A Generic Revision and Phylogenetic Analysis of the Primnoidae

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13. Thouarella (Diplocalyptra) Kinoshita, 1908 34 14. Thouarella (Epithouarella) Kükenthal, 1915 35 15. Metafannyella, new genus. 35 16. Fannyella (Fannyella) Gray, 1872 35 17. Fannyella (Scyphogorgia), new subgenus 36 18. Fannyella (Cyathogorgia), new subgenus 36 19. Onogorgia, new genus 37 20. Pyrogorgia, new genus 37 21. Amphilaphis Studer and Wright in Studer, 1887 38 22. Mirostenella Bayer, 1988 38 23. Acanthoprimnoa Cairns and Bayer, 2004 39 24. Plumarella Gray, 1870 39 25. Callogorgia Gray, 1858 40 26. Fanellia Gray, 1870 40 27. Paranarella Cairns, 2007 41 28. Primnoa Lamouroux, 1812 41 29. Australogorgia, new genus 42 Australogorgia aldersladei, new species 42 30. Narella Gray, 1870 43 31. Arthrogorgia Kükenthal, 1908 43 32. Paracalyptrophora Kinoshita, 1908 44 33. Calyptrophora Gray, 1866 44 34. Tokoprymno Bayer, 1996 45 35. Parastenella Versluys, 1906 45 36. Candidella Bayer, 1954 46 37. Microprimnoa Bayer and Stefani, 1989 46 38. Pterostenella Versluys, 1906 47 39. Perissogorgia Bayer and Stefani, 1989 47 40. Dasystenella Versluys, 1906 47 41. Pseudoplumarella Kükenthal, 1915 48 FIGURES 3–19 49 REFERENCES 67 INDEX 73
A Generic Revision and Phylogenetic Analysis of the Primnoidae (Cnidaria: Octocorallia) Stephen D. Cairns and † Frederick M. Bayer, Smithsonian Institution, Department of Invertebrate Zoology, P.O. Box 27012, MRC 163, Washington, D.C. 20013-7012, USA. Manuscript received 7 March 2008; accepted 25 June 2008. † Deceased 2 October 2007. INTRODUCTION Among the 44 families of Octocorallia (Williams and Cairns, 2005, updated), the Primnoidae ranks fourth in number of species (233/3200 = 7.3%, see Tables 1, 4) and third in number of genera (36/340 = 10.6%), resulting in an average of 6.5 species per genus. No confi rmed fossil species are known. Primnoids occur worldwide at depths of 8– 5,850 m, although they are most common at slope and upper abyssal depths (Table 1), the few shallow records being uncommon. The primnoids thus may be considered to be the quintessential deepwater octocoral family, only some pennatulaceans occurring in deeper water. It is tempting to speculate that the modifi cation of their imbricate external calycular scales is the key adaptation for their success in deep water; however, two isidid subfamilies, many of which occur at shelf depths, also have very similar sclerite morphology. Aside from the innate taxonomic interest of the family and the beauty of their calycular architecture, primnoids serve important ecological and geophysical roles. Because of their large size (e.g., Primnoa occurs up to 2 m in height and 7 m wide) and local abundance (sometimes occurring in large monospecifi c fi elds), some primnoids, such as Primnoa, Narella, and Callogorgia, form habitat for fi sh (Etnoyer and Morgan, 2005; Stone, 2006; Etnoyer and Warrick, 2007) and other invertebrates (Krieger and Wing, 2002; Buhl-Mortensen and Mortensen, 2004, 2005), especially on seamounts (Cairns and Baco, 2007; Rogers et al., 2007; Cairns and Bayer, 2007 [2008]), and on deepwater coral banks (Cairns and Bayer, 2005). Also, because of their solid, layered axis and purported long life span (Andrews et al., 2002; Risk et al., 2002), various isotopic analyses of their axes can be used to determine paleotemperatures (Heikoop et al., 2002; Sinclair et al., 2005; Sherwood et al., 2005). The following abbreviations are used in the text. Museums AM Australian Museum, Sydney BM The Natural History Museum, London MNHNP Muséum National d’Histoire Naturelle, Paris NTM Northern Territories Museum, Darwin
Page 1 and 2: Smithsonian Institution Scholarly P
Page 3 and 4: smithsonian contributions to zoolog
Page 5: Contents INTRODUCTION 1 A Brief His
Page 9 and 10: NMNH National Museum of Natural His
Page 11 and 12: nifi cant morphological and histori
Page 13 and 14: number 629 7 TABLE 2. Characters an
Page 15 and 16: TABLE 3. Character matrix used in p
Page 17 and 18: 78 90 Grade A 95 53 84 Clade B 99 1
Page 19 and 20: the genera, even though some genera
Page 21 and 22: nate to the branches, there is a te
Page 23 and 24: Kükenthal’s (1919) suggestion th
Page 25 and 26: smooth state transforms to granular
Page 27 and 28: number 629 21 a′′′. Branching
Page 29 and 30: 1. Primnoeides Studer and Wright in
Page 31 and 32: Discussion. Ainigmaptilon has such
Page 33 and 34: number 629 27 TABLE 4. List of the
Page 35 and 36: TABLE 4. (Continued) Genera and sub
Page 37 and 38: TABLE 4. (Continued) Genera and sub
Page 39 and 40: Discussion. Four species are known
Page 41 and 42: once again considered as a separate
Page 43 and 44: Discussion. Cyathogorgia is monotyp
Page 45 and 46: Although similar to the family Isid
Page 47 and 48: side; in some species adaxial and e
Page 49 and 50: Distribution. Same as for the genus
Page 51 and 52: two species facing downward. Well-d
Page 53 and 54: distinctive cruciform extinction pa
Page 55 and 56: FIGURES 3-19 FIGURE 3. Primnoeides
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FIGURE 5. Convexella magelhaenica (
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FIGURE 7. Thouarella (Diplocalyptra
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FIGURE 9. Fannyella (Cyathogorgia)
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FIGURE 11. Acanthoprimnoa goesi (a-
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FIGURE 13. Paranarella watlingi (a-
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FIGURE 15. Paracalyptrophora kerber
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FIGURE 17. Microprimnoa diabathra (
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FIGURE 19. Pseudoplumarella thetis
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68 SMITHSONIAN CONTRIBUTIONS TO ZOO
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REQUIREMENTS FOR SMITHSONIAN SERIES
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A Generic Revision and Phylogenetic Analysis of the Primnoidae

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