster repandus Cairns and Lind- close-up view of the same specimen in photo A. 360 Professional Paper NMFS 23 8. Errinopora undulata Lindner and Cairns, 2011 (continued) Figure 16-14 A map of the Aleutian Islands showing the distribution of Errinopora undulata () in Alaska waters. Chapter 16 361 9. Errinopora zarhyncha Fisher, 1938 Description (Adapted from Cairns and Lindner, 2011) Remarks Errinopora zarhyncha is distinctive among Colonies are typically uniplanar but occasionally bushy the Alaska Errinopora in having a robust colony with and quite robust with equal dichotomous branching. very large gastropores and tall dactylopore spines. Colonies are up to 26 cm in height and 24 cm in width. The coenosteum is composed principally (95%) of The branches are thick and have blunt tips. The coenos- high-magnesium calcite (10.6 mol% MgCO3) with very teal texture is reticulate-spinose and quite porous. The small amounts of aragonite and low-magnesium calcite coenosteum is light orange to orange. (Cairns and Macintyre, 1992; senior author and S. D. The gastropores are circular and quite variable in Cairns, unpubl. data). size, up to 1.1 mm in diameter, and often closely packed and linearly arranged. The gastrostyles are lanceolate Distribution Uncommon. In Alaska – endemic to the but relatively small due to the large diameter of the Aleutian Islands (Amukta Pass to Amchitka Island) in- gastropore tube. The dactylopore spines are quite tall cluding Bowers Bank and Ridge (Fig. 16-15). Elsewhere (up to 3 mm in height), thick-walled, and usually later- – not reported. ally fused into transverse tiers flanking the gastropores; compound dactylopore spines are common. Habitat Typically found singly but occasionally in The female ampullae are low hemispherical mounds small patches on hard rock, including bedrock, boul- (up to 1.25 mm in diameter), but are usually overshad- ders, and cobbles, in areas of moderate current, and at owed by an adjacent dactylopore spine; its efferent pore depths between 115 and 658 m. Video observations is lateral. The male ampullae are smaller (about 0.6–0.7 (Stone, 2014), however, indicate that this species occurs mm in diameter) and also superficial. as deep as 1520 m in the central Aleutian Islands. Figure 16-15 A map of the Aleutian Islands showing the distribution of Errinopora zarhyncha () in Alaska waters. 362 Professional Paper NMFS 23 Photos A) The dried E. zarhyncha holotype (USNM zarhyncha colony (center) photographed on Amchixtam 42874; whole colony) collected in Amukta Pass, eastern Chaxsxii in Amchitka Pass, central Aleutian Islands, at Aleutian Islands, at a depth of 518 m. B) Two mostly a depth of 862 m. E) Errinopora zarhyncha colonies whole dried E. zarhyncha colonies (USNM 1123446) (indicated by the white circle) photographed in northern collected south of Amlia Island, central Aleutian Is- Amchitka Pass, central Aleutian Islands, at a depth of lands, at a depth of 497 m. C) An E. zarhyncha colony 830 m. The distance between the red laser marks is 20 (center, top) photographed northwest of Tanaga Island, cm. F) A closer view of the same colonies in photo E. central Aleutian Islands, at a depth of 162 m. D) An E. The distance between the red laser marks is 10 cm. Chapter 16 363 10. Stylantheca papillosa (Dall, 1884) Description (Adapted from Cairns and Lindner, this species include Stylantheca porphyra Fisher, 1931 2011) Colonies consist of either thin (1–2 mm) en- and Allopora petrograpta Fisher, 1938. The blue top- crustations or thicker ones that produce short knobby snail (Calliostoma ligatum) is often associated with S. branches; colonies are up to 30 cm in diameter. The papillosa colonies and may feed on their polyps (senior coenosteum is reticulate-granular in texture and covered author, personal observ.). with short conical papillae (possibly nematopores). The The coenosteum is composed principally (86–99%) coenosteum is purple, red, or pink and occasionally has of aragonite with small amounts of low-magnesium cal- white tips on the papillae. cite and very small amounts of high-magnesium calcite The gastro- and dactylopores are arranged in cyclo- (senior author and S. D. Cairns, unpubl. data). systems, measure 0.9–1.2 mm in diameter, and are uni- formly distributed on the encrustation. The gastropore Distribution Uncommon. In Alaska – eastern Gulf of tube is about 0.3 mm in diameter, the ring palisade is well Alaska including the inside waters of Southeast Alaska developed, and the gastrostyles are globose or triangular. (Fig. 16-16). The Alaska records represent the northern- The dactylopores per cyclosystem range from 2 to 12, but most for the species. Elsewhere – British Columbia to usually average 4 to 8; the dactylostyles are quite robust. northern California (south of San Francisco Bay). Both female (0.7–0.9 mm in diameter) and male (0.4– 0.5 mm in diameter) ampullae are internal. Habitat In Alaska – most common on bedrock in ar- eas of high current and tidal surge, at depths between Remarks Stylantheca papillosa is the only encrusting 6 and 40 m. This is the shallowest stylasterid in Alaska stylasterid known in Alaska waters and may be the only waters. Elsewhere – on bedrock at depths between the encrusting stylasterid worldwide. Junior synonyms for lower intertidal zone and 27 m. Figure 16-16 A map of the eastern Gulf of Alaska showing the distribution of Stylantheca papillosa () in Alaska waters. 364 Professional Paper NMFS 23 Photos A) A whole dried S. papillosa colony (USNM a depth of 18 m. E) A patch of S. papillosa colonies (in- 1482029) collected near Cape Ommaney, eastern Gulf cluding USNM 1482029) at a depth of 18 m near Cape of Alaska, at a depth of 18 m. B) A whole dried S. pap- Ommaney. F) Adult lingcod (Ophiodon elongatus) rest illosa colony (USNM 1482032) collected near Cross on a large patch of S. papillosa in the Edgecumbe Pin- Sound, eastern Gulf of Alaska, at a depth of 18 m. C) nacles Marine Reserve near Sitka, Alaska. The depth is A close-up view of the same specimen in photo B. D) A approximately 40 m and thus extends the depth range close-up view of a dried S. papillosa specimen (USNM for the species. 148202) collected in Burrard Inlet, British Columbia, at Chapter 16 365 11. Stylaster alaskanus Fisher, 1938 Description (Adapted from Cairns and Lindner, ka species in having a reticulate colony and ornamented 2011) Colonies are uniplanar, or dichotomously female ampullae. It was previously known as Stylaster branched and highly anastomotic, forming a sieve-like cancellatus Fisher, 1938. reticulum. The largest known specimen is 60 cm in Associated fauna include ophiuroids. The repro- height and width but larger colonies have been observed ductive ecology of this species has been studied from in situ (see photo C on next page). The coenosteum is colonies collected in the Aleutian Islands (Brooke and linear-imbricate in texture and light orange, pink, or Stone, 2007). The coenosteum is composed principally creamy white in color. (99–100%) of aragonite with very small amounts of The gastro- and dactylopores are arranged in cyclo- low- and high-magnesium calcite (senior author and S. systems that are linearly arranged on the branch edges D. Cairns, unpubl. data). only. Cyclosystems are irregular in shape and 0.9–1.3 mm in diameter. The gastropore tubes are funnel- shaped, with a prominent ring palisade and a lanceolate Distribution Common. In Alaska – endemic to the gastrostyle. The range of dactylopores per cyclosystem Aleutian Islands (Amukta Pass to east of Stalemate is 7–14, but averages about 11; dactylostyles are rudi- Bank) including Petrel Bank and Bowers Ridge (Fig. 16- mentary. 17). Elsewhere – not reported. The female ampullae are superficial hemispheres up to 0.9 mm in diameter, often bear low ridges or short Habitat Stylaster alaskanus is often found in complex spines, and have a lateral efferent pore. The male ampul- habitat and is a good indicator species of coral garden lae are also superficial, up to 0.6 mm in diameter, and habitat. It is typically found singly but occasionally in usually densely clustered. small patches on hard rock, including bedrock, boul- ders, and cobbles, in areas of moderate current, and at Remarks Stylaster alaskanus differs from other Alas- depths between 146 and 2124 m. Figure 16-17 A map of the Aleutian Islands showing the distribution of Stylaster alaskanus () in Alaska waters. 366 Professional Paper NMFS 23 Photos A) A large fragment of a dried S. alaskanus ern Aleutian Islands, at a depth of 256 m. C) A large S. colony (USNM 1122455) collected south of Amchitka alaskanus colony (70 cm in height by 74 cm in width) Island, western Aleutian Islands, at a depth of 265 m. photographed in northern Amchitka Pass in the central B) A close-up view of a dried S. alaskanus specimen Aleutian Islands at a depth of 712 m. The distance be- (USNM 1122466) collected west of Kiska Island, west- tween the red laser marks is 20 cm. Chapter 16 367 12. Stylaster brochi (Fisher, 1938) Description (Adapted from Cairns and Lindner, branch surfaces, and thus belonged to the genus previ- 2011) Colonies are variable in shape, ranging from uni- ously called Allopora. The reproductive ecology of this planar branching to bushy, with irregular dichotomous species has been studied from colonies collected in the branching; the largest known colony is 28 cm in height. Aleutian Islands (Brooke and Stone, 2007). The coenos- All colonies are infested with the spionid polychaete teum is composed principally (88–99%) of aragonite Polydora, which forms characteristic binary tubes along with small amounts of low-magnesium calcite and very most branch axes. The coenosteum has a reticulate- small amounts of high-magnesium calcite (senior author granular texture and is pale orange in color. and S. D. Cairns, unpubl. data). The gastro- and dactylopores are arranged in cy- closystems that are uniformly arranged on all branch Distribution Very common and widespread. In Alas- surfaces; cyclosystems are 0.9–1.4 mm in greater diam- ka – eastern Gulf of Alaska including the inside wa- eter. The gastropore tubes are cylindrical, elongate, and ters of Southeast Alaska, western Gulf of Alaska, and usually curved, containing a diffuse ring palisade and throughout the Aleutian Islands including Petrel Bank an elongate gastrostyle. The range of dactylopores per (Fig. 16-18). Elsewhere – not reported. cyclosystem is 6–13, but averages about 9; the dactylo- styles are well developed. Habitat Typically found in small patches on hard rock The female ampullae are superficial mounds (0.8–1.1 including bedrock, boulders, and cobbles, in areas of mm in diameter) and each has a lateral efferent pore. moderate current, and at depths between 20 and 455 m. The male ampullae are clustered, primarily internal, and measure 0.3–0.5 mm in diameter. Photos A) A dried whole S. brochi colony (USNM 1482034) collected near Semisopochnoi Island, central Remarks Stylaster brochi is one of four Alaska spe- Aleutian Islands, at a depth of 22 m. B) A close-up view cies to have its cyclosystems arranged uniformly on all of the same specimen in photo A. 368 Professional Paper NMFS 23 12. Stylaster brochi (Fisher, 1938) (continued) Figure 16-18 A map of the North Pacific Ocean showing the distribution of Stylaster brochi () in Alaska waters. Chapter 16 369 13. Stylaster campylecus (Fisher, 1938) Description (Adapted from Cairns and Lindner, Remarks Stylaster campylecus is distinctive in having 2011) Colonies branch dichotomously in a uniplanar longitudinally arranged coenosteal strips on the abaxial or multiplanar fashion; in large colonies these branch- side of the cyclosystems. This species is also known as es sometimes anastomose. The largest known colony S. polyorchis (Fisher, 1938), S. moseleyanus (Fisher, (USNM 1482002) is 50 cm in height and 38 cm in 1938), S. campylecus tylotus (Fisher, 1938), and S. cam- width. The coenosteal texture is reticulate-granular to pylecus campylecus (Fisher, 1938). The reproductive porcelaneous; however, the coenosteal strips are longi- ecology of this species has been studied from colonies tudinally arranged on the abaxial side of cyclosystems. collected in the Aleutian Islands (Brooke and Stone, The coenosteum is white, pale orange, pale pink, and 2007). The coenosteum is composed principally (87– occasionally bright pink. 99%) of aragonite with small amounts of low-magne- The gastro- and dactylopores are arranged in cyclo- sium calcite and very small amounts of high-magnesium systems located on branch edges as well as the anterior calcite (Cairns and Macintyre, 1992; senior author and face; the cyclosystems measure 1.0–1.3 mm in diameter. S. D. Cairns, unpubl. data). The gastropore tubes are cylindrical and curved, often containing a poorly developed ring palisade and a short, Distribution Common, abundant in some areas, and stout gastrostyle. The range of dactylopores per cyclo- widespread. In Alaska – eastern Gulf of Alaska includ- system is 7–17, but averages about 12; dactylostyles are ing the inside waters of Southeast Alaska, western Gulf rudimentary. of Alaska, and throughout the Aleutian Islands includ- The female ampullae are large (1.0–1.1 mm in diam- ing Petrel and Bowers Banks (Fig. 16-19). Elsewhere – eter) superficial hemispheres and each has a lateral effer- not reported. ent pore. The male ampullae are also superficial, mea- sure 0.4–0.5 mm in diameter, and are usually clustered Habitat Typically found in patches on hard rock in- on anterior branch faces. cluding bedrock, boulders, cobbles, and pebbles, in Figure 16-19 A map of the North Pacific Ocean showing the distribution of Stylaster campylecus () in Alas- ka waters. 370 Professional Paper NMFS 23 areas of moderate current, and at depths between 79 specimen collected near Cape Moffet, Adak Island, cen- and 1396 m. tral Aleutian Islands, at a depth of 150 m. D) A patch of S. campylecus colonies at a depth of 681 m in the Photos A) A large portion of a S. campylecus colony central Aleutian Islands. E) A close-up view of a large (USNM 1481994) collected in southern Amchitka Pass, S. campylecus colony in photo D. F) A small patch of central Aleutian Islands, at a depth of 115 m. B) A large S. campylecus colonies near the Delarof Islands, south- portion of a S. campylecus colony (USNM 1481992) ern Amchitka Pass, central Aleutian Islands, at a depth collected on Petrel Bank, central Aleutian Islands, at a of 165 m. The distance between the red laser marks in depth of 145 m. C) A close–up view of a S. campylecus photos D–F is 10 cm. Chapter 16 371 14. Stylaster crassiseptum Cairns and Lindner, 2011 Description (Adapted from Cairns and Lindner, ing very broad pseudosepta. The coenosteum is com- 2011) Colony branching is uniplanar, in an irregular posed entirely of aragonite (senior author and S. D. dichotomous fashion; the largest known specimen is 24 Cairns, unpubl. data). cm in height and 19 cm in width. The coenosteal tex- ture is reticulate-granular, sometimes almost porcella- Distribution Rare. In Alaska – known from only neous; the coenosteum is pale orange. six specimens collected in the Aleutian Islands (south The gastropores and dactylopores are arranged in of Atka Island to west of Kiska Island) and on Bowers cyclosystems that occur exclusively on opposite branch Bank (Fig. 16-20). It is apparently endemic to the re- edges; the cyclosystems are only 0.7–1.0 mm in diam- gion. Elsewhere – not reported. eter. The gastropore tubes are cylindrical and slightly curved, each containing a well-developed ring palisade Habitat Largely unknown but likely found singly or and a lanceolate gastrostyle. The range of dactylopores in small patches on hard rock including bedrock and per cyclosystem is 6–12, but averages about 9; the dacty- boulders, in areas of moderate current, and at depths lostyles are robust. Pseudosepta are broad and rounded, between 291 and 531 m. and some are up to five times the width of the adjacent dactylotome. Photos A) A large portion of a dried S. crassiseptum The female ampullae are superficial hemispheres and colony (USNM 1122497; paratype) collected on Bowers measure 0.9–1.1 mm in diameter. The male ampullae Bank, central Aleutian Islands, at a depth of 291 m. B) are also superficial, measure 0.45–0.50 mm in diameter, A close-up view of the same specimen in photo A. C) A and are often clustered. large portion of a dried S. crassiseptum colony (USNM 1122527; paratype) collected south of Atka Island, cen- Remarks Stylaster crassiseptum is distinctive in hav- tral Aleutian Islands, at a depth of 531 m. 372 Professional Paper NMFS 23 14. Stylaster crassiseptum Cairns and Lindner, 2011 (continued) Figure 16-20 A map of the Aleutian Islands showing the distribution of Stylaster crassiseptum () in Alaska waters. Chapter 16 373 15. Stylaster elassotomus Fisher, 1938 Description The holotype is a bushy colony with ir- the Alaska species in having large gastropores and rela- regular, dichotomous branching and measures 5.5 cm in tively short dactylotomes. height and about 5 cm in width. The coenosteum has a reticulate-granular texture and is white in color. Distribution Extremely rare. In Alaska – known from The gastro- and dactylopores are arranged in cir- only a single specimen collected southeast of Agattu Is- cular to slightly elliptical cyclosystems located exclu- land in the western Aleutian Islands (Fig. 16-21). It is sively on the two branch edges; the cyclosystems are apparently endemic to the region. Elsewhere – not re- 1.0–1.2 mm in greater diameter. The gastropores are ported. quite large (up to 0.6 mm in diameter); the gastro- pore tubes are cylindrical and highly curved and each Habitat Largely unknown but likely found singly or contains a delicate ring palisade and a slender gastro- in small patches on hard rock including bedrock, boul- style. The range of dactylopores per cyclosystem is ders, cobbles, and pebbles, in areas of moderate current, 11–17, but averages about 14; the dactylostyles are and at depths around 881 m. inconspicuous. Female ampullae are unknown. The male ampullae Photos A) The dried S. elassotomus holotype (USNM are small (0.5–0.6 mm in diameter) superficial hemi- 43268; large portion of a colony) collected southeast spheres. of Agattu Island, western Aleutian Islands, at a depth of 881 m. B) A close-up view of the same specimen in Remarks Stylaster elassotomus is distinctive among photo A. 374 Professional Paper NMFS 23 15. Stylaster elassotomus Fisher, 1938 (continued) Figure 16-21 A map of the Aleutian Islands showing the distribution of Stylaster elassotomus () in Alaska waters. Chapter 16 375 16. Stylaster leptostylus (Fisher, 1938) Description Colonies branch in a uniplanar and equal lar to S. campylecus, but differs in having fewer dacty- dichotomous manner; the largest known colony is 13 lopores per cyclosystem. cm in height. The coenosteum has a reticulate-granular texture and is white in color. Distribution Rare. In Alaska – known only from the The gastro- and dactylopores are arranged in circu- seven type specimens collected at the same location in lar cyclosystems on branch edges and the anterior face; Amukta Pass in the eastern Aleutian Islands (Fig. 16- the cyclosystems are 1.0–1.1 mm in diameter. The gas- 22). It is apparently endemic to the region. Elsewhere tropore tubes are slightly curved and funnel-shaped and – not reported. each houses a lanceolate gastrostyle; a ring palisade is absent. The range of dactylopores per cyclosystem is Habitat Largely unknown but likely found singly or 7–12, but averages about 10; the dactylostyles are of in small patches on hard rock including bedrock, boul- moderate size. ders, cobbles, and pebbles, in areas of moderate current, The female ampullae are smooth superficial hemi- and at depths around 518 m. spheres measuring 0.9–1.1 mm in diameter. The male ampullae are superficial mounds and measure 0.45–0.55 Photos A) The dried S. leptostylus holotype (USNM mm in diameter. 43270; large portion of a colony) collected in Amukta Pass, eastern Aleutian Islands, at a depth of 518 m. B) Remarks This species was originally described as a Large portions of dried S. leptostylus colonies (USNM form of Stylaster moseleyanus but later elevated to spe- 76817; paratypes) collected in Amukta Pass at a depth cies rank by Cairns and Lindner (2011). It is most simi- of 518 m. 376 Professional Paper NMFS 23 16. Stylaster leptostylus (Fisher, 1938) (continued) Figure 16-22 A map of the Aleutian Islands showing the distribution of Stylaster leptostylus () in Alaska waters. Chapter 16 377 17. Stylaster parageus columbiensis Cairns and Lindner, 2011 Description (Adapted from Cairns and Lindner, ters of Southeast Alaska (Fig. 16-23). Elsewhere – Brit- 2011) See description of the nominate subspecies ish Columbia to northern Washington. (Stylaster parageus parageus) and Remarks. Colonies are relatively small, only measuring up to 8.5 cm in Habitat In Alaska – typically occurs in patches, often height and 8 cm in width. The color of the coenosteum on rocky bedrock pinnacles, in areas of moderate to is white in life but turns a creamy orange in ethanol- strong current, and at depths between 92 and 439 m. preserved specimens. Observed on video footage collected on Shutter Ridge as shallow as 88 m (senior author, personal observ.). Remarks This subspecies differs from the nominate Elsewhere – reported at depths between 246 and 288 m. subspecies by having larger cyclosystems (1.0–1.5 mm in diameter), larger gastropores (0.45–0.50 mm in di- Photos A) Large portions of dried S. p. columbien- ameter), and a slightly higher number of dactylopores sis colonies (USNM 1291726) collected on Shutter per cyclosystem (6–13, average=9). It also ranges Ridge, eastern Gulf of Alaska, at a depth of 92 m. B) considerably further south (approximately 1400 km) A close-up view of the leftmost specimen in photo A. than the nominate subspecies (Cairns and Lindner, C) Stylaster p. columbiensis colonies photographed on 2011). Shutter Ridge at a depth of 88 m. D) A close-up view The coenosteum is composed entirely of aragonite of a colony in photo C (between the red laser marks). (senior author and S. D. Cairns, unpubl. data). E) A closer view of the same specimen in photo D. F) A patch of S. p. columbiensis on a rocky pinnacle on Shut- Distribution Common, abundant in some areas. In ter Ridge. The distance between the red lasers marks in Alaska – eastern Gulf of Alaska including the inside wa- photos C, D, and F is 10 cm. Figure 16-23 A map of the eastern Gulf of Alaska showing the distribution of Stylaster parageus columbiensis () in Alaska waters. 378 Professional Paper NMFS 23 17. Stylaster parageus columbiensis Cairns and Lindner, 2011 (continued) Chapter 16 379 18. Stylaster parageus parageus (Fisher, 1938) Description (Adapted from Cairns and Lindner, Remarks Originally described as the “southern shal- 2011) Colonies branch in a uniplanar or multiplanar low-water race” of S. campylecus by Fisher (1938), the manner, often with short branchlets oriented perpen- subspecies parageus was elevated to species rank by dicular to the flabella (or fans); branching is delicate Cairns and Lindner (2011). It is distinctive in having and dichotomous. Colonies are relatively large, measur- very small gastropores. ing up to 19 cm in height and 22 cm in width. Spionid The coenosteum is composed almost entirely (92– polychaete tubes are common. The coenosteal texture 100%) of aragonite with very small amounts of low- is reticulate-granular; the coenosteum is white to light magnesium calcite (senior author and S. D. Cairns, un- pink in color. publ. data). Studies of this species growth indicate that The gastro- and dactylopores are arranged in circu- the periodicity of skeletal growth bands is monthly, ra- lar cyclosystems on branch edges and the anterior face; dial growth rates are 1.4 mm/year (standard deviation the cyclosystems are 0.9–1.0 mm in diameter. The gas- [SD] 0.1), and axial growth rates are 17.3 mm/year (SD tropores are quite small (0.3 mm in diameter); the gas- 1.1) (Aranha, 2010). tropore tubes are cylindrical, slightly curved, and each contains a well-developed ring palisade and a lanceolate Distribution Common, locally abundant. In Alaska gastrostyle. The range of dactylopores per cyclosystem – eastern Gulf of Alaska including the inside waters of is 5–11 and averages about 8; the dactylostyles are well Southeast Alaska to south of Resurrection Bay in the developed. western Gulf of Alaska (Fig. 16-24). Elsewhere – north- The female ampullae are superficial hemispheres ern British Columbia near the border with Southeast (0.8–1.0 mm in diameter) and each has a lateral efferent Alaska. pore. The male ampullae are mostly internal and about 0.5 mm in diameter. Habitat In Alaska – typically occurs in patches, some- times dense, on rocky bedrock pinnacles and other hard Figure 16-24 A map of the North Pacific Ocean showing the distribution of Stylaster parageus parageus () in Alaska waters. 380 Professional Paper NMFS 23 rock areas of moderate to strong current, and at depths in Stephens Passage at a depth of 35 m. C) A close-up between 26 and 445 m. Elsewhere – reported at a depth view of the same specimen in photo A. D) A close-up of 23 m. view of the same specimen in photo B. E) A small patch of S. p. parageus on the Fairweather Ground, eastern Photos A) A dried whole S. p. parageus colony Gulf of Alaska, at a depth of 143 m. F) A small patch (USNM 1482020) collected in Stephens Passage, South- of S. p. parageus on the Fairweather Ground at a depth east Alaska, at a depth of 26 m. B) A large portion of a of 157 m. The distance between the red lasers marks in dried S. p. parageus colony (USNM 1482021) collected photos E and F is 10 cm. Chapter 16 381 19. Stylaster repandus Cairns and Lindner, 2011 Description (Adapted from Cairns and Lindner, growth form, but not unlike Cyclohelia lamellata and 2011) Colonies are firmly attached to hard substrate Errinopora undulata in this regard. by a robust stem, which gives rise to several undulat- The coenosteum is composed principally (98%) of ing plates that result in a three-dimensional structure. high-magnesium calcite (9.7 mol% MgCO3) with very The largest known colony is 23 cm in height and 19 cm small amounts of aragonite and low-magnesium calcite in width. The coenosteal texture is reticulate-granular; (senior author and S. D. Cairns, unpubl. data). the coenosteum is light pink and covered with small pa- pillae. Branches are heavily infested with spionid poly- Distribution In Alaska – rare, known from only the chaete tubes. four type specimens collected in the same area near The gastro- and dactylopores are arranged in round Amukta Pass in the eastern Aleutian Islands (Fig. 16- cyclosystems on both the faces and edges of plates, and 25). It is apparently endemic to the region. Elsewhere are sometimes arranged in short transverse rows on – not reported. plate faces. The gastropore tubes are funnel-shaped, each containing a diffuse ring palisade and a short lan- Habitat Largely unknown but likely found singly or ceolate gastrostyle. The range of dactylopores per cy- in small patches on hard rock including bedrock, boul- closystem is 1–11 and averages about 4. Diastemas are ders, cobbles, and pebbles, in areas of moderate current, common, the dactylostyles are robust, and the tall ele- and at depths between 375 m and about 650 m. ments are unilinear. The female ampullae are low swellings on the coenos- Photos A) A large portion of a dried S. repandus col- teum and 0.9–1.1 mm in diameter. The male ampullae ony (USNM 1122741; paratype) collected in Amukta are low mounds, primarily internal, and 0.5–0.6 mm in Pass, eastern Aleutian Islands, at a depth of approxi- diameter. mately 650 m. B) The dried S. repandus holotype (USNM 1122740; large portion of a colony) collected Remarks Stylaster repandus is easily distinguished in Amukta Pass at a depth of 375 m. C) A close-up view as being the only species of Stylaster with a lamellar of the same specimen in photo B. 382 Professional Paper NMFS 23 19. Stylaster repandus Cairns and Lindner, 2011 (continued) Figure 16-25 A map of the Aleutian Islands showing the distribution of Stylaster repandus () in Alaska waters. Chapter 16 383 20. Stylaster stejnegeri (Fisher, 1938) Description (Adapted from Cairns and Lindner, 2011) Remarks This species is similar to S. brochi but has The holotype is an arborescent colony measuring 6 cm fewer dactylopores per cyclosystem and ridged female in height and 7 cm in width with equal dichotomous ampullae. branching; the branches are blunt-tipped. The coenos- teum is reticulate-granular in texture and is covered with Distribution Extremely rare, known only from the small papillae (possibly nematopores). Binary spionid holotype collected on Petrel Bank (Fig. 16-26). It is polychaete tubes are present along the branch axes and apparently endemic to the region. Elsewhere – not re- the coenosteum is light orange to pink in color. ported. The gastro- and dactylopores are arranged in cyclo- systems that occur uniformly on all sides of branches; Habitat Largely unknown but likely found singly or the cyclosystems are 1.0–1.2 mm in diameter. The gas- in small patches on hard rock, including bedrock, boul- tropores are circular; the gastropore tubes are cylindri- ders, cobbles, and pebbles, in areas of moderate current, cal and slightly curved and each contains a lanceolate and at depths around 95 m. gastrostyle but no ring palisade. The range of dacty- lopores per cyclosystem is 5–11 and averages about 6; Photos A) The dried S. stejnegeri holotype (USNM dactylostyles are rudimentary. Supernumerary dactylo- 43271; large portion of a colony) collected on Petrel pores are common. Bank, central Aleutian Islands, at a depth of 87 m. B) A The female ampullae are superficial hemispheres up close-up view of the same specimen in photo A. to 1.1 mm in diameter and are often covered with low radiating ridges. Male ampullae are unknown. 384 Professional Paper NMFS 23 20. Stylaster stejnegeri (Fisher, 1938) (continued) Figure 16-26 A map of the Aleutian Islands showing the distribution of Stylaster stejnegeri () in Alaska waters. Chapter 16 385 21. Stylaster trachystomus (Fisher, 1938) Description (Adapted from Cairns and Lindner, campylecus, Stylaster trachystomus was raised to spe- 2011) Colony branching is uniplanar, in a delicate ir- cies level by Cairns and Lindner (2011). This species is regular dichotomous manner; the largest known colony similar to S. campylecus and S. brochi, as discussed by is 12 cm in height and 8 cm in width. The coenosteum Cairns and Lindner (2011). is reticulate-granular in texture with short longitudinal The coenosteum is composed almost entirely (89– ridges present on most branches. The coenosteum is 100%) of aragonite with very small amounts of low- pink-orange in color. magnesium calcite (Cairns and Macintyre, 1992; senior The gastro- and dactylopores are arranged in cyclo- author and S. D. Cairns, unpubl. data). systems that occur on all branch faces, with fewer on the posterior face. Cyclosystems are circular to ellipti- Distribution Uncommon (17 specimens total collect- cal with the greater axis up to 1.8 mm in length. The ed). In Alaska – Aleutian Islands (northern Amukta Pass gastropore tubes are cylindrical, slightly curved, quite long, and contain a well-developed ring palisade and an to near Attu Island) including Bowers Ridge (Fig. 16- elongate lanceolate gastrostyle. The range of dactylo- 27). Elsewhere – not reported. pores per cyclosystem is 8–18 and averages about 12; the dactylostyles are inconspicuous. Habitat Typically occurs singly or in small patches on The female ampullae are superficial hemispheres hard rock (bedrock and boulders), in areas of moderate that measure 1.0–1.3 mm in diameter and often have current, and at depths between 110 and 366 m. a knobby or papillose surface. The male ampullae are much smaller (0.50–0.55 mm in diameter) mounds and Photos A) A whole dried S. trachystomus colony both types are clustered on anterior branch faces. (USNM 1481964) collected near the Delarof Islands, central Aleutian Islands, at a depth of 110 m. B) A Remarks Originally described as a subspecies of S. close-up view of the same specimen in photo A. 386 Professional Paper NMFS 23 21. Stylaster trachystomus (Fisher, 1938) (continued) Figure 16-27 A map of the Aleutian Islands showing the distribution of Stylaster trachystomus () in Alaska waters. Chapter 16 387 22. Stylaster venustus (Verrill, 1868) Description (Adapted from Fisher, 1938) Colonies magnesium calcite (senior author and S. D. Cairns, un- are small (rarely more than 7 cm in height), planar to publ. data). bushy, dichotomously branched, and have blunt branch tips. The coenosteum is reticulate-granular in texture Distribution Rare. In Alaska – known from only a and violet or pink in color; coenosteal papillae are com- single specimen collected near Cape Ommaney, Baranof mon. Internal spionid polychaete tubes are present. Island, in the eastern Gulf of Alaska (Fig. 16-28). This The gastro- and dactylopores are arranged in cyclo- specimen (see photo A) represents the northernmost re- systems that occur uniformly on all branch surfaces. cord for the species. Elsewhere – British Columbia to The cyclosystems are polygonal and 0.7–0.9 mm in di- central California (Big Sur). ameter. The gastropore tubes are straight and funnel- shaped, each containing a prominent ring palisade and Habitat In Alaska – largely unknown but likely found a lanceolate gastrostyle. The range of dactylopores per in small patches on hard rock, including bedrock, boul- cyclosystem is only four to eight and averages about six; ders, cobbles, and pebbles, in areas of moderate current the dactylostyles are robust. and tidal surge, and at shallow depths (around 20 m). The female ampullae are entirely internal and about Elsewhere – specimens have been collected at depths 0.6 mm in diameter. The male ampullae are also inter- between 4 m (southern British Columbia) and 117 m nal, quite abundant, and about 0.40 mm in diameter. (Heceta Bank, off Oregon). Remarks Stylaster venustus is most similar to S. ver- Photos A) Portions of dried S. venustus colonies rillii (Dall, 1884), but has larger cyclosystems, smaller (USNM 1482062) collected near Cape Ommaney, Ba- branches, and differs in the color of the coenosteum ranof Island, eastern Gulf of Alaska, at a depth of 20 (see Cairns and Lindner, 2011, p. 48, table 2, figure m. B) Four whole dried S. venustus colonies (USNM 16). The coenosteum is composed almost entirely 76544) collected in the Strait of Juan de Fuca, Washing- (96%) of aragonite with very small amounts of low- ton, at a depth of 73 m. 388 Professional Paper NMFS 23 22. Stylaster venustus (Verrill, 1868) (continued) Figure 16-28 A map of the eastern Gulf of Alaska showing the distribution of Stylaster venustus () in Alaska waters. Chapter 16 389 23. Stylaster verrillii (Dall, 1884) Description (Adapted from Cairns and Lindner, 2011) of that species above. The reproductive ecology of this Colonies are arborescent and dichotomously branched; species has been studied from colonies collected in the the branches have blunt to clavate tips. The largest Aleutian Islands (Brooke and Stone, 2007). known colony is only 3.5 cm in height and 7 cm in The skeleton (coenosteum) is composed principal- width. The coenosteum is reticulate-granular in texture ly (96.2–99.7%) of high-magnesium calcite (9.0–9.4 and coenosteal papillae are common. Spionid polychaete mol% MgCO3) with very small amounts of aragonite worm tubes are also common. The coenosteum is light and low- and high-magnesium calcite (senior author orange to bright pink in color. and S. D. Cairns, unpubl. data). The gastro- and dactylopores are arranged in cyclo- systems that occur uniformly on all branch surfaces; the Distribution Common, abundant in some areas. In cyclosystems are round and 1.0–1.2 mm in diameter. Alaska – this species exhibits an apparent disjunct dis- The gastropore tubes are elongate and cylindrical, and tribution in Alaska. Collected from the eastern Gulf of each contains a well-developed ring palisade and an Alaska including the inside waters of Southeast Alaska elongate spiny gastrostyle. The range of dactylopores to northwest of Portlock Bank in the western Gulf of per cyclosystem is 5–10 and averages about 7; the dac- Alaska and the Aleutian Islands (Akutan Pass to near tylostyles are robust. Kiska Island; Fig. 16-29). Elsewhere – northern Brit- The female ampullae are internal and 0.7–0.8 mm ish Columbia near the border with Southeast Alaska to in diameter. The male ampullae are also internal and the San Juan Islands of Washington. There is a single 0.3–0.5 mm in diameter. record from the Sea of Okhotsk, Russia. Remarks This species was originally described as Habitat In Alaska – typically occurs in patches, some- Allopora moseleyi Dall, 1884. It is most similar to S. times dense, on rocky bedrock pinnacles and other hard venustus and compared to that species in the account rock, including cobbles and pebbles, in areas of moderate Figure 16-29 A map of the North Pacific Ocean showing the distribution of Stylaster verrillii () in Alaska waters. 390 Professional Paper NMFS 23 to strong current, and at depths between 11 and 401 m. fet, Adak Island, central Aleutian Islands, at a depth of Elsewhere – reported at depths between 23 and 183 m. 155 m. C) A S. verrillii colony photographed on Shut- ter Ridge, eastern Gulf of Alaska, at a depth of 88 m. Photos A) A dried whole S. verrillii colony (USNM D) A small patch of S. verrillii photographed on Shut- 1482150) collected near Bobrof Island, central Aleutian ter Ridge at a depth of 143 m. The distance between Islands, at a depth of 17 m. B) A dried whole S. verril- the red laser marks is 10 cm. E) A close-up view of one lii colony (USNM 1027819) collected near Cape Mof- colony in photo D. Acknowledgments 391 Acknowledgments We thank J. Heard for preparing hundreds of photo- were provided by J. Orr, M. Boyd, C. Piotrowski, G. graphs for this publication and B. Wing, C. O’Clair, Hendler, E. Kools, L. Lundsten, B. Voss, G. Keel, H. J. Karinen, D. Carlson, S. France, L. Watling, K. Iken, Gartner, A. Mickle, S. McCue, and V. Delnavaz. We P. Etnoyer, A. Baco-Taylor, T. Marshall, K. Lowyck, thank Cara Mayo and Shelley Arenas for skillful edit- J. Hocevar, M. Ridgway, and countless others for as- ing and publication advice and two anonymous referees sistance with the collection of specimens. F. Sinniger for helpful reviews of the manuscript. Field collections Harri contributed substantially to the Zoantharia chap- were supported by NOAA Fisheries (Alaska Fisheries ter; J. Thoma, B. Horvath, and M. Everett shared taxo- Science Center), NOAA Undersea Research Program nomic insights; and A. Helbling and A. Cohen provided (West Coast and Polar Regions Undersea Research laboratory analyses on calcium carbonate mineralogy. Center), NOAA Ocean Exploration, the North Pacific D. Csepp, T. Warshaw, D. Harris, N. McDaniel, M. Research Board, and Greenpeace International. Fund- Chamberlain, L. Barr, P. Malecha, D. Geiger, and R. ing for the publication of this guide was provided by Langton provided in situ, topside, and laboratory pho- NOAA’s Alaska Fisheries Science Center and Deep Sea tographs. Specimen metadata and laboratory assistance Coral Research and Technology Program. 392 Professional Paper NMFS 23 Literature cited Andrews, A. H., E. E. Cordes, M. M. Mahoney, K. Munk, K. H. 1952b. Two new species of Arthrogorgia (Gorgonacea: Prim- Coale, G. M. Cailliet, and J. Heifetz. noidae) from the Aleutian Islands region. Proc. Biol. Soc. 2002. Age, growth and radiometric age validation of a deep- Wash. 65:63–70. sea, habitat-forming gorgonian (Primnoa resedaeformis) 1956. Descriptions and redescriptions of the Hawaiian octo- from the Gulf of Alaska. Hydrobiologia 471:101–110. corals collected by the U. S. Fish Commission steamer “Al- https://doi.org/10.1023/A:1016501320206 batross” (2. Gorgonacea: Scleraxonia). Pac. Sci. 10:67–95. Andrews, A. H., G. M. Cailliet, L. A. Kerr, K. H. Coale, C. Lund- 1981. Key to the genera of Octocorallia exclusive of Pennatu- strom, and A. P. DeVogelaere. lacea (Coelenterata: Anthozoa), with diagnoses of new 2005. Investigations of age and growth for three deep-sea cor- taxa. Proc. Biol. Soc. Wash. 94:902–947. als from the Davidson Seamount off central California. In 1990. A new isidid octocoral (Anthozoa: Gorgonacea) from Cold-water corals and ecosystems (A. Freiwald and J. M. New Caledonia, with descriptions of other new species Roberts, eds.), p. 1021–1038. Springer-Verlag, Berlin, Ger- from elsewhere in the Pacific Ocean. Proc. Biol. Soc. Wash. many. 103:205–228. Andrews, A. H., R. P. Stone, C. C. Lundstrom, and A. P. DeVoge- 1996. The gorgonacean genus Arthrogorgia (Octocorallia: laere. Primnoidae). Proc. Biol. Soc. Wash. 109:605–628. 2009. Growth rate and age determination of bamboo corals Bayer, F. M., and J. Stefani. from the northeastern Pacific Ocean using refined 210Pb 1987. New and previously known taxa of isidid octocorals dating. Mar. Ecol. Prog. Ser. 397:173–185. https://doi. (Coelenterata: Gorgonacea), partly from Antarctic waters. org/10.3354/meps08193 Proc. Biol. Soc. Wash. 100:937–991. Aranha, R. Bayer, F. M., M. Grasshoff, and J. Verseveldt. 2010. Geochemistry, microstructure and growth banding in 1983. Illustrated trilingual glossary of morphological and Stylaster campylecus parageus and Primnoa pacifica: im- anatomical terms applied to Octocorallia, 75 p. E. J. Brill, plications for commonly observed deep sea corals as pa- Leiden, Netherlands. leoceanographic archives. M.S. thesis, 264 p. Meml. Univ. Boutillier, P. D. G., J. A. Boutillier, and G. E. Gillespie. Nfld., St. John’s, NL, Canada. [Available from http://re- 2019. British Columbia corals: a synopsis of information on search.library.mun.ca/id/eprint/9617.] their taxonomy, occurrences, distribution, threats and gen- Aranha, R., E. Edinger, G. Layne, and G. Piercey. eral status. Can. Tech. Rep. Fish. Aquat. Sci. 3322, 260 p. 2014. Growth rate variation and potential paleoceanographic Bramanti, L., G. Magagnini, L. De Maio, and G. Santangelo. proxies in Primnoa pacifica: insights from high-resolution 2005. Recruitment, early survival and growth of the Medi- trace element microanalysis. Deep-Sea Res., II 99:213–226. terranean red coral Corallium rubrum (L. 1758), a 4-year https://doi.org/10.1016/j.dsr1012.2013.1007.1001 study. J. Exp. Mar. Biol. Ecol. 314:69–78. https://doi. Ardila, N. E., G. Giribet, and J. A. Sánchez. org/10.1016/j.jembe.2004.08.029 2012. A time-calibrated molecular phylogeny of the precious Breedy, O., S. D. Cairns, and V. Häussermann. corals: reconciling discrepancies in the taxonomic classifi- 2015. A new alcyonacean octocoral (Cnidaria, Anthozoa, Oc- cation and insights into their evolutionary history. BMC tocorallia) from Chilean fjords. Zootaxa 3919:327–334. Evol. Biol. 12:246. https://doi.org/10.1186/1471-2148-12- https://doi.org/10.11646/zootaxa.3919.2.5 246 Broch, H. Aurivillius, M. 1932. Über einige geographisch interessante Fundstellen von 1931. The gorgonarians from Dr. Sixten Bock’s expedition to Alcyonarien und Hydrokorallen im nördlichen Stillen Oz- Japan and Bonin Islands 1914. Kungliga Svenska Vetens- ean. Explor. Mers U. R. S. S., Leningrad 17:81–86. kapsAkademiens Handlingar, Tredje Serien 9:1–337. 1935. Oktokorallen des Nördlichsten Pazifischen Ozeans und Baco, A. R. ihre Beziehungen zur Atlantischen Fauna. Avhandlinger 2007. Exploration for deep-sea corals on North Pacific sea- utgitt av det Norske Videnskaps-Akaademi 1935(1):1–53. mounts and islands. Oceanography 20:108–117. https:// 1957. The northern octocoral, Paragorgia arborea (L.), doi.org/10.5670/oceanog.2007.11 in sub-Antarctic waters. Nature 179:1356. https://doi. Baco, A. R., and T. M. Shank. org/10.1038/1791356a0 2005. Population genetic structure of the Hawaiian precious Brodeur, R. D. coral Corallium lauuense (Octocorallia: Coralliidae) using 2001. Habitat-specific distribution of Pacific ocean perch microsatellites. In Cold-water corals and ecosystems (A. (Sebastes alutus) in Pribilof Canyon, Bering Sea. Cont. Freiwald and J. M. Roberts, eds.), p. 663–678. Springer- Shelf Res. 21:207–224. https://doi.org/10.1016/S0278- Verlag, Berlin, Germany. 4343(00)00083-2 Barr, L., and N. Barr. Brook, G. 1983. Under Alaskan seas: the shallow water marine inverte- 1889. Report on the Antipatharia collected by H.M.S. Chal- brates, 208 p. Alsk. Northwest Publ. Co., Anchorage, AK. lenger during the years 1873–76. In Report on the scien- Bayer, F. M. tific results of the voyage of the H.M.S. Challenger during 1952a. Descriptions and redescriptions of the Hawaiian oc- the years 1873–76. Zoology, vol. 32, part 80, p. 1–222. tocorals collected by the U. S. Fish Commission Steamer Neill and Co., Edinburgh, Scotland. “Albatross.” (1. Alcyonacea, Stolonifera, and Telestacea). Brooke, S., and R. P. Stone. Pac. Sci. 4:126–136. 2007. Reproduction of deep-water hydrocorals (family Literature Cited 393 Stylasteridae) from the Aleutian Islands, Alaska. Bull. Mar. 2009. A generic revision and phylogenetic analysis of the Sci. 81:519–532. Primnoidae (Cnidaria: Octocorallia). Smithson. Contrib. Busby, M. S., J. W. Orr, and D. M. Blood. Zool. 629, 79 p. 2006. Eggs and late-stage embryos of Allocareproctus unan- Cairns, S. D., and M. V. Kitahara. gas (family Liparidae) from the Aleutian Islands. Ichthyol. 2012. An illustrated key to the genera and subgenera of the Res. 53:423–426. https://doi.org/10.1007/s10228-006- Recent azooxanthellate Scleractinia (Cnidaria, Anthozoa), 0361-3 with an attached glossary. ZooKeys 227:1–47. https://doi. Cairns, S. D. org/10.3897/zookeys.227.3612 1981. Marine flora and fauna of the northeastern United Cairns, S. D., and A. Lindner. States. Scleractinia. NOAA Tech. Rep. NMFS Circ. 438, 2011. A revision of the Stylasteridae (Cnidaria, Hydrozoa, Fi- 14 p. lifera) from Alaska and adjacent waters. ZooKeys 158:1– 1982. Antarctic and Subantarctic Scleractinia. Antarct. Res. 88. https://doi.org/10.3897/zookeys.158.1910 Ser. 34, 74 p. Cairns, S. D., and I. G. Macintyre. 1991. Cyclohelia lamellata, new genus and species of Stylaster- 1992. Phylogenetic implications of the calcium carbonate idae (Cnidaria: Hydrozoa) from the Bering Sea. Pac. Sci. mineralogy in the Stylasteridae (Cnidaria: Hydrozoa). 45:383–388. Palaios 7:96–107. https://doi.org/10.2307/3514799 1994. Scleractinia of the temperate North Pacific. Smithson. Cairns, S. D., and M. L. Taylor. Contrib. Zool. 557, 150 p. 2019. An illustrated key to the species of the genus Narella 1995. The marine fauna of New Zealand: Scleractinia (Cni- (Cnidaria, Octocorallia, Primnoidae). ZooKeys 822:1–15. daria: Anthozoa). N. Z. Oceanogr. Inst. Mem. 103, 139 p. https://doi.org/10.3897/zookeys.822.29922 2001. Studies on western Atlantic Octocorallia (Coelenterata: Cairns, S. D., and H. H. Wirshing. Anthozoa). Part 1. The genus Chrysogorgia Duchassaing 2018. A phylogenetic analysis of the Primnoidae (Anthozoa: and Michelotti, 1864. Proc. Biol. Soc. Wash. 114:746–787. Octocorallia: Calcaxonia) with analyses of character evo- 2004. A new shallow-water species of Javania (Scleractinia: lution and a key to the genera and subgenera. BMC Evol. Flabellidae) from Indonesia. Raffles Bull. Zool. 52:7–10. Biol. 18:66. https://doi.org/10.1186/s12862-018-1182-5 2007. Calcaxonian octocorals (Cnidaria: Anthozoa) from the Cairns, S. D., R. P. Stone, H.-W. Moon, and J. H. Lee. Eastern Pacific seamounts. Proc. Calif. Acad. Sci. 58:511– 2018. Primnoidae (Octocorallia: Calcaxonia) from the 541. Emperor Seamounts, with Notes on Callogorgia el- 2010. A review of the Octocorallia (Cnidaria: Anthozoa) from egans (Gray, 1870). Pac. Sci. 72:125–142. https://doi. Hawai´i and adjacent seamounts. Part 3: genera Thouarel- org/10.2984/72.1.8 la, Plumarella, Callogorgia, Fanellia, and Parastenella. Pac. Cairns, S. D., R. T. S. Cordeiro, Y. Xu, Z. Zhan, and P. A. Alder- Sci. 64:413–440. https://doi.org/10.2984/64.3.413 slade. 2011a. A revision of the Primnoidae (Octocorallia: Alcyona- 2021. A new family and two new genera of calcaxonian oc- cea) from the Aleutian Islands and Bering Sea. Smithson. tocoral, including a redescription of Pleurogorgia militaris Contrib. Zool. 634, 55 p. (Cnidaria: Octocorallia: Chrysogorgiidae) and its place- 2011b. Global diversity of the Stylasteridae (Cnidaria: Hy- ment in a new genus. Invert. Syst. 35:282–297. https://doi. drozoa: Anthecatae). PLoS ONE 6(7):e21670. https://doi. org/10.1071/IS20066 org/10.1371/journal.pone.0021670 Chaytor, J. D., R. A. Keller, R. A. Duncan, and R. P. Dziak. 2015. Tropical deep-sea benthos volume 28: Stylasteridae 2007. Seamount morphology in the Bowie and Cobb hotspot (Cnidaria: Hydrozoa: Anthoathecata) of the New Caledo- trails, Gulf of Alaska. Geochem. Geophys. Geosyst. nian region. Mem. Mus. Natl. Hist. Nat., Paris 207, 362 p. 8:Q09016. https://doi.org/10.1029/2007GC001712 2016. The marine fauna of New Zealand: primnoid octo- Choy, E., K. Watanabe, B. Williams, R. Stone, P. Etnoyer, E. corals (Anthozoa, Alcyonacea) — Part 2. Primnoella, Cal- Druffel, T. Lorenson, and M. Knaak. lozostron, Metafannyella, Callogorgia, Fanellia and other 2020. Understanding growth and age of red tree cor- genera. NIWA Biodivers. Mem. 129, 131 p. als (Primnoa pacifica) in the North Pacific Ocean. PLoS 2018. Deep-water octocorals (Cnidaria, Anthozoa) from ONE 15(12):e0241692. https://doi.org/10.1371/journal. the Galápagos and Cocos Islands. Part 1. Suborder Cal- pone.0241692 caxonia. ZooKeys 729:1–46. https://doi.org/10.3897/zoo- CITES (Convention on International Trade in Endangered Spe- keys.729.21779 cies). 2021. The marine fauna of New Zealand: primnoid octo- 2007. Convention on international trade in endangered spe- corals (Anthozoa, Alcyonacea) — Part 3. Thouarella, and cies of wild fauna and flora. Consideration of proposals additional records of other primnoid species. NIWA Bio- for amendment of appendices I and II. Fourteenth meeting divers. Mem. 133, 67 p. of the conference of the parties. The Hague (Netherlands), Cairns, S. D., and A. Baco. 3–15 June, CoP14 prop. 21, 22 p. [Available from https:// 2007. Review and five new Alaskan species of the deep- cites.org/sites/default/files/eng/cop/14/prop/E14-P21.pdf.] water octocoral Narella (Octocorallia: Primnoidae). Cordeiro, R. T. S., S. D. Cairns, and C. D. Pérez. Syst. Biodivers. 5:391–407. https://doi.org/10.1017/ 2017. A revision of the genus Radicipes Stearns,1883 (Antho- s1477200007002472 zoa: Octocorallia: Chrysogorgiidae). Zootaxa 4319:1–26. Cairns, S. D., and F. M. Bayer. https://doi.org/10.11646/zootaxa.4319.1.1 2005. A review of the genus Primnoa (Octocorallia: Gorgona- Cordes, E. E., J. W. Nybakken, and G. Van Dykhuizen. cea: Primnoidae), with the description of two new species. 2001. Reproduction and growth of Anthomastus ritteri (Oc- Bull. Mar. Sci. 77:225–256. tocorallia: Alcyonacea) from Monterey Bay, California, 394 Professional Paper NMFS 23 USA. Mar. Biol. 138:491–501. https://doi.org/10.1007/ Ehrenberg, C. G. s002270000470 1834. Beiträge zur physiologischen Kenntniss der Corallen- Cutress, C. E., and W. E. Pequegnat. thiere im allgemeinen, und besonders des rothen Meeres, 1960. Three new species of Zoantharia from California. Pac. nebst einem Versuche zur physiologischen Systematik der- Sci. 14:89–100. selben. Abh. Königlichen Akad. Wiss., Berlin. 1:225–380. Dall, W. H. Esper, E. J. C. 1884. On some Hydrocorallinae from Alaska and California. 1794. Fortsetzungen der Pflanenthiere, vol. 1, parts 1–2, 64 Proc. Biol. Soc. Wash. 2:111–115. p., Nürnberg. Daly, M., M. R. Brugler, P. Cartwright, A. G. Collins, M. N. Etnoyer, P. J. Dawson, D. G. Fautin, S. C. France, C. S. McFadden, D. M. 2008. A new species of Isidella bamboo coral (Octocoral- Opresko, E. Rodriguez, et al. lia: Alcyonacea: Isididae) from northeast Pacific sea- 2007. A review of phylogenetic patterns and diversity 300 mounts. Proc. Biol. Soc. Wash. 121:541–553. https://doi. years after Linnaeus. Zootaxa 1668:127–182. https://doi. org/10.2988/08-16.1 org/10.11646/zootaxa.1668.1.11 Etnoyer, P., S. D. Cairns, J. A. Sanchez, J. K. Reed, J. V. Lopez, W. Danielssen, D. C. W. Schroeder, S. D. Brooke, L. Watling, A. Baco-Taylor, G. C. 1860. Beskrivelse over en ny Art Virgularia [meeting report]. Williams, et al. Forhandlinger i Videnskabs-selskabet i Christiania. 1859, 251 p. 2006. Deep-sea coral collection protocols. NOAA Tech. Dautova, T. N. Memo. NMFS-OPR-28, 53 p. 2007. Gorgonians (Anthozoa: Octocorallia) of the northwest- Everett, M. V., L. K. Park, E. A. Berntson, A. E. Elz, C. E. Whit- ern Sea of Japan. Russ. J. Mar. Biol. 33:297–304. https:// mire, A. A. Keller, and M. E. Clarke. doi.org/10.1134/S1063074007050045 2016. Large-scale genotyping-by-sequencing indicates high 2018. Two new species of deep-water Calcigorgia gorgoni- levels of gene flow in the deep-sea octocoral Swiftia sim- ans (Anthozoa: Octocorallia) from the Kurile Islands, Sea plex (Nutting 1909) on the West Coast of the United States of Okhotsk, with a review of the distinctive characters of PLoS ONE 11(10):e0165279. https://doi.org/10.1371/ the known species of the genus. Eur. J. Taxon. 408:1–22. journal.pone.0165279 https://doi.org/10.5852/ejt.2018.408 Fabricius, K., and P. Alderslade. Desonie, D. L., and R. A. Duncan. 2001. Soft corals and sea fans: a comprehensive guide to the 1990. The Cobb-Eikelberg seamount chain: hotspot vol- tropical shallow-water genera of the central-west Pacific, canism with mid-ocean ridge basalt affinity. J. Geo- the Indian Ocean and the Red Sea, 264 p. Aust. Inst. Mar. phys. Res. 95:12697–12711. https://doi.org/10.1029/ Sci., Townsville, Australia. JB095iB08p12697 Feehan, K. A., and R. G. Waller. Dolan, E. 2015. Notes on reproduction of eight species of Eastern Pacific 2008. Phylogenetics, systematics and biogeography of deep- cold-water octocorals. J. Mar. Biol. Assoc. UK 95(4):691– sea Pennatulacea (Anthozoa: Octocorallia): evidence from 696. https://doi.org/10.1017/S0025315415000053 molecules and morphology. Ph.D. diss., 195 p. Univ. South- Fisher, W. K. ampton, Southampton, UK. [Available from https://eprints. 1931. Californian hydrocorals. Ann. Mag. Nat. Hist., ser. 10, soton.ac.uk/65669/.] 8(46):391–399. Duchassaing, P., and J. Michelotti. 1938. Hydrocorals of the North Pacific Ocean. Proc. U. 1864. Supplément au mémoire sur les coralliaires des Antilles, S. Natl. Mus. 84:493–554. https://doi.org/10.5479/ ser. 2, vol. 23, 112 p. Mém. Acad. Sci. Turin (2)23:97–206. si.00963801.84-3024.493 Dueñas, L. F., P. Alderslade, and J. A. Sánchez. Flint, H. C., R. G. Waller, and P. A. Tyler. 2014. Molecular systematics of the deep-sea bamboo corals 2007. Reproductive ecology of Fungiacyathus marenzelleri (Octocorallia: Isididae: Keratoisidinae) from New Zealand from 4100 m depth in the northeast Pacific Ocean. Mar. with descriptions of two new species of Keratoisis. Mol. Biol. 151:843–849. https://doi.org/10.1007/s00227-006- Phylogenet. Evol. 74:15–28. https://doi.org/10.1016/j. 0539-2 ympev.2014.01.031 Försterra, G., and V. Häussermann. Du Preez, C., and V. Tunnicliffe. 2003. First report on large scleractinian (Cnidaria: Anthozoa) 2011. Shortspine thornyhead and rockfish (Scorpaenidae) distribution in response to substratum, biogenic structures accumulations in cold-temperate shallow water of south and trawling. Mar. Ecol. Prog. Ser. 425:217–231. https:// Chilean fjords. Zool. Verh. 345:117–128. doi.org/10.3354/meps09005 France, S. C. Durham, J. W. 2007. Genetic analysis of bamboo corals (Cnidaria: Octo- 1947. Corals from the Gulf of California and the North Pa- corallia: Isididae): does lack of colony branching distin- cific coast of America. Geol. Soc. Am. Mem. 20, 68 p. guish Lepidisis from Keratoisis? Bull. Mar. Sci. 81:323–333. Durham, J. W., and J. L. Barnard. Fujita, T., and S. Ohta. 1952. Stony corals of the eastern Pacific collected by the Vele- 1988. Photographic observations of the life style of the deep- ro III and Velero IV. Allan Hancock Pac. Exped. 16, 110 p. sea ophiuroid Asteronyx loveni (Echinodermata). Deep- Eckelbarger, K. J., P. A. Taylor, and R. W. Langton. sea Res. Oceanogr. Res., A 35:2029–2043. https://doi. 1998. Gonadal morphology and gametogenesis in the sea org/10.1016/0198-0149(88)90123-9 pen Pennatula aculeata (Anthozoa: Pennatulacea) from Goddard, P., R. Wilborn, C. Rooper, K. Williams, R. Towler, M. the Gulf of Maine. Mar. Biol. 132:677–690. https://doi. Sigler, and P. Malecha. org/10.1007/s002270050432 2016. Results of the 2014 underwater camera survey of the Literature Cited 395 eastern Bering Slope and Outer Shelf. NOAA Tech. Memo. Hoff, G. R., and B. Stevens. NMFS-AFSC-313, 304 p. 2005. Faunal assemblage structure of the Patton Seamount Goldsmith, D. B., C. A. Kellogg, C. L. Morrison, M. A. Gray, R. P. (Gulf of Alaska, USA). Alsk. Fish. Res. Bull. 11:27–36. Stone, R. G. Waller, S. D. Brooke, and S. W. Ross. Horowitz, W. L. 2018. Comparison of microbiomes of cold-water corals Prim- 2002. Evaluation of sub-sea physical environmental data for noa pacifica and Primnoa resedaeformis, with possible the Beaufort Sea OCS and incorporation into a Geograph- link between microbiome composition and host genotype. ic Information System (GIS) database. U.S. Dept. Interior, Sci. Rep. 8:12383. https://doi.org/10.1038/s41598-018- Miner. Manag. Serv., Alsk. Cont. Shelf Reg. OCS Study 30901-z MMS 2002-017, 66 p. [Available from https://www.boem. Gray, J. E. gov/sites/default/files/boem-newsroom/Library/Publica- 1860. II. Revision of the family Pennatulidae, with descrip- tions/2002/2002-017.pdf.] tions of some new species in the British Museum. Ann. Horvath, E. A. Mag. Nat. Hist., Zool. Bot. Geol., Series 3. 5(25–30):20– 2019. A review of gorgonian coral species (Cnidaria, Octo- 25. corallia, Alcyonacea) held in the Santa Barbara Museum of 1867. Additional note on Corallium johnsoni. Proc. Zool. Soc. Natural History research collection: focus on species from London. 1867:125–127. Scleraxonia, Holaxonia, Calcaxonia – Part III: suborder 1870. Catalogue of lithophytes or stony corals in the collec- Holaxonia continued, and suborder Calcaxonia. ZooKeys tion of the British Museum, 51 p. Order of the Trustees, 860:183–306. https://doi.org/10.3897/zookeys.860.34317 London. Horvath, E. A., and R. P. Stone. Gray, M. A., R. P. Stone, M. R. McLaughlin, and C. A. Kellogg. 2018. Another unusual new gorgonian (Anthozoa: Octo- 2011. Microbial consortia of gorgonian corals from the Aleu- corallia: Plexauridae) from the Aleutian Islands of Alaska. tian Islands. FEMS Microbiol. Ecol. 76:109–120. https:// Zootaxa 4524:112–120. https://doi.org/10.11646/zoo- doi.org/10.1111/j.1574-6941.2010.01033.x taxa.4524.1.8 Grigg, R. W. Johnson, T. 2002. Precious corals in Hawaii: discovery of a new bed and 2003. The Bering Sea and Aleutian Islands: region of wonders. revised management measures for existing beds. Mar. Fish. Alsk. Sea Grant Coll. Program, SG-ED-42, 199 p. Univ. Rev. 64(1):13–20. Alaska Fairbanks, Fairbanks, AK. Hartill, E. C., R. G. Waller, and P. J. Auster. Johnstone, J. W., R. G. Waller, and R. P. Stone. 2020. Deep coral habitats in Glacier Bay National Park and 2021. Shallow-emerged coral may warn of deep-sea coral re- sponse to thermal stress. Sci. Rep. 11:22439. https://doi. Preserve, Alaska. PLoS ONE 15(8):e0236945. https://doi. org/10.1038/s41598-021-01948-2 org/10.1371/journal.pone.0236945 Keller, N. B. Heifetz, J. 1977. New species of the genus Leptopenus and some pecu- 2002. Coral in Alaska: distribution, abundance, and spe- liarities of deep-sea ahermatypic corals. Tr. Inst. Okeanol. cies associations. Hydrobiologia 471:19–28. https://doi. 108:37–43. [In Russian.] org/10.1023/A:1016528631593 Kinoshita, K. Henry, L.-A., E. L. R. Kenchington, and A. Silvaggio. 1907. Vorläufige Mitteilung über einige neue japanische Prim- 2003. Effects of mechanical experimental disturbance on as- noid-Korallen. Annot. Zool. Jpn. 6:229–237. pects of colony responses, reproduction, and regeneration 1913. Beiträge zur Kenntnis der Morphologie und Stam- in the cold-water octocoral Gersemia rubiformis. Can. J. mesgeschichte der Gorgoniden. J. Coll. Sci. Univ. Tokyo Zool. 81:1691–1701. https://doi.org/10.1139/z03-161 32:1–50. https://doi.org/10.15083/00037767 Herrera, S., and T. M. Shank. Kitahara, M. V., S. D. Cairns, and D. J. Miller. 2016. RAD sequencing enables unprecedented phylogenetic 2010. Monophyletic origin of Caryophyllia (Scleractinia, resolution and objective species delimitation in recalcitrant Caryophylliidae), with descriptions of six new spe- divergent taxa. Mol. Phylogenet. Evol. 100:70–79. https:// cies. Syst. Biodivers. 8:91–118. https://doi.org/10.1080/ doi.org/10.1016/j.ympev.2016.03.010 14772000903571088 Herrera, S., A. Baco, and J. A. Sánchez. Kölliker, A. von. 2010. Molecular systematics of the bubblegum coral genera 1870. Anatomisch-systematische Beschreibung der Alcyonar- (Paragorgiidae, Octocorallia) and description of a new ien. Erste Abtheilung: Die Pennatuliden. Abhandlungen der deep-sea species. Mol. Phylogenet. Evol. 55:123–135. Senckenbergischen Naturforschenden Gesellschaft 7:487– https://doi.org/10.1016/j.ympev.2009.12.007 602. Herrera, S., T. M. Shank, and J. A. Sánchez. 1872. Anatomisch-systematische Beschreibung der Alcyonar- 2012. Spatial and temporal patterns of genetic variation in ien, Erste Abtheilung: Die Pennatuliden. Abhandlungen der the widespread antitropical deep-sea coral Paragorgia ar- Senckenbergischen Naturforschenden Gesellschaft 8:85– borea. Mol. Ecol. 21:6053–6067. https://doi.org/10.1111/ 275, plates 18–24. mec.12074 1875. Die Pennatulide Umbellula und zwei neue Typen der Al- Hickson, S. J. cyonarien. Festschrift zur Feier des fünfundzwanzigjähri- 1915. Some Alcyonaria and a Stylaster from the west coast gen Bestehens der Physicalisch-medicinischen Gesellschaft of North America. Proc. Zool. Soc. Lond. 85:541–557. in Würzburg, 23 p., plates 1–2. https://doi.org/10.1111/j.1469-7998.1915.00541.x 1880. Report on the Pennatulida dredged by H.M.S. Chal- 1921. On some Alcyonaria in the Cambridge Museum. Proc. lenger during the years 1873–1876. In Report on the sci- Camb. Philos. Soc., Math. Phys. Sci. 20:366–373. entific results of the voyage of H.M.S. Challenger during 396 Professional Paper NMFS 23 the years 1873–76. Zoology, vol. 1, part 2 (prepared by C. Malecha, P. W., and R. P. Stone. W. Thomson), 41 p., 11 plates. Neill and Co., Edinburgh, 2009. Response of the sea whip Halipteris willemoesi to simu- Scotland. lated trawl disturbance and its vulnerability to subsequent Koren, J., and D. C. Danielssen. predation. Mar. Ecol. Prog. Ser. 388:197–206. https://doi. 1877. Contribution to the natural history of the Pennatuli- org/10.3354/meps08145 dae living on the Norwegian coast. Fauna Littoralis Nor. Marshall, A. M. 3:82–102, plates 3–4, 9–12. 1883. Report on the Pennatulida dredged by H.M.S. “Triton.” 1883. Nye Alcyonider, Gorgonider og Pennatulider tilhorende Trans. R. Soc. Edinb. 32:119–152. Norges Fauna. Bergens Mus. Skr. 2:1–38, plates 1–3. Masuda, M. M., and R. P. Stone. Krieger, K. J. 2015. Bayesian logistic mixed-effects modelling of transect 2001. Coral (Primnoa) impacted by fishing gear in the Gulf of data: relating red tree coral presence to habitat charac- Alaska. In Proceedings of the first international symposium teristics. ICES J. Mar. Sci. 72:2674–2683. https://doi. on deep-sea corals; Halifax, Nova Scotia, 30 July–2 August org/10.1093/icesjms/fsv163 2000 (J. H. M. Willison, J. Hall, S. E. Gass, E. L. R. Kench- Matsumoto, A. K. ington, M. Butler, and P. Doherty, eds.), p. 106–116. Ecol. 2005. Recent observations on the distribution of deep-sea Action Cent. and Nova Scotia Mus., Halifax, NS, Canada. coral communities on the Shiribeshi Seamount, Sea of Ja- Krieger, K. J., and B. L. Wing. pan. In Cold-water corals and ecosystems (A. Freiwald and 2002. Megafauna associations with deepwater corals (Prim- J. M. Roberts, eds.), p. 345–356, Springer-Verlag, Berlin, noa spp.) in the Gulf of Alaska. Hydrobiologia 471:83–90. Germany. https://doi.org/10.1023/A:1016597119297 2007. Effects of low water temperature on growth and mag- Kükenthal, W. nesium carbonate concentrations in the cold-water gorgo- 1906. Diagnosen neuer japanischer Alcyonaceen. Zool. Anz. nian Primnoa pacifica. Bull. Mar. Sci. 81:423–435. 30(8/9):280–289. Matsumoto, A. K., L. P. van Ofwegen, and F. M. Bayer. 1908. Diagnosen neuer Gorgoniden (4 Mittelung.). Zool. Anz. 2019. A revision of the genus Calcigorgia (Cnidaria, Octo- 33(1):9–20. corallia, Acanthogorgiidae) with the description of three 1915a. System und Stammesgeschichte der Isididae. Zool. new species. Zootaxa 4571:1–27. https://doi.org/10.11646/ Anz. 46:116–126. [Although the publication date of this zootaxa.4571.1.1 work is 1916, the article was published in 1915.] McFadden, C. S., and F. G. Hochberg. 1915b. Pennatularia. Das Tierreich 43:1–132.1919. Wissen- 2003. Biology and taxonomy of encrusting alcyoniid soft schaftliche Ergebnisse der Deutschen Tiefsee-Expedition corals in the northeastern Pacific Ocean with descriptions auf dem Dampfer “Valdivia” 1898–1899. Vol. 13, part 2, of two new genera (Cnidaria, Anthozoa, Octocorallia). Gorgonaria, first half, 646 p. Invertebr. Biol. 122:93–113. https://doi.org/10.1111/j. Lamb, A., and B. P. Hanby. 2005. Marine life of the Pacific Northwest: a photographic 1744-7410.2003.tb00076.x encyclopedia of invertebrates, seaweeds and selected fishes, McFadden, C. S., L. P. van Ofwegen, and A. M. Quattrini. 398 p. Harbour Publ., Madeira Park, BC, Canada. 2022. Revisionary systematics of Octocorallia (Cnidaria: Langton, R. W., E. W. Langton, R. B. Theroux, and J. R. Uzmann. Anthozoa) guided by phylogenomics. Bull. Soc. Syst. Biol. 1990. Distribution, behavior and abundance of sea pens, Pen- 1(3):8735. https://doi.org/10.18061/bssb.v1i3 natula aculeata, in the Gulf of Maine. Mar. Biol. 107:463– Miller, R. J., J. Hocevar, R. P. Stone, and D. V. Fedorov. 469. https://doi.org/10.1007/BF01313430 2012. Structure-forming corals and sponges and their use Lapid, E. D., and N. E. Chadwick. as fish habitat in Bering Sea submarine canyons. PLoS 2006. Long-term effects of competition on coral growth ONE 7(3):e33885. https://doi.org/10.1371/journal. and sweeper tentacle development. Mar. Ecol. Prog. Ser. pone.0033885 313:115–123. https://doi.org/10.3354/meps313115 Molodtsova, T. N. Lapointe, A., and L. Watling. 2013. Deep-sea mushroom corals (Octocorallia: Alcyonacea: 2015. Bamboo corals from the abyssal Pacific: Bathygor- Alcyoniidae) of the Northern Mid-Atlantic Ridge. Mar. gia. Proc. Biol. Soc. Wash. 128:125–136. https://doi. Biol. Res. 9:488–515. https://doi.org/10.1080/17451000. org/10.2988/0006-324X-128.2.125 2012.750427 Linnaeus, C. Molodtsova, T. N., and D. M. Opresko. 1758. Systema naturae per regna tria naturae, secundum 2017. Black corals (Anthozoa: Antipatharia) of the Clarion- classes, ordines, genera, species, cum characteribus, differ- Clipperton fracture zone. Mar. Biodivers. 47:349–365. entiis, synonymis, locis (10th revised ed.), vol. 1, 824 p. https://doi.org/10.1007/s12526-017-0659-6 Holmiae, Laurtenii Salvii. Molodtsova, T. N., and F. A. Pasternak. Lumsden, S. E., T. F. Hourigan, A. W. Bruckner, and G. Dorr 2005. Redescription of Parantipathes euantha (Pasternak, (eds.). 1958) (Anthozoa: Antipatharia) from the Kurile-Kam- 2007. The state of deep coral ecosystems of the United States. chatka Trench. Invertebr. Zool. 2:169–179. https://doi. NOAA Tech. Memo. CRCP-3, 365 p. org/10.15298/invertzool.02.2.02 Madsen, F. J. Moroff, T. 1944. Octocorallia (Stolonifera – Telestacea – Xeniidae – Al- 1902. Studien über Octocorallien. I. Ueber die Pennatulaceen cyonacea – Gorgonacea). In The Danish Ingolf-Expedition, des Münchener Museums. Zoologische Jahrbücher (Syst.) vol. 5, part 13, 65 p. H. Hagerup, Copenhagen, Denmark. 17:363–403, plates 14–15. Literature Cited 397 Morrison, C. L., M. J. Springmann, K. M. Shroades, and R. P. and black coral morphology. N. Z. Aquat. Environ. Bio- Stone. divers. Rep. 131, 20 p. [Available from https://www.mpi. 2015. Development of twelve microsatellite loci in the red tree govt.nz/dmsdocument/4377-aebr-131-antipatharia-black- corals Primnoa resedaeformis and Primnoa pacifica. Con- corals-for-the-new-zealand-region.] serv. Genet. Resour. 7:763–765. https://doi.org/10.1007/ Pante, E., and S. C. France. s12686-015-0455-1 2010. Pseudochrysogorgia bellona n. gen., n. sp.: a new ge- Mortensen, P. B., and L. Buhl-Mortensen. nus and species of chrysogorgiid octocoral (Coelenterata, 2005. Morphology and growth of the deep-water gorgonians Anthozoa) from the Coral Sea. Zoosystema 32:595–612. Primnoa resedaeformis and Paragorgia arborea. Mar. Biol. https://doi.org/10.5252/z2010n4a4 147:775–788. https://doi.org/10.1007/s00227-005-1604-y Pante, E., S. C. France, A. Couloux, C. Cruaud, C. S. McFadden, Moseley, H. N. S. Samadi, and L. Watling. 1881. Report on certain hydroid, Alcyonarian, and Madrepo- 2012. Deep-sea origin and in-situ diversification of chrys- rarian corals procured during the voyage of H.M.S. Chal- ogorgiid octocorals. PLoS ONE 7(6):e38357. https://doi. lenger, in the years 1873–1876. In Report on the scientific org/10.1371/journal.pone.0038357 results of the voyage of H.M.S. Challenger during the years Pasternak, F. A. 1873–76, Zoology, vol. 2 (prepared by C. W. Thomson), 1958. Die Tiefsee Antipatharien des Kurilen-Kamchatka Gra- 248 p. Neill and Co., Edinburgh, Scotland. [While the pub- bens. Trud. Inst. Oceanol. 27:180–191. lication date of this volume is dated 1881, the section ref- 1977. Antipatharia. In Galathea report. Scientific results of erenced here is dated 1880.] the Danish deep-sea expedition around the world, 1950– Normark, W. R., and P. R. Carlson. 52, vol. 14 (T. Wolff, ed.), p. 157–164. Scand. Sci. Press, 2003. Giant submarine canyons: is size any clue to their im- Copenhagen, Denmark. portance in the rock record? In Extreme depositional envi- Penn, K., D. Wu, J. A. Eisen, and N. Ward. ronments: mega end members in geologic time. Geol. Soc. 2006. Characterization of bacterial communities associ- Am. Spec. Pap. 370 (M. A. Chan and A. W. Archer, eds.), p. ated with deep-sea corals on Gulf of Alaska seamounts. 175–190. Geol. Soc. Am., Boulder, CO. Appl. Environ. Microbiol. 72:1680–1683. https://doi. Nørrevang, A. org/10.1128/AEM.72.2.1680-1683.2006 1973. On a peculiar mode of germ-layer formation: the Pérez, C. D., R. T. S. Cordeiro, G. C. Williams, and P. B. Gomes. early embryology of Gersemia rubiformis (Ehrenberg 2021. Revised nomenclature of the sea pen genus Bal- 1834) (Octocorallia). Acta Zool. 54:65–71. https://doi. ticina Gray, 1870 (= Halipteris Kölliker, 1870) (Antho- org/10.1111/j.1463-6395.1973.tb00443.x zoa: Octocorallia). Zootaxa 4966:237–244. https://doi. NRC (National Research Council). org/10.11646/zootaxa.4966.2.10 1990. Assessment of the U.S. outer continental shelf en- Pires, D. O., C. B. Castro, and J. C. Silva. vironment studies program: 1. physical oceanography, 2009. Reproductive biology of the deep-sea pennatulacean 144 p. Natl. Acad. Press, Washington, DC. https://doi. Anthoptilum murrayi (Cnidaria, Octocorallia). Mar. org/10.17226/1609 Ecol. Prog. Ser. 397:103–112. https://doi.org/10.3354/ Nutting, C. C. meps08322 1908. Descriptions of the Alcyonaria collected by the U.S. Bu- Reece, R. S., S. P. S. Gulick, B. K. Horton, G. L. Christeson, and reau of Fisheries steamer Albatross in the vicinity of the L. L. Worthington. Hawaiian Islands in 1902. Proc. U. S. Natl. Mus. 34:543– 2011. Tectonic and climatic influence on the evolution of the 601. Surveyor Fan and Channel system, Gulf of Alaska. Geo- 1909. Alcyonaria of the California coast. Proc. U. S. Natl. Mus. sphere 7:830–844. https://doi.org/10.1130/GES00654.1 35(1658):681–727. https://doi.org/10.5479/si.00963801. Reimer, J. D., and F. Sinniger. 35-1658.681 2010. Unexpected diversity in Canadian Pacific zoanthids 1912. Descriptions of the Alcyonaria collected by the U. S. (Cnidaria: Anthozoa: Hexacorallia): a molecular examina- Fisheries steamer “Albatross,” mainly in Japanese wa- tion and description of a new species from the waters of ters, during 1906. Proc. U.S. Natl. Mus. 43(1923):1–104. British Columbia. Mar. Biodivers. 40:249–260. https://doi. https://doi.org/10.5479/si.00963801.43-1923.1 org/10.1007/s12526-010-0045-0 Opresko, D. M. Roark, E. B., T. P. Guilderson, S. Flood-Page, R. B. Dunbar, B. L. 2003. Revision of the Antipatharia (Cnidaria: Anthozoa). Part Ingram, S. J. Fallon, and M. McCulloch. III. Cladopathidae. Zool. Meded. 77:495–536. 2005. Radiocarbon-based ages and growth rates of bam- 2005. New genera and species of antipatharian corals (Cni- boo corals from the Gulf of Alaska. Geophys. Res. Lett. daria: Anthozoa) from the North Pacific. Zool. Meded. 79- 32:L04606. https://doi.org/10.1029/2004GL021919 2:129–165. Roark, E. B., T. P. Guilderson, R. B. Dunbar, and B. L. Ingram. Opresko, D. M., and T. N. Molodtsova. 2006. Radiocarbon-based ages and growth rates of Hawaiian 2021. New species of deep-sea antipatharians from the deep-sea corals. Mar. Ecol. Prog. Ser. 327:1–14. https://doi. North Pacific (Cnidaria: Anthozoa: Antipatharia), Part 2. org/10.3354/meps327001 Zootaxa 4999:401–422. https://doi.org/10.11646/zoo- Roberts, J. M., A. Wheeler, A. Freiwald, and S. Cairns. taxa.4999.5.1 2009. Cold-water corals: the biology and geology of deep-sea Opresko, D., D. Tracey, and E. Mackay. coral habitats, 334 p. Cambridge Univ. Press, Cambridge, 2014. Antipatharia (black corals) for the New Zealand region. UK. A field guide of commonly sampled New Zealand black Rossin, A. M., R. G. Waller, and R. P. Stone. corals including illustrations highlighting technical terms 2019. The effects of in-vitro pH decrease on the gameto- 398 Professional Paper NMFS 23 genesis of the red tree coral, Primnoa pacifica. PLoS Slephkova, N. A., and L. N. Seravin. ONE 14(4):e0203976. https://doi.org/10.1371/journal. 1983. Spontaneous and induced stretching of polyps in the pone.0203976 coral Gersemia fruticosa (Alcyonacea, Nephthyidae). Zool. Ryland, J. S., and J. E. Lancaster. Zh. 62:675–680. [In Russian.] 2003. Revision of methods for separating species of Protopa- Squires, D. F. lythoa (Hexacorallia: Zoanthidea) in the tropical West Pa- 1962. The fauna of the Ross Sea. Part 2. Scleractinian corals. cific. Invertebr. Syst. 17:407–428. https://doi.org/10.1071/ N. Z. Oceanogr. Inst. Mem. 19, 28 p. IS02008 Stearns, R. E. C. Saenger, C., R. I. Gabitov, J. Farmer, J. M. Watkins, and R. Stone. 1883. Descriptions of a new genus and species of alcyonoid 2017. Linear correlations in bamboo coral δ13C and δ18O polyp, from Japanese waters, with remarks on the struc- sampled by SIMS and micromill: evaluating paleoceano- ture and habits of related forms, etc. Proc. U.S. Nat. Mus. graphic potential and biomineralization mechanisms using 6:96–101. δ11B and ∆47 composition. Chem. Geol. 454:1–14. https:// Stone, R. P. doi.org/10.1016/j.chemgeo.2017.02.014 2006. Coral habitat in the Aleutian Islands of Alaska: depth Sanamyan, K. E., and N. P. Sanamyan (eds.). distribution, fine-scale species associations, and fisher- 2020. Flora and fauna of Matua Island (middle Kuril Island): ies interactions. Coral Reefs 25:229–238. https://doi. the field guide. Volume 1: Sea, 495 p. OOO Intron, Chere- org/10.1007/s00338-006-0091-z povets, Russia. [Available from http://sanamyan.com/ma- 2014. The ecology of deep-sea coral and sponge habitats of tua/eng.php, in Russian.] the central Aleutian Islands of Alaska. NOAA Prof. Pap. Sánchez, J. A. NMFS 16, 52 p. 2005. Systematics of the bubblegum corals (Cnidaria: Oc- Stone, R. P., and S. D. Cairns. tocorallia: Paragorgiidae) with description of new spe- 2017. Deep-sea coral taxa in the Alaska region: depth and cies from New Zealand and the Eastern Pacific. Zootaxa geographic distribution, 9 p. Natl. Mar. Fish. Serv., Silver 1014:1–72. https://doi.org/10.11646/zootaxa.1014.1.1 Spring, MD. [Available from https://deepseacoraldata. Sánchez, J. A., and S. D. Cairns. noaa.gov/library/2017-state-of-deep-sea-corals-report.] 2004. An unusual new gorgonian coral (Anthozoa: Octo- Stone, R. P., and J. Mondragon. corallia) from the Aleutian Islands, Alaska. Zool. Meded. 2018. Deep-sea emergence of red tree corals (Primnoa paci- 78:265–274. fica) in Southeast Alaska glacial fjords. NOAA Prof. Pap. Saucier, E. H., S. C. France, and L. Watling. NMFS 20, 33 p. 2021. Toward a revision of the bamboo corals: part 3, de- Stone, R. P., and C. N. Rooper. constructing the family Isididae. Zootaxa 5047:247–272. 2017. State of the deep-sea coral and sponge ecosystems of the https://doi.org/10.11646/zootaxa.5047.3.2 Alaska Region. In The state of deep-sea coral and sponge Sebens, K. P., and J. S. Miles. ecosystems of the United States (T. F. Hourigan, P. J. Etnoy- 1988. Sweeper tentacles in a gorgonian octocoral: morpholog- er, and S. D. Cairns, eds.), p. 57–92. NOAA Tech. Memo. ical modifications for interference competition. Biol. Bull. NMFS-OHC-4. 175:378–387. https://doi.org/10.2307/1541729 Stone, R. P., and S. K. Shotwell. Seravin, L. N., and A. V. Gudkov. 2007. State of deep coral ecosystems in the Alaska Region: 1990. Spontaneous and periodic changes in the form and size Gulf of Alaska, Bering Sea and the Aleutian Islands. In of the alcyonarian colony of Gersemia fruticosa (Alcyona- The state of deep coral ecosystems of the United States (S. cea, Nephthyidae). Translated by the Laboratory of Inver- E. Lumsden, T. F. Hourigan, A. W. Bruckner, and G. Dorr, tebrate Zoology, Biological Research Institute, Leningrad eds.), p. 65–108. NOAA Tech. Memo. CRCP-3. University, Leningrad, Staryi Petergof. Biol. Morya 3:9–12. Stone, R. P., and B. L. Wing. Sherwood, O. A., and E. N. Edinger. 2001. Growth and recruitment of an Alaskan shallow-water 2009. Ages and growth rates of some deep-sea gorgonian and gorgonian. In Proceedings of the first international sympo- antipatharian corals of Newfoundland and Labrador. Can. sium on deep-sea corals; Halifax, Nova Scotia, 30 July–2 J. Fish. Aquat. Sci. 66:142–152. https://doi.org/10.1139/ August 2000 (J. H. M. Willison, J. Hall, S. E. Gass, E. L. R. F08-195 Kenchington, M. Butler, and P. Doherty, eds.), p. 88–94. Sinniger, F., and V. Häussermann. Ecol. Action Cent. and Nova Scotia Mus., Halifax, NS, 2009. Zoanthids (Cnidaria: Hexacorallia: Zoantharia) from Canada. shallow waters of the southern Chilean fjord region, Stone, R. P., M. M. Masuda, and P. W. Malecha. with descriptions of a new genus and two new species. 2005. Effects of bottom trawling on soft-sediment epibenthic Org. Divers. Evol. 9:23–36. https://doi.org/10.1016/j. communities in the Gulf of Alaska. Am. Fish. Soc. Symp. ode.2008.10.003 41:439–453. https://doi.org/10.47886/9781888569605. Sinniger, F., J. D. Reimer, and J. Pawlowski. ch70 2010. The Parazoanthidae (Hexacorallia: Zoantharia) DNA Stone, R. P., H. Lehnert, and H. Reiswig. taxonomy: description of two new genera. Mar. Biodivers. 2011. A guide to the deep-water sponges of the Aleutian Is- 40:57–70. https://doi.org/10.1007/s12526-009-0034-3 land Archipelago. NOAA Prof. Pap. NMFS 12, 187 p. Sinniger, F., O. V. Ocaña, and A. R. Baco. Stone, R. P., K. W. Conway, D. J. Csepp, and J. V. Barrie. 2013. Diversity of zoanthids (Anthozoa: Hexacorallia) on Ha- 2013. The boundary reefs: glass sponge (Porifera: Hexacti- waiian seamounts: description of the Hawaiian gold coral nellidae) reefs on the international border between Can- and additional zoanthids. PLoS ONE 8(1):e52607. https:// ada and the United States. NOAA Tech. Memo. NMFS- doi.org/10.1371/journal.pone.0052607 AFSC-264, 31 p. Literature Cited 399 Stone, R. P., D. Stevenson, and S. Brooke. Vallier, T. L., D. W. Scholl, M. A. Fisher, T. R. Bruns, F. H. Wilson, 2015a. Assessment of a pilot study to collect coral bycatch R. von Huene, and A. J. Stevenson. data from the Alaska commercial fishing fleet. NOAA Tech. 1994. Geologic framework of the Aleutian arc, Alaska. In Memo. NMFS-AFSC-296, 45 p. The geology of Alaska (G. Plafker and H. C. Berg, eds.), p. Stone, R. P., M. M. Masuda, and J. F. Karinen. 367–388. Geol. Soc. Am., Boulder, CO. 2015b. Assessing the ecological importance of red tree coral Vaughan, T. W. thickets in the eastern Gulf of Alaska. ICES. J. Mar. Sci. 1900. A new fossil species of Caryophyllia from California, 72:900–915. https://doi.org/10.1093/icesjms/fsu190 and a new genus and species of turbinolid coral from Japan. Stone, R. P., P. W. Malecha, and M. M. Masuda. Proc. U.S. Natl. Mus. 22:199–203. https://doi.org/10.5479/ 2017. A five-year, in situ growth study on shallow-water pop- si.00963801.22-1194.199 ulations of the gorgonian octocoral Calcigorgia spiculifera 1906. Reports on the scientific results of the expedition to the in the Gulf of Alaska. PLoS ONE 12(1):e0169470. https:// Eastern Tropical Pacific, in charge of Alexander Agassiz, doi.org/10.1371/journal.pone.0169470 by the U. S. Fish Commission Steamer “Albatross” from Studer, T. October, 1904, to March, 1905, Lieut. Commander L. M. 1878. Ubersicht der Anthozoa Alcyonaria welche wahrend Garrett, U. S. N. commanding. Part 6: Madreporaria. Bull. der Reise S.M.S. Gazelle um die Erde gesammelt wurden. Mus. Comp. Zool. Harv. Univ. 50:59–72. Mon. K. Preuss. Akad. Wiss. Berlin. 1878:632–688. 1941. New corals: one Recent, Alaska; three Eocene, Alabama 1894. Note préliminaire sur les Alcyonaires. Reports on the and Louisiana. J. Paleontol. 15:280–284. dredging operations off the West Coast of Central America Verrill, A. E. to the Galapagos, to the West Coast of Mexico, and in the 1864. List of the polyps and corals sent by the Museum of Gulf of California, in charge of Alexander Agassiz, carried Comparative Zoology to other institutions in exchange, on by the U.S. Fish Commission steamer “Albatross,” dur- with annotations. Bull. Mus. Comp. Zool. Harv. Univ. ing 1891. Bull. Mus. Comp. Zool. 25:53–69. 1(3):29–60. Taylor, M. L., S. D. Cairns, D. J. Agnew, and A. D. Rogers. 1866. Synopsis of the polyps and corals of the North Pacific 2013. A revision of the genus Thouarella Gray, 1870 (Octo- exploring expedition, under Commodore C. Ringgold and corallia: Primnoidae), including an illustrated dichotomous Captain John Rodgers, U. S. N., from 1853 to 1856. Col- key, a new species description, and comments on Plumarel- lected by Dr. Wm. Stimpson, naturalist to the expedition. la Gray, 1870 and Dasystenella, Versluys, 1906. Zootaxa With descriptions of some additional species from the west 602:1–105. https://doi.org/10.11646/zootaxa.3602.1.1 coast of North America. Part 2: Alcyonaria. With two Tendal, O. S. plates. Proc. Essex Inst. 4:181–196. [While the publica- 1992. The North Atlantic distribution of the octocoral Para- tion date of this work is 1866, this chapter was written gorgia arborea (L., 1758) (Cnidaria, Anthozoa). Sarsia in 1865.] 77:213–217. https://doi.org/10.1080/00364827.1992.104 13506 1871. Notes on the Radiata in the Museum of Yale College, Thoma, J. N. with description on new genera and species. No. 6. Review 2013. Molecular and morphological diversity of sea fans with of the corals and polyps of the West Coast of America. emphasis on deep-sea octocorals of the order Alcyonacea Trans. Conn. Acad. Arts Sci. 1(2):377–558. [The descrip- Lamouroux, 1812. Ph.D. diss., 170 p. Univ. Louisiana, tion was presented in 1868 but the publication that it ap- Lafayette, LA. [Available from https://ull.ent.sirsi.net/cli- pears in was published in 1871.] ent/en_US/ull/search/detailnonmodal/ent:$002f$002fSD_ 1879. Notice of recent additions to the marine fauna of the ULL$002f0$002fSD_ULL:2196153/one.] eastern coast of North America, No. 3. Am. J. Sci. Arts, Ser. Thomson, J. A. 3, 17:239–243. 1927. Alcyonaires provenant des campagnes scientifiques du 1922. Report of the Arctic expedition 1913–18. Volume 8. Prince Albert Ier de Monaco. Résultats des Campagnes Sci- Mollusks, echinoderms, coelenterates, etc. Part G. Alcyo- entifiques Accomplies sur son yacht par Albert Ier. 73:1– naria and Actinaria, 165 p. F. A. Acland, Ottawa, ON, 77, 6 plates. Canada. Tracey, D., H. Neil, D. Gordon, and S. O’Shea. Verseveldt, J., and L. P. van Ofwegen. 2003. Chronicles of the deep: ageing deep-sea corals in New 1992. New and redescribed species of Alcyonium Linnaeus, Zealand waters. Water Atmos. 11(2):22–24. 1758 (Anthozoa: Alcyonacea). Zool. Meded. 66(7):155– Turner, D. L., R. B. Forbes, and C. W. Naeser. 181. 1973. Radiometric ages of Kodiak seamount and Giacomini Versluys, J. guyot, Gulf of Alaska: implications for circum-Pacific tec- 1902. Die Gorgoniden der Siboga-Expedition. I. Die Chrys- tonics. Science 182:579–581. https://doi.org/10.1126/sci- ogorgiiden, Siboga Exped. Monogr. 13, 120 p. ence.182.4112.579 Von Huene, R. Tuskes, P. M., and R. Clark. 1989. Continental margins around the Gulf of Alaska. In 2018. A new species of Otukaia (Calliostomatidae) from Alas- Eastern Pacific Ocean and Hawaii (E. L. Winterer, D. M. ka. Festivus 50:236–238. Hussong, and R. W. Decker, eds.), p. 383–401. Geol. Soc. Utinomi, H. Am., Boulder, CO. 1976. A review of the Japanese species of Alcyonium, with Waller, R. G., R. P. Stone, J. Johnstone, and J. Mondragon. descriptions of two new species and an almost forgotten 2014. Sexual reproduction and seasonality of the Alaskan rare species (Octocorallia, Alcyonacea). Publ. Seto Mar. red tree coral, Primnoa pacifica. PLoS ONE 9(4):e90893. Biol. Lab. 23(3/5):191–204. https://doi.org/10.1371/journal.pone.0090893 400 Professional Paper NMFS 23 Waller, R. G., R. P. Stone, L. N. Rice, J. Johnstone, A. M. Rossin, 1878, with the description of a new species from the north- E. Hartill, K. Feehan, and C. L. Morrison. east Pacific and a review of two additional species (Octo- 2019. Phenotypic plasticity or a reproductive dead end? corallia: Alcyonacea). Zool. Meded. 83:1067–1081. Primnoa pacifica (Cnidaria: Alcyonacea) in the south- Williams, G. C., B. W. Hoeksema, and L. P. van Ofwegen. eastern Alaska region. Front. Mar. Sci. 6:709. https://doi. 2012. A fifth morphological polyp in pennatulacean octo- org/10.3389/fmars.2019.00709 corals, with a review of polyp polymorphism in the genera Watling, L., S. C. France, E. Pante, and A. Simpson. Pennatula and Pteroeides (Anthozoa: Pennatulidae). Zool. 2011. Biology of deep-water octocorals. Adv. Mar. Biol. Stud. 51:1006–1017. 60:41–122. https://doi.org/10.1016/B978-0-12-385529- Wilson, M. T., A. H. Andrew, A. L. Brown, and E. E. Cordes. 9.00002-0 2002. Axial rod growth and age estimation of the sea pen, Williams, B., M. Risk, R. Stone, D. Sinclair, and B. Ghaleb. Halipteris willemoesi Kölliker. Hydrobiologia 471:133– 2007. Oceanographic changes in the North Pacific Ocean over 142. the past century recorded in deep-water gorgonian corals. Wing, B. L., and D. B. Barnard. Mar. Ecol. Prog. Ser. 335:85–94. https://doi.org/10.3354/ 2004. A field guide to Alaskan corals. NOAA Tech. Memo. meps335085 NMFS-AFSC-146, 67 p. Williams, G. C. Wood, R. L. 1986. Morphology, systematics, and variability of the south- 1957. Identification and microanatomical study of a new spe- ern African soft coral Alcyonium variabile (J. Stuart Thom- cies of Epizoanthus (Zoanthidea). Diss. Abstr. 18(2):707– son, 1921) (Octocorallia, Alcyoniidae). Ann. South Afr. 708. Mus. 96:241–270. 1995. Living genera of sea pens (Coelenterata: Octocorallia: WoRMS Editorial Board. Pennatulacea): illustrated key and synopses. Zool. J. Linn. 2022. World Register of Marine Species. https://doi. Soc. 113:93–140. https://doi.org/10.1006/zjls.1995.0004 org/10.14284/170 2005. New taxa of octocorals (Anthozoa: Octocorallia) from Wright, E. P. the northeastern Pacific Ocean. Proc. Calif. Acad. Sci. 1885. The Alcyonaria. In Report of the scientific results of the 56:53–65. voyage of H.M.S. Challenger. Narrative – volume 1, part 2013. New taxa and revisionary systematics of alcyona- 2 (C. W. Thomson and J. Murray, eds.), p. 689–693. Neill cean octocorals from the Pacific coast of North America and Co., Edinburgh, Scotland. (Cnidaria, Anthozoa). ZooKeys 283:15–42. https://doi. Wright, E. P., and T. Studer. org/10.3897/zookeys.283.4803 1889. Report on the Alcyonaria collected by H.M.S. Chal- Williams, G. C., and P. Alderslade. lenger during the years 1873–1876. In Report of the sci- 2011. Three new species of pennatulacean octocorals with entific results of the voyage of H.M.S. Challenger during the ability to attach to rocky substrata (Cnidaria: An- the years 1873–76. Zoology, vol. 31, 314 p. Neill and Co., thozoa: Pennatulacea). Zootaxa 3001:33–48. https://doi. Edinburgh, Scotland. org/10.11646/zootaxa.3001.1.2 Yamada, M. Williams, G. C., and L. Lundsten. 1950. Descriptions of two Alcyonium from northern Japan. 2009. The nephtheid soft coral genus Gersemia Marenzeller, Annot. Zool. Jpn. 23:114–116. Appendix I 401 Appendix I. Glossary Order Antipatharia (Adapted from Opresko et al., Lateral tentacles: the pairs of tentacles at the distal and 2014) proximal extremities of a polyp Abpolypar: on the side of the skeletal axis opposite to Mesentery (septa): six, 10, or 12 partitions of tissue the side bearing the polyps which extend across from the inside wall of the polyp to the actinopharynx in a radial pattern (when viewed in Actinopharynx: cylindrical tube of tissue leading down cross sectional view) from the mouth and supported internally by radial mes- enteries (also referred to as septa in earlier literature) Monopodial: having a stem extending to the top of the corallum Alternating (arrangement of pinnules): pinnules in lat- eral rows offset from each other Mutual distance (of spines): the distance between the midpoints of the bases of adjacent spines or branches Anisomorphic: unequal development of the spines in the immediate area of the polyps; the hypostomal spines are Oral cone: an elevated area of tissue around the mouth reduced in size; the circumpolypar spines are enlarged; of a polyp and the interpolypar spines are of an intermediate size Peristome: the area on the surface of a polyp between Basal: in a direction towards the attachment point of a the mouth and tentacles pinnule, branchlet, branch, or stem Pinnulate: having symmetrically arranged, simple or Basal diameter: diameter of the axis just above the at- branched ramifications of subequal size on the stem and tachment plate or holdfast branches of the corallum Branchlets: smallest ramifications of a corallum, usually Pinnules: symmetrically arranged, simple or branched of varying size and orientation ramifications of nearly equal length located on the stem and branches of the corallum Circumpolypar: area around the outer edge of the pol- yps, beneath the tentacles Polypar spines: spines on the side of the axis that bear the polyps; not restricted to the immediate area of each Complete mesenteries (septa): radially arranged endo- individual polyp dermal partitions joining the actinopharynx to the inte- rior body wall of the polyp Primary mesenteries: six internal partitions of tissue each extending across from the actinopharynx radially Corallum: the skeletal structure of the colony to the interior wall of the polyp and down to the ab- oral margin of the actinopharynx, thereby defining the Distal: direction towards the tip of a pinnule, branchlet, boundaries of the six tentacles, and appearing in cross branch, or stem section as spokes of a wheel Distal angle: the angle formed by the upper side of the Primary pinnules: small branchlets of a colony that spines or branches, and the axis are usually of equal size and arranged in a symmetric pattern around the axis (they may be simple or bear Distal lateral tentacles: the pair of tentacles at the distal subpinnules) end of a polyp Proximal lateral tentacles: the pair of tentacles on the Hypostomal: the area on the skeletal axis directly below side of the polyp closest to the base of the branch/pin- the mouth of the polyp nule bearing the polyp Interpolypar: the area between the distal lateral tenta- Sagittal axis: a plane through the center of the actino- cles of one polyp and the proximal lateral tentacles of pharynx, perpendicular to the direction of the stem or the adjoining one branch bearing the polyp 402 Professional Paper NMFS 23 Sagittal tentacles: the median pair of tentacles whose Verticillate (arrangement of pinnules): pinnules grouped members are placed on either side of the mouth and in in whorls around the circumference of the axis a plane, which is perpendicular to the direction of the branch bearing the polyp Order Scleractinia (Adapted from Cairns and Sclerenchyme: skeletal axis composed of concentric lay- Kitahara, 2012) ers of protein and chitin Axial septal edge: the inner edge of a septum, adjacent Secondary mesenteries: complete mesenteries which to the fossa and/or columella (see Fig. 3-1) are not attached all the way to the aboral edge of the actinopharynx Basal plate: the thin encrusting calcareous plate secreted by the planula when first attaching to a substrate Secondary pinnules: very small ramifications that arise from the primary pinnules; often arranged uniserially or Calice: the skeletal analog of the polyp, cupping the bilaterally and sometimes giving rise to tertiary pinnules polyp from below, and consisting of the septa, and, if present, the columella and pali (see Fig. 3-1) Semispiral (arrangement of pinnules): groups of pin- nules (one from each of the longitudinal rows) on ei- Ceratoid: the triangular shape of a solitary corallum cre- ther side of a branch or stem which are offset slightly ated if the sides of the corallum diverge at an angle of to those from which they form a curved ascending or 10°–40° descending pattern around the circumference of the axis Columella: an axial structure of diverse shape and com- Septa: see mesentery position that projects from the center of a calice; if com- posed of twisted lamellae, it is termed fascicular (see Fig. Simple (pinnule): without further sub-ramifications 3-1) Siphonoglyph: a ciliary tract extending in an oral-aboral Corallum (coralla): the aragonitic calcium carbonate direction on the sagittal border of the actinopharynx skeleton of a scleractinian coral Stalk: the lowermost unpinnulated part of the stem in Costa (costae): continuation of a septum on the out- monopodial pinnulated colonies, especially those of side of the corallite wall, often as a ridge or low linear Bathypathes species mound (see Fig. 3-1) Stem: the primary axis of a colony that arises from the Cycle: see septum holdfast and from which branches or pinnules may or may not arise Epitheca: thin, external smooth or wrinkled, non-tra- becular sheath (wall) surrounding individual corallites Subopposite (arrangement of pinnules): pinnules in lat- eral rows grouped together in pairs whose members are Exsert septum: a septum whose distal edge projects almost directly opposite one another above the thecal rim (Fig. 3-1) Subpinnules: general term for one or more orders of Fascicular: see columella small ramifications which can arise from the primary pinnules, and consisting of secondary, tertiary, and qua- Fossa: the central region of a corallite, in which the pol- ternary pinnules yp sits (see Fig. 3-2) Tertiary pinnules: very small ramifications that arise Half-system: one twelfth of the septa of a calice, consist- from the secondary pinnules; usually arranged uniseri- ing of the septa between an S1 and an adjacent S2 (see ally, and sometimes giving rise to quaternary pinnules Fig. 3-2) Transverse axis: a plane through the center of the ac- Palus (pali): small flattened lobe on the axial edge of tinopharynx in the direction of the branch bearing the various septal cycles, always one per septum polyp Appendix I 403 Pedicel: the stem-like region of a solitary coral just above Order Zoantharia (Adapted from Ryland and the base and below the calicular surface (see Fig. 3-1) Lancaster, 2003) Polycyclic base: the lower several millimeters of a soli- Capitulum: top part of the contracted polyp formed by tary corallum that is composed of concentric rings of the contracted tentacles partitioned chambers Cnidae: explosive cells containing one giant secretory Polyp: the fleshy non-skeletal part of the coral, consist- organelle that can deliver a sting to other organisms; zo- ing of tentacles, mouth, and mesenteries antharians possess five main types of cnidae with many variations: spirocysts, spirulae, penicilli, basitrichs, and Porcelaneous: smooth, polished homotrichs (equivalent to nematocysts) Pourtalès plan: a form of septal arrangement in which Cnidome: the specific complement of cnidae for a given the axial edges of pairs of higher cycle septa bend in species front of and unite before their adjacent lower cycle sep- tum (see Fig. 3-2) Gastrodermal canals: canals that connect the individual polyps through the coenenchyme (also referred to as me- Protosepta: the first septa deposited by a planula after sogloeal canals) settlement, usually 6 or 12 in number Macrocnemes: fully developed mesenteries Septal formula: a shorthand notation to indicate the relative widths of the respective septal cycles, thus S1– Mesogloea: the connective tissue layer between the epi- 2>S3>S4 would mean that the septa of the first two cy- dermis and gastrodermis cles are equal in width and wider than those of the third cycle septa, which in turn are wider than those of the Scapulus: the oral disc of each polyp in the colony fourth cycle septa Scapus: the column of the polyp from the base to the Septum (septa): radially arranged longitudinal partitions capitulum of a corallite, usually arranged in hexameral symmetry; septa are added in cycles, with the first cycle composed Siphonoglyph: the strongly ciliated grooves extending of 6 septa, the second of 6, the third of 12, the fourth down one side of the pharynx of 24, etc.; the first cycle septa are designated as S1, the second as S2, etc. (see Fig. 3-1) Subclass Octocorallia (Adapted from Bayer et al., Stereome: a general term for thick calcareous deposits 1983) on various parts of the corallum Abaxial: away from the axis Synapticulotheca: a porous wall of a solitary corallum, characteristic of the family Dendrophylliidae Acanthose: thorny System: one sixth of the septa of a corallum, consisting Adaxial: towards the axis of the septa between two adjacent S1 (see Fig. 3-2) Anthocodia: the distal part of a polyp bearing the mouth Theca: the wall of a solitary corallum (see Fig. 3-1) and the tentacles Trabecular spine: linearly arranged spines that project Anthostele: the proximal, rigid part of some polyps, of- from the upper septal edges ten stiffened by sclerites, and into which the anthocodia may be withdrawn (equivalent to calyx) Trochoid: the triangular shape of a solitary corallum created if the sides of the corallum diverge at an angle Arborescent: tree-like colonies with a definite stalk of 40°–60° Autozooid: a polyp with eight well-developed ten- tacles and mesenteries; the larger polyp in dimorphic 404 Professional Paper NMFS 23 species that is typically responsible for food capture and Cross: a stellate sclerite with four rays in one plane reproduction Crown: the ring of transversely placed, usually bow- Axial sheath: the coenenchyme immediately surround- shaped sclerites encircling the anthocodia below the ten- ing the axis tacles (equivalent to collaret) Axis: the inner supporting structure that in octocorals Crown and points: the combined arrangement of the may be composed of aragonite, high-magnesium calcite, sclerites of the crown and the points low-magnesium calcite, horny material, or a combina- tion of these compounds Dichotomous branching: a branching pattern of repeat- ed bifurcation Bottlebrush: pinnately branched colonies in which nu- merous, crowded, short branchlets arise all around the Dimorphism: the presence of two types of polyps: auto- main stalk zooids and siphonozooids Buccal: pertaining to the mouth cavity Distad: toward or near the distal part or end Bushy: colonies with abundant branches arising imme- Double star: a rod sclerite with two whorls of irregu- diately above the holdfast and not forming an obvious larly shaped, more or less spiny projections main stem En chevron: V-shaped Calyx (calyces): cylindrical or wart-like projecting anthostele Flagelliform: stout, unbranched, whip-like colonies Capitate: unbranched colonies with a broad distal sec- Fusiform: tapered at each end; spindle-shaped tion on a distinctly narrower stalk Gastric cavity: the interior space of a polyp Capitulum: disc-shaped or hemispherical, polypiferous section of an alcyonacean colony (family Alcyoniidae) Holdfast: the part of the colony that attaches to the substrate Central chord: the central part of the axis in order Mala- calcyonacea, made up of horny material and to some Internode: the rigid calcareous segment of the Keratoisi- degree calcareous material didae axis that is white or creamy white in color Clavate: club-shaped or a terminally enlarged branch Introvert: (equivalent to neck zone) Clubs: monaxial sclerites enlarged at the head end and Irregular branching: branching that originates irregular- tapered at the handle end ly (neither pinnate nor dichotomous) in planar or nearly planar colonies Coenenchyme: the colonial tissue between polyps con- sisting of mesogloea usually containing sclerites and Loculus (loculi): a calcified or fiber-filled space in the penetrated by a network of gastrodermic canals axes of malacalcyonaceans (particularly those in the former family Plexauridae) that in cross section may ap- Collaret: (see crown) pear crescentic or lenticular Colony: a group of interconnected, genetically identical, Medulla: the inner supporting structure of some corals elementary functional units (i.e., the polyps) (equivalent to central chord of malacalcyonaceans and axis in scleralcyonaceans) Contractile polyp: a polyp that can diminish in size without introversion; the tentacles alone may be folded Mesenteries: the thin, radial, non-calcareous partitions inward over the mouth, but there is no neck zone or that join the pharynx to the body wall and divide the introvert that provides for withdrawal of the entire an- gastrovascular cavity of the polyp thocodia into the anthostele or coenenchyme Appendix I 405 Mesogloea: a jelly-like substance separating the two epi- Retractile polyp: a polyp in which the anthocodia can thelial layers and containing many cells invert completely into the anthostele or coenenchyme Mesozooid: a polyp intermediate between an autozooid Rhizoids: the rootlike processes extending from the base and a syphonozooid of some colonies Monopodial branching: branching that occurs when the Rods: straight or curved monaxial sclerites that are main axis of the colony maintains a single line of growth blunt at both ends but gives rise to a few or many lateral branches Scales: thin, flat, or nearly flat sclerites Morph: a subset of individuals of a particular variant of a species that exhibits a distinct morphological feature(s) Sclerite: multi-formed calcareous structures found in the mesogloea and elsewhere in the colony Neck zone: the soft, thin-walled basal part of an an- thocodia, with few sclerites and providing for the intro- Siphonoglyph: the strongly ciliated grooves extending version of the anthocodia into the anthostele (equivalent down one side of the pharynx to introvert) Siphonozooid: a polyp with a strongly developed si- Needles: long, thin, nearly smooth monaxial sclerites phonoglyph and reduced or no tentacles, usually much smaller than the autozooids, with the primary function Node: the flexible horny joint of the calcified axis in of irrigating colonies with seawater Keratoisididae that is typically black or brown in color Spindles: straight or curved monaxial sclerites that are Obconic: in the form of an inverted cone, attached at pointed at both ends the pointed end Stalk: the barren basal part of some colonies or the nar- Operculum: eight triangular scales covering the with- row proximal part of a non-retractile polyp drawn tentacles in species of the family Primnoidae Stem: the main part of the colony, from which branches Oral disc: the area of the polyp immediately surrounding may or may not arise, or the polyp-bearing part of the the mouth and formed by the inner parts of the tentacles colony, usually giving rise to the branches Pharynx: the tubular passageway between the mouth Stolon: a rounded or flattened elongate or membranous and the gastric cavity coenenchymal expansion growing over the substrate and peripherally producing new polyps Pinnate tentacles: tentacles bearing pinnules that are characteristic of all Octocorallia Stoloniferous: colonies consisting of multiple polyps in- terconnected by stolons Pinnules: the lateral processes of the tentacle Sulcal side: the side of a polyp nearest the siphonoglyph Plates: flat sclerites that are much thicker than scales, circular to oval, but often irregular in shape Sympodial branching: the apparent main axis of the col- ony formed by the proximal parts of a series of shorter Points: the eight rows of en chevroned (i.e., V-shaped) lateral branches (successive secondary axes) and having sclerites in the distal part of the anthocodia and super- a zigzag or spiral form posing the crown (if present) Teeth: the more or less pointed lobes of the calicular Polyp: an individual of an octocoral colony (equivalent margin, usually stiffened by sclerites, and typically num- to autozooid, siphonozooid, and mesozooid) bering between two and eight when present Radiates: sclerites with processes radiating in one plane Thornstar: a type of flat sclerite in which one or more (i.e., stellate forms) or in various planes in a more or less projecting thorns arise vertically from the middle of the symmetrical order divided, rootlike base and may be more-or-less leaflike; occurs in Isidella tentaculum 406 Professional Paper NMFS 23 Trimorphism: in Heteropolypus only, the presence of End bulb: terminal (basal) swelling of the stalk three types of polyps (autozooids, siphonozooids, and mesozooids) Longitudinal canals: the four main canals in some spe- cies formed by the gastric cavity of the primary polyp Variant: a subset of individuals of a particular species or oozooid, one dorsal, two lateral, and one ventral, ex- that deviates in form, typical anatomy, or behavior tending the length of the colony Verrucae: wart-like projections Mesozooid: a polyp intermediate in size and structure between autozooids and siphonozooids, found on the Warty club: a club (sclerite) with the head end orna- rachis and polyp leaves in some species of Pennatula mented with warts and Pteroeides; apparently having an exhalent current function (Williams et al., 2012) Superfamily Pennatuloidea (Adapted from Oozooid: the primary polyp, also referred to as axial Bayer et al., 1983 and Dolan, 2008) polyp, supported by the calcareous axis Acrozooid: large, white globular polyps clustered at the Peduncle: the lower part of the colony devoid of polyps distal-most region of the ventral rachis in some shallow- and polyp leaves, also referred to as the foot, that an- water species of Pteroeides; they are presumably incipi- chors in the substrate ent vegetative buds and have an asexual reproductive function (Williams et al., 2012) Pentamorphic: possessing five types of polyps (oo- zooids, autozooids, siphonozooids, mesozooids, and Anthocodia: the distal part of a polyp bearing the mouth acrozooids) and the tentacles Polyp leaves: the flattened expansions bearing the sec- Asulcal side: the side of the polyp opposite the ondary polyps siphonoglyph Rachis: the polypiferous (polyp bearing) part of the Autozooid: a polyp with eight well-developed tentacles colony and mesenteries; the larger polyp in dimorphic species, often termed simply “polyp” and typically responsible Rays: radiating bundles of large sclerites on the under- for food capture and reproduction side of polyp leaves Axis: the calcareous inner supporting structure, some- Siphonozooid: a polyp with a strongly developed sipho- times called a rod, that in pennatuloideans is composed noglyph and with reduced or no tentacles, usually much of high-magnesium calcite with very small amounts of smaller than the autozooids, with the primary function low-magnesium calcite and aragonite of irrigating colonies with seawater Calyces: cylindrical or wartlike projecting anthostele; Stalk: the barren basal part of the colony above the pe- some have between two and eight teeth and contain duncle or holdfast sclerites (as in Balticina and Pennatula) Stipules: clusters of siphonozooids forming small acces- Clavate: a club-shaped or terminally enlarged branch sory lobes at the dorsal ends of polyp leaves Dorsal side: the side of the colony derived from the asul- Teeth: the more or less pointed lobes of the calicular cal side of the oozooid; the side adjacent to the oldest margin, usually stiffened by sclerites, and typically num- (and often largest) autozooids in pennatuloideans hav- bering between two and eight when present ing well-developed leaves Terete: cylindrical but usually slightly tapered at both Dorsal track: the more or less naked strip extending ends and with a smooth surface along the rachis between the oldest polyps of the polyp leaves (corresponding to the asulcal side of the primary Terminal polyp: the distal end of the primary polyp zooid) Appendix I 407 Trimorphism: the presence of three types of polyps (au- Dactylozooid: one of three types of stylasterid polyps tozoids, siphonozooids, and mesozooids) that specializes in defense and food acquisition, each composed of a single mouthless tentacle Ventral track: the narrow naked strip extending along the rachis between the youngest polyps of the polyp Dichotomous: bifurcating leaves Distichoporine: see pore rows and Fig. 16-2 Family Stylasteridae (Adapted from Cairns, Flabellate: fan-shaped 2011b) Gastropore: the surface pore (usually round) of the Abcauline: on the side facing toward the growing tip of stylasterid skeleton that is associated with the gastrozo- a branch oid (see Figs. 16-1 and 16-2) Adcauline: on the side facing away from the growing tip Gastropore tube: the tube that contains the gastrozooid of a branch (see Fig. 16-5) and, when present, the gastrostyle; it is usually a simple cylinder but may be double-chambered (see Fig. 16-4) Ampulla (ampullae): the skeletal encasement of the gonophore, often forming a prominent hemispherical Gastrostyle: the vertical, spinose, axial structure that structure (see Fig. 16-2) projects from the base of the gastropore tube of various genera (see Fig. 16-4) Anastomotic: forming a reticulate sieve Gastrozooid: one of three types of stylasterid polyps Branchlet: a smaller distal branch of a colony that specializes in obtaining food, usually consisting of a polyp having a ring of tentacles that encircle a mouth Clavate: club-shaped Gonochoric: either male or female, not hermaphroditic Coenosteum: the calcium carbonate skeleton of the stylasterid colony, usually aragonitic, but occasionally Gonophore: one of three types of stylasterid polyps that calcitic or partially calcitic specializes in housing the reproductive gametes Corallum: the calcareous skeleton of the entire colony Lamellate: composed of or arranged in layers (sometimes called the colony) Lanceolate: in reference to a gastrostyle, narrow and ta- Cyclosystem: a functional unit of some stylasterid gen- pered, like a lance era in which a gastropore is surrounded by a circle of dactylopores (see Fig. 16-1) Linear-imbricate coenosteal texture: branch surface cov- ered with linear parallel strips of imbricating scales (see Dactylopore: the surface pore of the stylasterid skeleton Fig. 16-3) that is associated with a dactylozooid (see Fig. 16-2) Monotypic: a genus having but one species Dactylopore spine: a projection from the branch surface usually adjacent to a dactylopore, U-shaped or horse- Nematophore: large concentrations of nematocysts shoe-shaped in cross section (see Fig. 16-5) Nematopore: a shallow skeletal pit in the coenosteum Dactylostyle: a single row or crowded multiple rows of that houses a nematophore (see Fig. 16-3) small cylindrical pillars that occur on the outer wall of the dactylopore tube and/or dactylopore spine Papilla (papillae): conical, apically perforate nemato- cyst-bearing structures that occur on the coenosteum of Dactylotome: the open area, usually a slit, which allows some species the dactylozooid to bend toward the polyp (see Figs. 16-3 and 16-4) Porcellaneous: smooth and without pores 408 Professional Paper NMFS 23 Pore rows: gastropores unilinearly arranged on branch Reticulate-granular coenosteal texture: branch surface edges, flanked on either side by a row of dactylopore covered with granular strips arranged in a reticulate spines; sometimes called a distichoporine arrangement manner (see Fig. 16-1) (see Fig. 16-2) Ring palisade: a ring or girdle of small cylindrical pillars Pseudocyclosystem: a semi-circular arrangement of dac- that project from the wall of the gastropore tube near tylopore spines that resembles a cyclosystem the level of the gastrostyles tip (see Fig. 16-4) Pseudoseptum (pseudosepta): the roughly triangular- Supernumerary: additional, or extra, isolated dactylo- shaped coenosteum that separates the dactylotomes in a pores beyond those found in the cyclosystem (see Fig. cyclosystem (see Figs. 16-1, 16-3, and 16-4) 16-1) Appendix II 409 Appendix II. Coral species reported from Alaska waters This list includes the current known deep-sea coral spe- Steamer Albatross expeditions and National Marine cies (Phylum Cnidaria, Class Anthozoa, and Class Hy- Fisheries Service stock assessment surveys, and past re- drozoa) and their reported distribution and depth range search expeditions. Single values in the depth range col- in the 6 regions of Alaska: eastern Gulf of Alaska (1), umn represent depths known from only a single speci- western Gulf of Alaska (2), Gulf of Alaska Seamount men or several specimens from the same location. The Province (3), Aleutian Islands (4), Bering Sea (5), and World Register of Marine Species was used as the taxo- Arctic (6). The list was drawn from historical col- nomic authority. lections, including specimens from the U.S. Fisheries Higher taxon Species Distribution Depth range (m) Phylum Cnidaria Class Anthozoa Subclass Hexacorallia Order Antipatharia Family Cladopathidae Chrysopathes formosa Opresko, 2003 1 619–756 Chrysopathes speciosa Opresko, 2003 1 648–914 Heteropathes pacifica (Opresko, 2005) 3 3563–4663 Trissopathes pseudotristicha Opresko, 2003 3,4 2306–2828 Family Schizopathidae Alternatipathes mirabilis Opresko and Molodtsova, 2021 3 4600–4685 Bathypathes alaskensis Opresko and Molodtsova, 2021 1,3,4 329–1837 Bathypathes patula Brook, 1889 2 4950 Bathypathes ptiloides Opresko and Molodtsova, 2021 3 4491–4676 Bathypathes tiburonae Opresko and Molodtsova, 2021 2 3280–3356 Bathypathes sp. A 2 3275 Bathypathes sp. B 3 4497 Dendrobathypathes boutillieri Opresko, 2005 1,3,4 859–2161 Dendrobathypathes sp. 3 1652–2634 Lillipathes wingi Opresko, 2005 1,3,5 600–909 Lillipathes sp. A 1,3 600–2738 Parantipathes euantha (Pasternak, 1958) 3,4 862–2762 Parantipathes pluma Opresko and Molodtsova, 2021 4,5 602–1562 Parantipathes sp. A 3 2715–2819 Order Scleractinia Family Caryophylliidae Caryophyllia (Caryophyllia) alaskensis Vaughan, 1941 1,4,5 90–1397 Caryophyllia (Caryophyllia) arnoldi Vaughan, 1900 1,2,3,4 21–1702 Crispatotrochus foxi (Durham and Barnard, 1952) 4 135–702 Desmophyllum dianthus (Esper, 1794) 1 398 Family Dendrophylliidae Balanophyllia (Balanophyllia) elegans Verrill, 1864 1 15–22 Family Flabellidae Flabellum (Flabellum) oclairi Cairns, sp. nov. 4 27–507 Javania borealis Cairns, 1994 4 17–1266 Javania cailleti (Duchassaing and Michelotti, 1864) 3,4 150–2635 Family Fungiacyathidae Fungiacyathus (Bathyactis) marenzelleri (Vaughan, 1906) 2,3 2370–6328 Family Micrabaciidae Leptopenus discus Moseley, 1880 2 4820–5000 Order Zoantharia Family Epizoanthidae Epizoanthus scotinus Wood, 1957 4 116 Appendix continued 410 Professional Paper NMFS 23 Higher taxon Species Distribution Depth range (m) Family Parazoanthidae Mesozoanthus sp. 1,2,5 123–625 Zibrowius cf. ammophilus 3 745–2511 gen. nov., sp. nov. 1 87–107 Subclass Octocorallia Order Malacalcyonacea Family Alcyoniidae Alcyonium pacificum Yamada, 1950 4 6–12 Alcyonium sp. 4 5–600 Gersemia fruticosa (Sars, 1860) 2,4 1005–2564 Gersemia lambi Williams, 2013 1 1–30 Gersemia rubiformis (Ehrenberg, 1834) 1,2,4,5,6 3–90 Gersemia sp. 3 897–1095 Family Clavulariidae Clavularia armata Thomson, 1927 3 2667–2730 Clavularia eburnea Kükenthal, 1906 4 715 Clavularia rigida Broch, 1935 4 3277 Clavularia sp. A 2,4,5 82–990 Family Gorgoniidae Callistephanus pacificus Nutting, 1912 1,2,3,4,5 96–2778 Callistephanus simplex (Nutting, 1909) 1,3,4 500–1352 Family Malacalcyonacea Alaskagorgia aleutiana Sánchez and Cairns, 2004 4 87–1255 incertae sedis Alaskagorgia splendicitrina Horvath and Stone, 2018 4 184 Calcigorgia beringi (Nutting, 1912) 1,4 87–1933 Calcigorgia gigantea Matsumoto et al., 2019 4 135–381 Calcigorgia japonica Dautova, 2007 4 57–2210 Calcigorgia matua Dautova, 2018 4 300 Calcigorgia spiculifera Broch, 1935 1,4 12–512 Cryogorgia koolsae Williams, 2005 4 18–412 Elenanthus cf. violaceus 4 22–352 Family Paramuriceidae Acanthogorgia spissa Kükenthal, 1908 4 1092–2087 Acanthogorgia sp. 4 843–1692 Muriceides cylindrica Nutting, 1912 4 393–881 Muriceides nigra Nutting, 1912 4 78–1195 Order Scleralcyonacea Family Chrysogorgiidae Chrysogorgia sp. A 4 1913–2514 Chrysogorgia sp. B 3 3358–4768 Chrysogorgia sp. C 4 1359 Pseudochrysogorgia sp. A 3 1854–2227 Pseudochrysogorgia sp. B 3 2787 Pseudochrysogorgia sp. C 3 4103-4712 Radicipes stonei Cordeiro et al., 2017 3,4 1612–3580 Family Coralliidae Hemicorallium sp. 3 1677–1779 Paragorgia jamesi Herrera and Shank, 2016 1,2,3 372–944 Paragorgia arborea var. pacifica Verrill, 1922 2,3,4,5 21–1115 Paragorgia arborea var. pacifica morph nodosa 2,3,4,5 299–2022 Paragorgia stephencairnsi Sánchez, 2005 1,2,4 171–741 Paragorgia sp. A 3 716–1112 Paragorgia sp. B 3 730–1376 Sibogagorgia cauliflora Herrera et al., 2010 3 2766 Appendix continued Appendix II 411 Higher taxon Species Distribution Depth range (m) Subfamily Anthomastinae Heteropolypus japonicus (Nutting, 1912) 4 400–424 Heteropolypus ritteri (Nutting, 1909) 4 241–429 Heteropolypus sp. 1,3,4,5 29–4216 Pseudoanthomastus sp. 3 1122–2642 Family Keratoisididae Bathygorgia profunda Wright, 1885 3,4 2372–4784 Bathygorgia sp. A 3,4 603–1925 Isidella tentaculum Etnoyer, 2008 1,2,3,4,5 115–1608 Isidella sp. A 3 3839–4643 Isidella sp. B 3 3909–4643 Keratoisis sp. A 4 1584–2035 Keratoisis sp. B 4 1716–2044 Keratoisis sp. C 4 1715–1988 Keratoisis sp. D 3 598–2689 Keratoisis sp. E 3 3990–4373 Keratoisis sp. F 1,4 574–1563 Keratoisis sp. G 3 1689–1726 Orstomisis sp. 4 2674–2826 Family Primnoidae Arthrogorgia kinoshitai Bayer, 1952 4 150–1309 Arthrogorgia otsukai Bayer, 1952 4 1332–1348 Arthrogorgia utinomii Bayer, 1996 4 163–882 Callogorgia compressa (Verrill, 1865) 4 82–1341 Callogorgia fraseri (Hickson, 1915) 2,4 52–1341 Calyptrophora laevispinosa Cairns, 2007 3 1754–3531 Narella abyssalis Cairns and Baco, 2007 3 4559–4633 Narella alaskensis Cairns and Baco, 2007 3 2216–3204 Narella arbuscula Cairns and Baco, 2007 3 2710–3432 Narella bayeri Cairns and Baco, 2007 2,3 3280–4092 Narella cristata Cairns and Baco, 2007 3 3022–3358 Parastenella doederleini (Wright and Studer, 1889) 4 1746–2539 Parastenella gymnogaster Cairns, 2007 3 2193–3074 Parastenella ramosa (Studer, 1894) 1,3,4,5 667–1937 Plumarella aleutiana Cairns, 2011 3,4,5 79–2828 Plumarella echinata Cairns, 2011 4 150–1692 Plumarella hapala Cairns, 2011 4 120–407 Plumarella nuttingi Cairns, 2011 4 350–888 Plumarella profunda Cairns, 2011 4 2514 Plumarella robusta Cairns, 2011 4 115–1061 Plumarella spicata Nutting, 1912 4 712–1913 Plumarella superba (Nutting, 1912) 4 40–715 Primnoa pacifica Kinoshita, 1907 1,2,4,5 6–899 Primnoa pacifica var. willeyi (Hickson, 1915) 1,2,3,4 27–863 Primnoa wingi Cairns and Bayer, 2005 4,5 217–1280 Thouarella cristata Cairns, 2011 4 91–744 Thouarella trilineata Cairns, 2011 4 97–1266 Family Sarcodictyonidae Sarcodictyon incrustans (Broch, 1935) 1 15–238 Sarcodictyon sp. A 3 2665–2811 Appendix continued 412 Professional Paper NMFS 23 Higher taxon Species Distribution Depth range (m) Superfamily Pennatuloidea Family Anthoptilidae Anthoptilum grandiflorum (Verrill, 1879) 1,4,5 600–2511 Anthoptilum murrayi Kölliker, 1880 3,4 744–2266 Anthoptilum sp. A 3 898–1095 Anthoptilum sp. B 3 797–1784 Anthoptilum sp. C 4 1667–2707 Anthoptilum sp. D 5 230–1116 Family Balticinidae Balticina californica (Moroff, 1902) 1,2,4 35–823 Balticina willemoesi (Kölliker, 1880) 1,2,4,5 18–310 Balticina sp. A 1,2,4,5 253–1391 Family Kophobelemnidae Kophobelemnon sp. 1 2869 Family Pennatulidae Pennatula aculeata Danielssen, 1860 1,5 2869–2972 Pennatula sp. 4 2239–2930 Ptilosarcus gurneyi (Gray, 1860) 1,2,4 3–274 Family Protoptilidae Protoptilum sp. 1,2 18–142 Family Stachyptilidae Stachyptilum superbum Studer, 1894 4 388 Family Umbellulidae Umbellula sp. 1,2,3,4 840–4656 Family Veretillidae Cavernularia vansyoci Williams, 2005 4 94–126 Family Virgulariidae Virgularia bromleyi Kölliker, 1880 5 1889 Virgularia cf. bromleyi 1,2,4 20–472 Virgularia cf. glacialis 4 122 Class Hydrozoa Order Anthoathecata Family Stylasteridae Crypthelia trophostega Fisher, 1938 4 138–1913 Cyclohelia lamellata Cairns, 1991 4 27–691 Distichopora borealis Fisher, 1938 1,4 137–1361 Errinopora dichotoma Lindner and Cairns, 2011 4 175–405 Errinopora disticha Lindner and Cairns, 2011 4 176–536 Errinopora fisheri Lindner and Cairns, 2011 4 455–458 Errinopora nanneca Fisher, 1938 4 22–594 Errinopora undulata Lindner and Cairns, 2011 4 366–425 Errinopora zarhyncha Fisher, 1938 4 115–1520 Stylantheca papillosa (Dall, 1884) 1 6–40 Stylaster alaskanus Fisher, 1938 4 146–2124 Stylaster brochi (Fisher, 1938) 1,2,4 20–455 Stylaster campylecus (Fisher, 1938) 1,2,4 79–1396 Stylaster crassiseptum Cairns and Lindner, 2011 4 291–531 Stylaster elassotomus Fisher, 1938 4 881 Stylaster leptostylus (Fisher, 1938) 4 518 Stylaster parageus columbiensis Cairns and Lindner, 2011 1 88–439 Stylaster parageus parageus (Fisher, 1938) 1,2 26–445 Stylaster repandus Cairns and Lindner, 2011 4 375–650 Stylaster stejnegeri (Fisher, 1938) 4 95 Stylaster trachystomus (Fisher, 1938) 4 110–366 Stylaster venustus (Verrill, 1868) 1 20 Stylaster verrillii (Dall, 1884) 1,2,4 11–401 Appendix II 413 Appendix III. Additional supporting references on Alaska corals Baco, A. R., and S. D. Cairns. and species associations. Am. Fish. Soc. Symp. 41:289–299. 2012. Comparing molecular variation to morphological species Miller, R. J., C. Juska, and J. Hocevar. designations in the deep-sea coral Narella reveals new in- 2015. Submarine canyons as coral and sponge habitat on the sights into seamount coral ranges. PLoS ONE 7(9):e45555. eastern Bering Sea slope. Global Ecol. Conserv. 4:85–94. https://doi.org/10.1371/journal.pone.0045555 https://doi.org/10.1016/j.gecco.2015.05.009 Bright-Diaz, L. M., K. B. Strychar, and T. C. Shirley. NMFS (National Marine Fisheries Service). 2011. Compounds from deep-sea bubblegum corals, Para- 2011. U.S. national bycatch report. NOAA Tech. Memo. gorgia arborea, elicit anti-predation behavior in fish. NMFS-F/SPO-117E, 508 p. Open Mar. Biol. J. 5:58–67. https://doi.org/10.2174/ 2013. U.S. national bycatch report, first edition update 1 (L. 1874450801105010058 R. Benaka, C. Rilling, E. E. Seney, and H. Winarsoo, eds.), Brugler, M. R., D. M. Opresko, and S. C. France. 57 p. Natl. Mar. Fish. Serv., U.S. Dep. Commer., Silver 2013. The evolutionary history of the order Antipatharia Spring, MD. [Available from https://www.fisheries.noaa. (Cnidaria: Anthozoa: Hexacorallia) as inferred from mi- gov/resource/document/national-bycatch-report.] tochondrial and nuclear DNA: implications for black coral 2016. U.S. national bycatch report, first edition update 2 (L. taxonomy and systematics. Zool. J. Linn. Soc. 169:312– R. Benaka, D. Bullock, J. Davis, E. E. Seney, and H. Winar- 361. https://doi.org/10.1111/zoj.12060 Bryan, T. L., and A. Metaxas. soo, eds.), 90 p. Natl. Mar. Fish. Serv., U.S. Dep. Commer., 2007. Predicting suitable habitat for deep-water gorgonian Silver Spring, MD. [Available from https://www.fisheries. corals on the Atlantic and Pacific continental margins of noaa.gov/resource/document/national-bycatch-report.] North America. Mar. Ecol. Prog. Ser. 330:113–126. https:// NOAA (National Oceanic and Atmospheric Administration). doi.org/10.3354/meps330113 2010. Report of the NOAA deep-sea coral and sponge eco- Cairns, S. D., R. P. Stone, E. A. Berntson, and S. A. Pomponi. systems exploration and research priorities workshop for 2017. Species discovery of deep-water corals and sponges in Alaska. Anchorage, AK, Sept. 14–15, 2010, 23 p. [Available U.S. waters (2007–2016). In The state of deep-sea coral from https://repository.library.noaa.gov/view/noaa/677.] and sponge ecosystems of the United States (T. F. Hourigan, Reynolds, J. R., S. C. Rooney, J. Heifetz, H. G. Greene, B. L. P. J. Etnoyer, and S. D. Cairns, eds.), p. 39–55. NOAA Tech. Norcross, and S. K. Shotwell. Memo. NMFS-OHC-4. 2012. Habitats and demersal fish communities in the vicin- Cimberg, R. L., T. Gerrodette, and K. Muzik. ity of Albatross Bank, Gulf of Alaska. In Seafloor geomor- 1981. Habitat requirements and expected distribution of phology as benthic habitat: GeoHAB Atlas of seafloor geo- Alaska coral. Final Report, Research Unit 601, VTN Ore- morphic features and benthic habitat (P. T. Harris and E. K. gon, Inc. U. S. Dep. Commer., NOAA, OCSEAP, Final Rep. Baker, eds.), p. 539–553. Elsevier, London. 54:207–308. Rooper, C. N., M. E. Wilkins, C. S. Rose, and C. Coon. Dunton, K. H. 2011. Modeling the impacts of bottom trawling and the 2012. Chukchi Sea offshore monitoring in drilling area subsequent recovery rates of sponges and corals in the (COMIDA): chemical and benthos (CAB) final report. Bu- Aleutian Islands, Alaska. Cont. Shelf Res. 31:1827–1834. reau of Ocean Energy Management, OCS Study BOEM https://doi.org/10.1016/j.csr.2011.08.003 2012-012, 311 p. [Available from https://espis.boem.gov/ Rooper, C. N., M. Zimmermann, M. M. Prescott, and A. J. final%20reports/5182.pdf.] Hermann. Goddard, P., R. E. Wilborn, C. N. Rooper, K. Williams, and R. 2014. Predictive models of coral and sponge distribution, Towler. abundance and diversity in bottom trawl surveys of the 2017. Results of the 2012 and 2014 underwater camera sur- Aleutian Islands, Alaska. Mar. Ecol. Prog. Ser. 503:157– veys of the Aleutian Islands. NOAA Tech. Memo. NMFS- 176. https://doi.org/10.3354/meps10710 AFSC-351, 505 p. Rooper, C. N., M. F. Sigler, P. Goddard, P. Malecha, R. Towler, K. Heifetz, J., B. L. Wing, R. P. Stone, P. W. Malecha, and D. L. Williams, R. Wilborn, and M. Zimmermann. Courtney. 2016. Validation and improvement of species distribution 2005. Corals of the Aleutian Islands. Fish. Oceanogr. 14 (Sup- models for structure-forming invertebrates in the eastern pl. 1):131–138. https://doi.org/10.1111/j.1365-2419.2005. Bering Sea with an independent survey. Mar. Ecol. Prog. 00371.x Ser. 551:117–130. https://doi.org/10.3354/meps11703 Heifetz, J., D. Woodby, J. R. Reynolds, and R. P. Stone. Sigler, M. F., C. N. Rooper, G. R. Hoff, R. P. Stone, R. A. McCon- 2007. Deep sea coral distribution and habitat in the Aleutian naughey, and T. K. Wilderbuer. Archipelago. North Pac. Res. Board Final Rep. 304, 303 p. 2015. Faunal features of submarine canyons on the eastern Heifetz, J., R. P. Stone, and S. K. Shotwell. Bering Sea slope. Mar. Ecol. Prog. Ser. 526:21–40. https:// 2009. Damage and disturbance to coral and sponge habitat of doi.org/10.3354/meps11201 the Aleutian Archipelago. Mar. Ecol. Prog. Ser. 397:295– Stone, R. P. 303. https://doi.org/10.3354/meps08304 2005. Exploring deep-sea coral habitat on the edge—Alaska’s Hoff, G. R., and B. Stevens. Aleutian Islands. J. Mar. Educ. 21(4):18–21. 2005. Faunal assemblage structure of the Patton Seamount Woodby, D., D. Carlile, and L. Hulbert. (Gulf of Alaska, USA). Alsk. Fish. Res. Bull. 11:27–36. 2009. Predictive modeling of coral distribution in the Central Malecha, P. W., R. P. Stone, and J. Heifetz. Aleutian Islands, USA. Mar. Ecol. Prog. Ser. 397:227–240. 2005. Living substrate in Alaska: distribution, abundance, https://doi.org/10.3354/meps08358