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Community structure of shallow water Alcyonacea (Anthozoa: Octocorallia) from the southern Tropical Eastern Pacific


Alcyonacea are sessile invertebrates, which can significantly shape the boundary layer in coral reefs and rocky habitats. Ecological aspects in this taxon have been well studied in the Caribbean, Mediterranean, and Indo-Pacific. With few recent exceptions, studies in the Eastern Pacific focused on taxonomy. We present a quantitative assessment of Alcyonacea communities from the southern Tropical Eastern Pacific, based on video transects in the Marine Reserve El Pelado. Seventeen species from the Plexauridae (8), Gorgoniidea (8), and Clavularidae (1) were identified, comprising 6963 colonies dominated by Muricea (86.7%), particularly M. plantaginea (48.6%). The overwhelming dominance of M. plantaginea was the most striking and previously unreported community trait, which contributed to a moderate Shannon entropy (n = 31, H mean 1.40, SD 0.22), equitability (n = 31, HE mean 0.16, SD 0.4), and species diversity expressed as effective number of species (n = 31, mean 4.16, SD 0.87). Few common species overprinted a more variable and subtle community pattern among rarer species, suggested in agglomerative hierarchical cluster analyses. Four species (M. plantaginea, M. purpurea, M. fruticosa and Leptogorgia alba) had the strongest influence on site groupings in the correspondence analysis between a principal component analysis of a Hellinger-transformed Alcyonacea species matrix and substrate categories, with filamentous turf algae and crustose coralline algae being the main determinants of site differentiation. Muricea plantaginea’s qualities of a keystone species, and the eurytopic and stenoecious distribution traits among some species are discussed. The invasive Carijoa riisei was confirmed as biological thereat to other Alcyonacea, and possible physiological distribution limitations are indicated.

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Fig. 1

Adapted from Glynn (2003)

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  1. Abeytia R, Guzmán HM, Breedy O (2013) Species composition and bathymetric distribution of gorgonians (Anthozoa: Octocorallia) on the Southern Mexican Pacific coast. Rev Biol Trop 61:157–1166

  2. Alvarez-Filip L, Dulvy NK, Gill JA, Côté IM, Watkinson AR (2009) Flattening of Caribbean coral reefs: region-wide declines in architectural complexity. Proc R Soc B 276:3019–3025.

  3. Bayer FM (1961) The shallow water Octocorallia of the West Indian region. Martinus Nijhoff, The Hague.

  4. Bielschowsky E (1929) Die Gorgonarien Westindien. 6. Die Familie Gorgoniidae, zugleich eine Revision. Zoologische Jahrbücher Supplement 16:63–234

  5. Blunt JW, Copp BR, Keyzers RA, Munro MHG, Prinsep MR (2014) Marine natural products. Nat Prod Rep 31:160–258

  6. Bo M, Lavorato A, Di Camillo CG, Poliseno A, Baquero A, Bavestrello G, Irei Y, Reimer JD (2012) Black coral assemblages from Machalilla National Park (Ecuador). Pac Sci 66:63–81

  7. Borcard D, Gillet F, Legendre P (2011) Numerical ecology with R. Springer, Berlin

  8. Breedy O, Cortés J (2008) Octocorals (Coelenterata: Anthozoa: Octocorallia) of Isla del Coco, Costa Rica. Rev Biol Trop 56:71–77

  9. Breedy O, Cortés J (2014) Gorgonias (Octocorallia: Gorgoniidae) de las aguas someras del Pacifico Norte de Costa Rica. Rev Biol Trop 62:43–62

  10. Breedy O, Guzman HM (2002) A revision of the genus Pacifigorgia (Coelenterata: Octocorallia: Gorgoniidae). Proc Biol Soc Wash 115:782–839

  11. Breedy O, Guzman HM (2003) Octocorals from Costa Rica: the genus Pacifigorgia (Coelenterata: Octocorallia: Gorgoniidae). Zootaxa 281:1–10

  12. Breedy O, Guzman HM (2005a) A new species of alcyonacean octocoral from the Galápagos Archipelago. J Mar Biol Assoc UK 85:801–807

  13. Breedy O, Guzman HM (2005b) A new species of Leptogorgia (Coelenterata: Octocorallia: Gorgoniidae) from shallow waters of the eastern Pacific. Zootaxa 899:1–11

  14. Breedy O, Guzman HM (2007) A revision of the genus Leptogorgia Milne Edwards and Haime, 1857 (Coelenterata: Octocorallia: Gorgoniidae) in the eastern Pacific. Zootaxa 1419:1–90

  15. Breedy O, Guzman HM (2011) A revision of the genus Heterogorgia Verrill, 1868 (Anthozoa: Octocorallia: Plexauridae). Zootaxa 2995:27–44

  16. Breedy O, Guzman HM (2014) A new species of alcyonacean octocoral from the Peruvian zoogeographic region. J Mar Biol Assoc UK.

  17. Breedy O, Guzman HM (2015) A revisión of the genus Muricea Lamouroux, 1821 (Anthozoa, Octocorallia) in the eastern Pacific. Part I: Eumuricea Verrill, 1869 revisited. ZooKeys 537:1–32

  18. Breedy O, Guzman HM (2016) A revisión of the genus Muricea Lamouroux, 1821 (Anthozoa, Octocorallia) in the eastern Pacific. Part II. ZooKeys 581:1–69

  19. Breedy O, Guzman HM, Vargas S (2009a) A revision of the genus Eugorgia Verrill, 1868 (Coelenterata: Octocorallia: Gorgoniidae). Zootaxa 2151:1–46

  20. Breedy O, Hickman CP Jr, Williams GC (2009b) Octocorals in the Galapagos Islands. Galapagos Res 66:27–31

  21. Breedy O (2009) Octocorals. In: Wehrtmann IS, Cortés J (eds) Marine biodiversity of Costa Rica, Central America. Springer, Berlin, pp 161–167.

  22. Cantera JR, Prahl VH, Neira OR (1987) Moluscos, crustaceos y equinodermos asociados a la gorgonia Leptogorgia alba Duchassaing y Michelotti 1864, en la isla de Gorgona, Colombia. Bol Ecotropica 17:3–23

  23. Cerrano C, Bavestrello G, Bianchi CN et al (2000) A catastrophic mass-mortality episode of gorgonians and other organisms in the Ligurian Sea (northwestern Mediterranean), summer 1999. Ecol Lett 3:284–293.

  24. Coll JC (1992) The chemistry and chemical ecology of octocorals (Coelenterata, Anthozoa, Octocorallia). Chem Rev 92:613–631

  25. Concepcion GT, Kahng SE, Crepeau MW, Franklin EC, Coles SL, Toonen RJ (2010) Resolving natural ranges and marine invasions in a globally distributed octocoral (genus Carijoa). Mar Ecol Prog Ser 401:113–127.

  26. Cuddington K (2012) Ecosystem engineers. In: Hastings A, Gross LJ (eds) Encyclopedia of theoretical ecology. University of California Press, Berkeley, pp 230–234

  27. Etnoyer P, Warernchuck J (2007) A catshark nursery in deep gorgonian fields in the Mississippi Canyon, Gulf of Mexico. Bull Mar Sci 88:553–559

  28. Fabricius K, Alderslade P (2001) Soft corals and sea fans: a comprehensive guide to the tropical shallow water genera of the central-west Pacific, the Indian Ocean and the Red Sea. Australian Institute of Marine Science, Townsville

  29. Fabricius KE, McCorry D (2006) Changes in octocoral communities and benthic cover along a water quality gradient in the reefs of Hong Kong. Mar Pollut Bull 52:22–33.

  30. Fautin DG (2002) Reproduction of cnidaria. Can J Zool 80:1735–1754

  31. Fiedler PC, Lavine MF (2016) Oceanographic conditions of the Eastern Tropical Pacific. In: Glynn PW, Manzello D, Enochs IC (eds) Coral reefs of the Eastern Tropical Pacific. Springer, Berlin, pp 59–83

  32. Fiedler P (1992) Climatologies and variability of eastern tropical Pacific surface waters. NOAA Tech Rpt. NMFS 109

  33. Figueroa J (2015) Contribución al conocimiento de la fauna de octocorales (Octocorallia: Gorgoniidae) en el arrecife rocos del islote Los Ahorcados, Manabí, Ecuador. Tesis de grado, Univesridad Laica Eloy Alfaro de Manabí, Manbí Ecuador

  34. Fréchette M, Butman CA, Geyer WR (1989) The importance of boundary-layer flows in supplying phytoplankton to the benthic suspension feeder, Mytilus edulis. Limnol Oceanogr 34:19–36

  35. García-Parrado P, Alcolado PM (1996) Catálogo de los octocorales (Cnidaria) de Cuba, con comentarios sobre su taxonomía. Avicennia 4:41–45

  36. Glynn PW, Alvarado JJ, Banks S et al (2016a) Eastern Pacific Coral Reef Provinces, coral community structure and composition. An overview. In: Glynn PW, Manzello D, Enochs IC (eds) Coral reefs of the Eastern Tropical Pacific. Springer, Berlin, pp 107–176

  37. Glynn PW (2003) Coral communities and coral reefs of Ecuador. In: Cortés J (ed) Latin American Coral Reefs. Elsevier, Amsterdam, pp 449–472

  38. Glynn PW, Mones AB, Podésta GP et al (2016b) El Nino Southern oscillation: effects on Eastern Pacific coral reefs and associated biota. In: Glynn PW, Manzello D, Enochs IC (eds) Coral reefs of the Eastern Tropical Pacific. Springer, Berlin, pp 251–290

  39. Gomez CG, Guzman HM, Gonzalez A, Breedy O (2014) Survival, growth, and recruitment of octocoral species (Coelenterata: Octocorallia) in Coiba National Park, Pacific Panama. Bull Mar Sci 90(2):623–650.

  40. González-Ortiz V, Egea LG, Jiménez-Ramos R et al (2014) Interactions between seagrass complexity, hydrodynamic F1 and biomixing alter food availability for associated filter-feeding organisms. PLoS ONE 9:e104949.

  41. Government of Ecuador (2017) Accessed 10 Feb 2017

  42. Grigg RW (1975) Age structure of a longevous coral: a relative index of habitat suitability and stability. Am Nat 109:647–657.

  43. Grigg RW (1977) Population dynamics of two gorgonian corals. Ecology 58:278–290.

  44. Grigg RW (2003) Invasion of a deep black coral bed by an alien species, Carijoa riisei, off Maui, Hawaii. Coral Reefs 22:121–122.

  45. Grigg RW (1970) Ecology and population dynamics of the gorgonians, Muricea california and M. fruticosa. Dissertation, University of California at San Diego

  46. Gutierrez M, Capson TL, Guzman HM et al (2006) Antiplasmodial metabolites isolated from the marine octocoral Muricea austera. J Nat Prod 69:1379–1383.

  47. Guzman HM, Breedy O (2008) Leptogorgia christiae (Octocorallia: Gorgoniidae) a new shallow water gorgonian from Pacific Panama. J Mar Biol Assoc UK 88:719–722.

  48. Guzman HM, Breedy O (2011) Pacifigorgia marviva (Anthozoa: Octocorallia) a new species from Coiba National Park, Pacific Panama. J Mar Biol Assoc 92:693–698.

  49. Guzman HM, Guevara CA, Breedy O (2004) Distribution, diversity, and conservation of coral reefs and coral communities in the largest marine protected area of the Pacific Panama (Coiba Island). Environ Conserv 31:111–121.

  50. Gálvez H, Regalado J (2007) Características de las precipitaciones, la temperatura del aire y los vientos en la costa Ecuatoriana. Acta Oceanográfica del Pacífico 14:201–205

  51. Harrison PL (2011) Sexual reproduction of scleractinian corals. In: Dubinsky Z, Stambler N (eds) Coral reefs: an ecosystem in transition. Springer, Dordrecht

  52. Hickman CP Jr (2008) A field guide to corals and other radiates of Galápagos. Sugar Spring Press, Lexington

  53. Hickson SJ (1928) The Gorgonacea of Panama Bay together with a description of one species from the Galápagos Islands and one of Trinidad. Videnskavelige Meddelelser fra den naturhistoriske Forening i Kovenhavn for Aarene 85:325–422

  54. Hill M (1973) Diversity and evenness: a unifying notation and its consequences. Ecology 54:427–432

  55. Irlandi EA (1996) The effects of seagrass patch size and energy regime on growth of a suspension-feeding bivalve. J Mar Res 54:161–185

  56. Jost L (2006) Entropy and diversity. Oikos 113:363–375

  57. Kahng SE, Benayahu Y, Lasker HR (2011) Sexual reproduction in octocorals. Mar Ecol Prog Ser 443:265.

  58. Kahng SE, Benayahu Y, Wagner D, Rothe N (2008) Sexual reproduction in the invasive octo-coral Carijoa riisei in Hawaii. Bull Mar Sci 82:1–17

  59. Kahng SE, Grigg W (2005) Impact of an alien octocoral, Carijoa riisei, on black corals in Hawaii. Coral Reefs 24:556–562.

  60. Kahng SE (2006) Ecology and ecological impact of an alien octocoral, Carijoa riisei, in Hawaii. Dissertation, University of Hawaii

  61. Kelman D, Kashman Y, Hill RT, Rosenberg E, Loya Y (2009) Chemical warfare in the sea: the search for antibiotics from Red Sea corals and sponges. Pure Appl Chem 81:1113–1121

  62. Kohler KE, Gill SM (2006) Coral point count with excel extensions (CPCe): a visual basic program for the determination of coral and substrate coverage using random point count methodology. Comput Geosci 32:1259–1269.

  63. Lasker H (1985) Prey preferences and browsing pressure of the butterflyfish Chaetodon capistratus on Caribbean gorgonians. Mar Ecol Prog Ser 21:213–220

  64. Lasker HR, Coffroth MA (1988) Temporal and spatial variability among grazers: variability in the distribution of the gastropod Cyphoma gibbosum on octocorals. Mar Ecol Prog Ser 3:285–295.

  65. Legendre P, Legendre LFJ (2012) Numerical ecology, vol 24. Elsevier, Amsterdam

  66. Lenz EA, Bramanti L, Lasker HR, Edmunds PJ (2015) Long-term variation of octocoral populations in St. John, US Virgin Islands. Coral Reefs 34:1099.

  67. MacArthur R (1965) Patterns of species diversity. Biol Rev 40:510–533

  68. Mosher C, Watling L (2009) Partners for life: a brittle star and its octocoral host. Mar Ecol Prog Ser 397:81–88.

  69. Neira OR, Barba PR, Pardo RA (1992) Equinodermos del Parque Nacional Natural Ensenada de Utria (Pacifico Colombiano). An Inst Invest Mar Punta Betin 21:77–83

  70. Oksanen J, Blanchet FG, Friendly M et al (2017) vegan: Community Ecology Package. R package version 2.4-2. Accessed 12 July 2017

  71. Paradis E, Claude J, Strimmer K (2004) APE: analyses of phylogenetics and evolution in R language. Bioinformatics 20:289–290

  72. Patton WK (1972) Studies on the animal symbionts of the gorgonian coral, Leptogorgia virgulata (Lamarck). Bull Mar Sci 22:419–431

  73. Prahl HV, Escobar D, Molina G (1986) Octocorales (Octocorallia: Gorgoniidae y Plexauridae) de aguas someras del Pacífico Colombiano. Rev Biol Trop 34:3–33

  74. R Development Core Team (2012) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Austria. ISBN 3-900051-07-0.

  75. Ramos GE (1995) Neopontonides henryvonprahli, una nueva especie de camarón pontoníno del Pacífico de Colombia (Decapoda: Palaemonidae) simbionte de las gorgonias Muricea robusta y Leptogorgia alba. Rev Biol Trop 43:231–237

  76. Reyes-Bonilla H, Sinsel-Duarte F, Arizpe O (1997) Gorgonias y corales pétreos (Anthozoa: Gorgonacea y Scleractinia) de Cabo Pulmo, México. Rev Biol Trop 45:1439–1443

  77. Rivera F, Martínez PC (2011) Guía photográfica de corales y octocorales, Parque Nacional Machalilla y Reserva de Producción Faunística Marino Costera Puntilla de Santa Elena, Ecuador

  78. Rocha J, Peixe L, Gomes NCM, Calado R (2011) Cnidarians as a source of new marine bioactive compounds—an overview of the last decade and future steps for bioprospecting. Mar Drugs 9:1860–1886.

  79. Santangelo G, Carletti E, Maggi E, Bramanti L (2003) Reproduction and population sexual structure of the overexploited Mediterranean red coral Corallium rubrum. Mar Ecol Prog Ser 248:99–108

  80. Shannon CE (1948) A mathematical theory of communication. Bell Syst Tech J 27(3):379–423.

  81. Simpson A (2009) Reproduction in Octocorals (subclass Octocorallia): a review of published literature. Version 16 July 2009. In: Deep-sea corals portal. Accessed 19 Feb 2017

  82. Sinsel-Duarte F (1991) Taxonomía, distribución, abundáncia y diversidad de los gorgóniddos de Cabo Pulmo, Baja California Sur, Mexoco. BSc thesis, Departamento de Biología Marina, Universidad Autónoma de Baja California Sur, La Paz, Mexico

  83. Soler-Hurtado MM, López-González PJ (2012) Two new gorgonian species (Anthozoa: Octocorallia: Gorgoniidae) from Ecuador (Eastern Pacific) Mar. Biol Res 8:380–387

  84. Soler-Hurtado MM, Machordon A, Munus J, López Golnzáles PJ (2016) New records of the genera Leptogorgia, Pacifigorgia and Eugorgia (Octocorallia: Gorgoniidae) from Ecuador, with a description of a new species. Scientia Marian 80:369–394.

  85. Stiasny G (1941) Studien über alcyonaria und gorgonarias I–V. (Parerga und Paralipomena). Zoologosche Anzeiger 133:268–271

  86. Stiayny G (1943) Gorgonaria von Panama. Aus der Sammlung Dr. Th. Mortensen, Zoologisk Museum, Kopenhagen. Videnskavelige Meddelelser fra den naturhistoriske Forening I Kovenhaven for Aarene 197:59–103

  87. Strub PT, Mesias JM, Montecino V, Rutllant J, Salinas SH (1998) Coastal ocean circulation off western South America: 273–313. In: Robinson AR, Brink KH (eds) The Sea, vol 11. The global coastal ocean: regional studies and syntheses. Wiley, New York, pp 273–313

  88. Sánchez JA, Ardila NE, Andrade J, Dueñas LF, Navas R, Ballesteros D (2014) Octocoral densities and mortalities in Gorgona Island, Colombia, Tropical Eastern Pacific. Rev Biol Trop 62(1):209–219

  89. Sánchez JA, Ballesteros D (2014) The invasive snowflake coral (Carijoa riisei) in the Tropical Eastern Pacific, Colombia. Rev Biol Trop 62:197–207

  90. Sánchez JA, Gómez CE, Escobar D, Dueñas LF (2011) Diversidad, abundancia y amenazas de los octocorales de la isla Malpelo, Pacífico Oriental Tropical, Colombia. Boletín de Investigaciones Marinas y Costeras 40:139–154

  91. Taylor ML, Yesson C, Angew DJ, Mitchell RE, Rogers AD (2013) Using fisheries by-catch data to predict octocoral habitat suitability around South Georgia. J Biogeogr 40:1600–1701

  92. Tsounis G, Edmunds PJ (2017) Three decades of coral reef community dynamics in St. John, USVI: a contrast of scleractinians and octocorals. Ecosphere 8:e01646.

  93. Valenciennes A (1846) Zoophytes. In: Dupetit-Thouars A. Voyage autour du monde sur la frégatte la Vénus, pendant les annés 1836–1839. Atlas de Zoologie pp 1–15

  94. Van Oppen MJH, Miegog JC, Sanchez CA, Fabricius KE (2005) Diversity of algal endosymbionts (zooxanthellae) in octocorals: the roles of geography and host relationships. Mol Ecol 14:2403–2417.

  95. Vargas S, Breedy O, Guzman HM (2010) The phylogeny of Pacifigorgia (Coelenterata, Octocorallia, Gorgoniidae): a case study of the use of continuous characters in the systematic of the Octocorallia. Zoosystema 32:5–18.

  96. Vargas S, Guzman HM, Breedy O (2008) Distribution patterns of the genus Pacifigorgia (Octocorallia: Gorgoniidae): track compatibility analysis and parsimony analysis of endemicity. J Biogeogr 35:241–247.

  97. Vogel S (1994) Life in moving fluids. Princeton University Press, Princeton

  98. Vreeland HV, Lasker HR (1989) Selective feeding of the polychaeta Hermodice carunculata Pallas on Caribbean gorgonians. J Exp Mar Biol Ecol 129:265–267

  99. Wainwright SA, Biggs WD, Currey JD, Gosline JM (1976) Mechanical design in organisms. Princeton University Press, Princeton

  100. Wanatabe S, Metaxas A, Sameoto J, Lawton P (2009) Patterns in abundance and size of two deep-water gorgonian octocorals, in relation to depth and substrate features off Nova Scotia. Deep Sea Res Part 1 Oceanogr Res Pap 56:2235–2248

  101. Wang C, Deser C, Yu J-Y, DiNezio P, Clement A (2016) El Nino and Southern oscillation (ENSO): a review. In: Glynn PW, Manzello D, Enochs IC (eds) Coral reefs of the Eastern Tropical Pacific. Springer, Berlin, pp 85–106

  102. Williams GC, Breedy O (2004) The Panamic gorgonian genus Pacifigorgia (Octocorallia: Gorgoniidae) in the Galápagos Archipelago, with descriptions of three new species. Proc Calif Acad Sci 55:55–88

  103. Yoshioka PM (1994) Size-specific life history pattern of a shallow water gorgonian. J Exp Mar Biol Ecol 184:111–222

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Sascha Steiner was supported by the Prometeo Program (B 2015–0296) of the National Secretariat of Higher Education, Science and Technology (SENESCYT), Ecuador. Field surveys and the collection of samples were carried out under the research permit MAE013–15 IC–FAU–DPSE–MA, granted to the S. Steiner by the Ministry of the Environment of Ecuador. Dive excursions were funded by SENESCYT Project PIC14–CENAIM–001 granted to J. Rodríguez, at the National Center for Aquaculture and Marine Research (CENAIM). Sclerites were examined at the Research Centre in Microscopic Structures (CIEMIC), University of Costa Rica. We are grateful to O. Breedy for the assistance with the identification of specimens.

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Correspondence to Sascha C. C. Steiner.

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Steiner, S.C.C., Riegl, B., Lavorato, A. et al. Community structure of shallow water Alcyonacea (Anthozoa: Octocorallia) from the southern Tropical Eastern Pacific. Ecol Res 33, 457–469 (2018).

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  • Alcyonacea
  • Community structure
  • Eastern Pacific
  • Equatorial Front
  • Keystone species