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A Revision of the Primnoidae 

(Octocorallia: Alcyonacea) 

from the Aleutian Islands 

and Bering Sea

Stephen D. Cairns

smithsonian contributions to zoology • number 634
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s m i t h s o n i a n c o n t r i b u t i o n s t o z o o l o g y • n u m b e r 6 3 4

A Revision of the Primnoidae

(Octocorallia: Alcyonacea)

from the Aleutian Islands

and Bering Sea

Stephen D. Cairns

washington d.c.

2011
ABSTRACT

Cairns, Stephen D. A Revision of the Primnoidae (Octocorallia: Alcyonacea) from the Aleutian 

Islands and Bering Sea. Smithsonian Contributions to Zoology, number 634, vi + 55 pages, 17 

figures, 3 tables, 2011.—Nineteen of the 31 octocoral species (61%) known from the Aleutian 

Islands belong to family Primnoidae, which form large deep-water (9–2,514 m) colonies providing 

habitat for one of the most productive fisheries in the North Pacific. These 19 species are described 

or redescribed and figured unless recent descriptions have been published. Eight new species are 

described, two in the genus Thouarella and six in the genus Plumarella. These two genera are 

redefined and differentiated by emphasizing the nature (keeled or not) of the inner side of their 

marginal scales and the inner side’s articulation or lack thereof with the outer surface of the underlying opercular scales. Dicholaphis is proposed as a new subgeneric rank of the genus Plumarella, 

characterized by having polyps arranged on all sides of its branchlets, not on alternate opposite 

sides. A key is provided for the eight species of Aleutian Plumarella, and all 35 valid species in the 

genus are listed, including three new combinations: P. superba, P. abietina, and P. recta. A history 

of the Aleutian primnoids is recounted. Six primnoid species are reported from the Bering Sea, all 

of which also occur in the Aleutian Islands. Two-thirds of the Aleutian–Bering Sea primnoids are 

endemic to this region.

Cover images, from left to right: Figures 12b, 14d, and 13h (detail from each).

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

P.O. Box 37012, MRC 957

Washington, D.C. 20013-7012

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Text and images in this publication may be protected by copyright and other restrictions or owned by individuals and entities other than, and in addition to, the Smithsonian Institution. Fair use of copyrighted material includes the use of protected materials for personal, educational, or noncommercial purposes. Users must 

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any other use.

Library of Congress Cataloging-in-Publication Data

Cairns, Stephen D. (Stephen Douglas), 1949–

A revision of the Primnoidae (Octocorallia:Acyonacea) from the Aleutian Islands and Bering Sea / 

Stephen D. Cairns.

 p. cm. — (Smithsonian contributions to zoology ; no. 634)

Includes bibliographical references and index.

1. Primnoidae—Alaska—Aleutian Islands—Classification. 2. Primnoidae—Bering Sea—Classification.

I. Title. 

QL377.C6C344 2011

593.6—dc22

2010045753

ISSN: 0081-0282 (print); 1943-6696 (online)

The paper used in this publication meets the minimum requirements of the American National Standard 

for Permanence of Paper for Printed Library Materials Z39.48–1992.
Contents

LIST OF Tables v

INTRODUCTION 1

Abbreviations 2

History of Primnoid Octocorals from the Aleutian Islands 3

Acknowledgments 4

MATERIAL AND METHODS 4

SYSTEMATIC ACCOUNTS 4

Subclass Octocorallia 4

Order Alcyonacea 4

Suborder Calcaxonia Grasshoff, 1999 4

Family Primnoidae Milne Edwards, 1857 4

Genus Thouarella Gray, 1870 4

 Thouarella cristata, new species 5

 Thouarella trilineata, new species 6

Genus Plumarella Gray, 1870 7

Subgenus Plumarella (Plumarella) Gray, 1870 7

 Plumarella (P.) spicata Nutting, 1912 7

Plumarella (Dicholaphis) Kinoshita, 1907, new subgenus rank 9

 Key to the Species of the Subgenus Plumarella (Dicholaphis) 10

 Plumarella (D.) profunda, new species 10

 Plumarella (D.) hapala, new species 11

 Plumarella (D.) aleutiana, new species 11

 Plumarella (D.) superba (Nutting, 1912), new combination 13

 Plumarella (D.) nuttingi, new species 14

 Plumarella (D.) echinata, new species 15

 Plumarella (D.) robusta, new species 16

Genus Fanellia Gray, 1870 16

 Fanellia compressa (Verrill, 1865) 17

 Fanellia fraseri (Hickson, 1915) 18
i v • smithsonian contributions to zoology

Genus Primnoa Lamouroux, 1812 19

 Primnoa pacifica Kinoshita, 1907 19

 Primnoa pacifica var. willeyi Hickson, 1915 19

 Primnoa wingi Cairns and Bayer, 2005 20

Genus Arthrogorgia Kükenthal, 1908 20

 Arthrogorgia utinomii Bayer, 1996 20

 Arthrogorgia kinoshitai Bayer, 1952 21

 Arthrogorgia otsukai Bayer, 1952 22

Genus Calyptrophora Gray, 1866 22

 Calyptrophora laevispinosa Cairns, 2007 23

Genus Parastenella Versluys, 1906 23

 Parastenella doederleini (Wright and Studer, 1889) 24

 Parastenella ramosa (Studer, 1894) 24

 Parastenella gymnogaster Cairns, 2007 25

FIGURES 1–17 27

APPENDIX: STATION LIST 45

REFERENCES 49

INDEX 53
Tables

1. Species of Alaskan Primnoidae 2

2. Species of Plumarella Gray, 1870 8

A.1. Stations list 46

A Revision of the Primnoidae 

(Octocorallia: Alcyonacea) from the 

Aleutian Islands and Bering Sea

INTRODUCTION

The Gulf of Alaska, particularly the Aleutian Islands, harbors one of the 

most productive fisheries in the world (Sanchez and Cairns, 2004; Stone and 

Shotwell, 2007). Their large size (some species up to 7 m; Krieger, 2001), longevity (some species over a hundred years; Andrews et al., 2002), and tendency 

to form high-density “coral gardens” make Aleutian deep-water octocorals important formers of habitat in deep water, providing vital refuges for fish and invertebrates (Heifetz, 2002; Krieger and Wing, 2002; Auster, 2005; Stone, 2006) 

but making them vulnerable to fishing impacts (Stone and Shotwell, 2007). Over 

the last decade, NOAA has sponsored research to determine the distribution, 

abundance, and species associations of these deep-water octocorals in an effort 

to assess their contribution to fisheries interactions and the need for conservation of these important ecosystems. However, the taxonomy of the Aleutian 

octocorals has not been revised since Nutting (1912), and most lists of species 

(e.g., Cimberg et al., 1981; Wing and Barnard, 2004; Heifetz et al., 2005; Stone 

and Shotwell, 2007) include a high percentage (almost half) of unidentified species as well as dated taxonomy. It is the purpose of this paper to taxonomically 

revise the Aleutian and Bering Sea primnoid species so that they can be discussed 

in a more meaningful way.

The Aleutian Islands certainly has the highest coral diversity in Alaska (Heifetz, 2002) relative to octocorals, stylasterids, and Scleractinia, but later claims of 

having “the highest abundance and diversity of coldwater corals in the world” 

(Heifetz et al., 2005:131; Stone, 2006) are overly exaggerated. In fact, using as a 

comparison the number of 31 as the total of valid, named nonpennatulacean octocoral species found in the Aleutian Islands (Stone and Shotwell, 2007; see below), 

there are 91 deep-water octocoral species known from the Hawaiian Islands 

(Cairns and Bayer, 2007), 89 species in Sagami Bay, Japan (A. K. Matsumoto 

Stephen D. Cairns, Department of Invertebrate 

Zoology, National Museum of Natural History, 

Smithsonian Institution, P.O. Box 37012, MRC 

163, Washington, D.C. 20013-7012, USA; 

cairnss@si.edu. Manuscript received 6 May 

2010; accepted 1 October 2010.
2 • smithsonian contributions to zoology

et al., unpublished poster [Bathymetric Distribution of the 

Deep-Water Octocorals (Cnidaria, Anthozoa) of SagamiBay, Tokyo, Japan] presented at Third International Symposium on Deep-Sea Corals, Science and Management, 

University of Miami, 2005), 214 from the larger Japanese 

region (Imahara, 1996), and 166 deep-water species in the 

northwest Atlantic (Cairns, 2006, 2007a). However, the 

Aleutian Islands are distinctive in having a relatively high 

percentage of their octocoral fauna composed of primnoid 

species (i.e., 61%). The primnoid component of the Hawaiian Islands deep-water octocorals is only 31%, of Sagami 

Bay (36%), of the Japanese region (19%), and of the northwest Atlantic (22%). Of the 18 Aleutian primnoid species, 

seven also occur in the Bering Sea (Table 1). Twelve of the 

19 (63%) are endemic to the Aleutian Islands–Bering Sea 

region (Table 1), whereas five species have broader distributions to the west (Okhotsk Sea, Sakhalin Island, Japan, and 

Indonesia), four have broader distributions to the east (two 

to the Alaskan coast, one to Panama, and one to California), and two are also found on Gulf of Alaska seamounts.

Abbreviations

The following abbreviations are used in the text.

Museums/Institution

AB Auke Bay Laboratory, Juneau, Alaska

BM The Natural History Museum, London

table 1. The twenty-six primnoid species and one subspecies known from Alaska, including regions in which they have been collected 

(indicated with an x) and known depth range. A dash (–) indicates species is not known from that region; AI = Aleutian Islands.

West of Gulf of Alaska East of Bering Depth range

Alaskan primnoid Aleutians AI seamounts Aleutians Sea a (m)

Thouarella cristata, n. sp – x – – x 94–768

T. trilineata, n. sp. – x – – – 97–642

Plumarella (Plumarella) spicata Nutting, 1912 – x – – x 712–1,912

P. (Dicholaphis) profunda, n. sp. – x – – – 2,514

P. (D.) hapala, n. sp. – x – – – 120–384

P. (D.) aleutiana, n. sp. – x – – x 79–517

P. (D.) superba (Nutting, 1912), x x – – x 29–888

n. comb. (1,258) b

P. (D.) nuttingi, n. sp. – x – – – 492–888

P. (D.) echinata, n. sp. – x – – x 150–1,692

P. (D.) robusta, n. sp. – x – – – 712–1,061

Fanellia compressa (Verrill, 1865) – x – – – 82–1,061

F. fraseri (Hickson, 1915) – x – x – 86–1,341

Primnoa pacifica Kinoshita, 1907 x x – x x 9–800

P. pacifica var. willeyi Hickson, 1915 – x x x – 27–863

P. wingi Cairns and Bayer, 2005 – x – x x 110–914

Narella bayeri Cairns and Baco, 2007 – – x – – 3,277–4,091

N. arbuscula Cairns and Baco, 2007 – – x – – 2,775–3,465

N. cristata Cairns and Baco, 2007 – – x – – 3,385

N. abyssalis Cairns and Baco, 2007 – – x – – 4,594

N. alaskensis Cairns and Baco, 2007 – – x – – 2,377–3,075

Arthrogorgia utinomii Bayer, 1996 – x – – – 163–882

A. kinoshitai Bayer, 1952 x x – – – 220–1,309

A. otsukai Bayer, 1952 – x – – – 1,359–1,372

Calyptrophora laevispinosa Cairns, 2007b – – x x – 2,672–3,531

Parastenella doederleini (Wright and Studer, 1889) x x – – – 400–3,423

P. ramosa (Studer, 1894) x x x x – 665–1,766

P. gymnogaster Cairns, 2007b – – x x – 1,962–2,869

a Bering Sea (Bowers Bank, Pribilof Canyon, Zhemchug Canyon).

bThe depth of 1,258 m is for the pinnate growth form, which may be a station error (see text); 29–888 m is typical range.
n u m b e r 6 3 4 • 3

CAS California Academy of Sciences, San Francisco

NMNH National Museum of Natural History, 

Smithsonian Institution, Washington, D.C.

NOAA National Oceanic and Atmospheric Administration

USNM United States National Museum (now the 

NMNH) 

ZMA Zöologisch Museum, Amsterdam

Vessels

Alb USFWS Albatross

DW Deep Worker (submersible)

MF R/V Miller Freeman

Other Terms

GOA Gulf of Alaska

L:W ratio of length to width of a polyp or coenenchymal scale

SEM scanning electron microscope

History of Primnoid Octocorals

from the Aleutian Islands

The first primnoid reported from the Aleutian Islands, 

Prymnoa verticillaris by Ehrenberg (1834), is by inference 

only. Ehrenberg’s account does not list a collecting locality, but Studer (1878), who examined and illustrated this 

specimen from the Chamisso collection in the Berlin Museum, stated that it was collected on the Rurik Expedition (1816–1817), which made collections in the Bering 

Sea and eastern Aleutian Islands. According to Studer, this 

species was the same as Verrill’s (1865) Primnoa compressa, which was described from the Aleutian Islands and 

collected on the U.S. North Pacific Exploring Expedition 

but without precise locality or depth. Primnoa compressa

has now been transferred to the genus Fanellia. According 

to Bayer (1982), Ehrenberg’s specimens might have been 

either of the two Fanellia known from this region but 

more likely F. fraseri. It was not until 1915 that Hickson 

described Caligorgia fraseri (=Fanellia fraseri) from Albatross Bank in the Gulf of Alaska.

In his report on the octocorals collected by the Albatross in Japanese waters, Nutting (1912) also included 

nine extralimital records from the Aleutian Islands and 

Bering Sea, including three species of Plumarella, two of 

Thouarella, and a new genus and species, Primnodendron 

superbum (transferred to Plumarella herein). All of these 

records are discussed in the text below, but some samples 

of the specimens were too small or damaged to identify, 

such as P. flabellata and T. hilgendorfi. Much later Bayer 

(1952) also reported two new species of Arthrogorgia

from the same Albatross expedition of 1906, both from 

the Aleutian Islands.

In an infrequently cited “final report,” Cimberg et 

al. (1981) listed all primnoids previously reported from 

Alaska and discussed some of the ecological factors (e.g., 

depth, substrate, temperature) that affect their distribution, but their taxonomy is uncritical and the publication is 

more similar to an informal agency report, and thus these 

references are not included in the species synonymies.

Bayer’s (1982) second contribution to this topic was 

a redescription of the two species of Fanellia known from 

the Aleutian Islands, accompanied by detailed SEM illustrations and several new records for each species. He 

(Bayer and Stefani, 1989) later included these two species 

in a key to the species of Fanellia. Similar to his contribution of Fanellia, Bayer (1996) described three species 

(including one new) of Arthrogorgia from the Aleutian 

Islands, including copious SEM illustrations, several new 

records, and a key to the species in the genus. 

In 2002, Heifetz listed and indicated coarse geographic 

abundance of the common primnoid octocorals collected by 

NOAA vessels from 1975 through 1998, showing a correlation to fish that were collected at the same stations. Many 

of the primnoids were not identified to species, and unfortunately voucher specimens were not indicated. Later Heifetz 

et al. (2005) amplified this discussion, listing 52 nonpennatulacean octocorals from Alaskan waters, 21 of which 

were primnoids. They also pointed out a zoogeographic 

and physical boundary at Samalga Pass (between Umnak 

and Chuginadak Islands) and that coral diversity was much 

higher to the west of this pass, an observation supported by 

the distributions of the primnoid species reported herein. 

Many of the specimens they used for their observations 

were collected by the Delta and Jason II, some of which 

are included in this study. And, although not taxonomic 

in nature, Stone (2006) studied the depth distribution of 

western Aleutian cold-water corals (including octocorals) 

and their fine scale associations with economically important fish species. Most of these specimens were collected by 

the Delta submersible, many of which are reported herein.

Wing and Barnard’s (2004) field guide to the Alaskan 

corals listed 55 nonpennatulacean octocorals, 21 of which 

(38%) belonged to the family Primnoidae. They keyed all 

species and illustrated several, but, in general, their taxonomy was uncritical and voucher specimens were not 

designated.

In a revision of the genus Primnoa, Cairns and Bayer 

(2005) reported additional records of Primnoa pacifica
4 • smithsonian contributions to zoology

and the new species P. wingi from the Aleutian Islands. 

And, although not from the Aleutian Islands, Cairns 

(2007b) reported Parastenella ramosa and P. gymnogaster

from seamounts in the adjacent Gulf of Alaska. Likewise, 

Cairns and Baco (2007) reported five species of Narella

from deep seamounts in the Gulf of Alaska.

Finally, Stone and Shotwell (2007) provided a very 

useful status of the deep coral ecosystems of Alaska, and 

in their Appendix 2.1 list 68 nonpennatulacean octocorals 

from the Aleutian Islands, 21 of which were primnoids, 

however, almost half (31 taxa) of their listed taxa were 

undescribed, and their taxonomy was uncritical.

In summary, if one does not count unidentified species, only 14 (9 valid) primnoid species had been previously reported from the Aleutian Islands out of a total of 

27 (22 valid) nonpennatulacean species (based on list of 

Stone and Shotwell, 2007). The addition of eight new species and a new record (Parastenella doederleini) increase 

the total number of valid Aleutian species to 31 and the 

number of primnoids to 19 species, or 61% (19/31 species) of the nonpennatulacean octocorals. Another eight 

primnoid species occur in the Gulf of Alaska, bringing the 

total of Alaskan primnoids to 26 (Table 1). 

Acknowledgments

I am very grateful to Bob Stone (Auke Bay Laboratory, 

Juneau, Alaska) for the gift to the Smithsonian of a large 

percentage of the specimens used in this study and for his 

provision of metadata for those specimens. I also thank 

Amy Baco (Florida State University) for providing the 

Alvin specimens collected from seamounts in the Gulf of 

Alaska; these specimens were obtained by two NOAA-OE 

cruises: UAF 040118 and NA 04OAR4600051. I am, 

as always, grateful to Tim Coffer (Smithsonian) for his 

meticulous Photoshop work in composing the figures. Finally, I thank Michele Taylor (Imperial College London) 

and Rebeca Zapata (University of Seville) for their many 

discussions of the vexing relationships between and within 

Thouarella and Plumarella.

MATERIAL AND METHODS

Specimens were examined from over 124 stations (see 

Appendix) and logged by 17 vessels, most of which were 

government research vessels and submersibles sponsored 

by NOAA, covering the entire length of the Aleutian Islands at depths of 40–2,514 m. Most of the specimens 

are deposited at the NMNH, but in some cases, especially 

specimens from the Jason IV and Delta, only a small fragment is on deposit at the NMNH, with the main colony 

residing at Auke Bay Laboratory, Juneau, Alaska. Types of 

21 of the 26 Aleutian species are deposited at the NMNH. 

Most of the SEM images were taken by the author (stub 

number prefaced with C), but 19 images were used from 

the Bayer archive (stub number prefaced with a B). In the 

material examined sections, specimens are presumed to 

be preserved in 70% ethanol unless otherwise noted. A 

“confirmed” depth range is reported for each species (i.e., 

the depths of the shallowest deep to the deepest shallow 

ranges of individual stations).

SYSTEMATIC ACCOUNTS

Subclass Octocorallia

Order Alcyonacea

Suborder Calcaxonia Grasshoff, 1999

Family Primnoidae Milne Edwards, 1857

Genus Thouarella Gray, 1870

Thouarella Gray, 1870:45.—Kükenthal, 1919:405–408.—Bayer, 1956: 

F220.—Cairns, 2006:175–176.—Taylor et al., in press.

Rhopalonema Roule, 1908:2–3.

Thouarella (Diplocalyptra) Kinoshita, 1908a:454.—Cairns and Bayer, 2009: 

34–35.

Thouarella (Euthouarella) Kükenthal, 1915:149.—Cairns and Bayer, 2009: 

34.

Thouarella (Parathouarella) Kükenthal, 1915:150.

Thouarella (Epithouarella) Kükenthal, 1915:150–151.—Cairns and Bayer, 

2009:35.

Thouarella (Thouarella).—Bayer, 1956:F220.—Cairns and Bayer, 2009:33–

34.

Diagnosis. Colonies bottlebrush, uniplanar 

pinnate, or dichotomous in branching. Polyps isolated, 

that is, occurring on all sides of branchlets (species Group 

1 sensu Taylor et al., in press) or arranged in pairs or 

whorls on branchlets (species Group 2 sensu Taylor et al., 

in press), pointing upward. Polyps covered by 6 or 8 longitudinal rows of body wall scales, the scales of adaxial row 

often reduced in size and number to accommodate inward 

flexing of polyp. Marginal scales often arranged in 2 circlets of 4 scales that alternate with one another, the circumference of the distal polyp not being large enough to 
n u m b e r 6 3 4 • 5

accommodate 8 adjacent marginal scales. Each marginal 

bears a simple or ornate longitudinal keel or multiple keels 

that, when the marginals fold over the operculum, glide 

along outer concave surface of underlying opercular scales 

in a tongue-and-groove fashion.

Discussion. The various subgenera of 

Thouarella were previously distinguished (see synonym 

above, especially Kükenthal, 1915, 1919, 1924; Bayer, 

1956; Cairns, 2006; Cairns and Bayer, 2009) by the relative length of their marginal spines, and the grouping or 

nongrouping of polyps on the branchlets. However, Taylor et al. (in press) showed that the relative length of the 

marginal spine was a variable and continuous (nondiscrete) character. This led them to abandon that system 

of subgeneric differentiation in preference to dividing 

the 24 Thouarella species into two species groups, the 

groups based solely on the arrangement of polyps on the 

branchlets, a much easier character to interpret. Further 

differentiation of species within these groups is based on 

branching mode and characteristics of the polyp sclerites. 

The two new species of Thouarella described herein both 

belong to Group 1, that is, species having a random or 

isolated arrangement of polyps, which currently contains 

18 of the 26 known species in this genus.

The specimen reported as T. hilgendorfi by Nutting 

(1912:66), Heifetz (2002), Wing and Barnard (2004:24), 

Heifetz et al. (2005), and Stone and Shotwell (2007) as 

questionably from the Bering Sea (Alb-4771: USNM 

30133) could not be confidently identified. Likewise, 

Thouarella sp. A listed by Heifetz (2002) and Stone and 

Shotwell (2007) is uncertain.

Distribution. Worldwide, from Antarctic 

to the Aleutian Islands, 73–2,100 m (Cairns and Bayer, 

2009; Taylor et al., in press).

Thouarella cristata, new species

Figures 1a, 3a–i

?Plumarella sp.—Wing and Barnard, 2004:24, 42, fig. 12.

Description. Colonies uniplanar, consisting 

of relatively few main branches, from which branchlets 

originate in an irregularly pinnate fashion also in plane of 

colony; however, associated with the presence of a commensal polychaete, which occasionally lives along one of 

the main branches, a third set of branchlets develop, causing the branches to appear somewhat bushy or bottlebrush 

in shape (Figure 1a). Branchlets up to 8 cm in length and 

not further subdivided, usually regularly arranged parallel 

to one another. Largest colony (USNM 1010273) 50 cm 

in height, with a basal branch axis diameter of 11 mm. 

Holotype 32 cm tall and 16 cm wide, consisting of two 

main branches but lacking holdfast. Axis pale yellow to 

brown-black in color.

Polyps occurring on main stems and on all sides of 

branchlets in a crowded manner (Figure 3a), resulting in 

24–26 polyps/cm, even more toward tips of branchlets. 

Polyps 1.7–2.5 mm in length and flared distally as well as 

curving upward along branchlet axis. Brooding polyps (or 

those with large eggs) common; these polyps cylindrical or 

basally swollen.

Polyps covered with 8 longitudinal rows of body wall 

scales: 6-9 scales in each abaxial row, 6-7 in each outer 

lateral row, 4-6 in each inner lateral row, and 3-4 in 2 

adaxial rows, the proximal part of adaxial side of polyp 

being further covered with small elliptical scales 0.21–0.42 

mm in diameter (Figure 3c, i). Opercular scales triangular, 0.47–0.69 mm in length (L:W = 2.0–2.45), forming a 

prominent, peaked operculum easily visible in lateral view 

(Figure 3b,e). Outer surface of operculars prominently 

ridged, the finely serrate ridges radiating from a central 

basal point; proximal inner surface tuberculate, but distal inner surface bearing prominent multiple longitudinal 

ridges; edge of operculars finely serrate. Marginal scales 

(Figure 3f) triangular and larger than operculars (0.53–

0.73 mm in length) but with a smaller L:W (1.3–1.9), thus 

closer to an equilateral triangle. Outer surface of marginals prominently ridged, as the operculars, but distal inner 

surface bearing one or more prominent keels, easily visible in an opercular polyp view (Figure 3c, d); marginal 

scales flaring outward from polyp. The 2 adaxial marginals smaller than the other 6. Submarginal scales similar 

in shape to marginals, often flaring outward and having 

small keels, but shorter and with lower L:W ratio (0.9–

1.5). Remaining body wall scales (Figure 3g) progressively 

shorter (L:W around 1) and less ornamented (smooth exteriorly) toward base of polyp. All body wall scales thick 

and robust.

Coenenchymal scales (Figure 3h) irregular in shape 

and flat, with a granular outer surface and a tuberculate 

inner surface; scales up to 0.9 mm in length.

Remarks. According to the conventional 

organization of the genus (Cairns and Bayer, 2009), 

Thouarella cristata would fall within the nominate subgenus (i.e., those species having pointed marginal scales 

and isolated polyps) or according to the revised classification of the genus (Taylor et al., in press) as within species 

group 1 (i.e., those species having isolated polyps). Regardless, T. cristata differs from all previously described 
6 • smithsonian contributions to zoology

species in having thick and highly ridged body wall and 

opercular scales, a relatively high number of polyps per 

centimeter, and a tendency to form quasi-bottlebrush colonies (i.e., main branches often having three distinct rows 

of branchlets).

Types and Type locality. Holotype: Delta

6004-11C-6, 1 dry colony, 1 alcohol fragment, SEM stubs 

C1440-1443, USNM 1115600. Paratypes: 51°26.55′N, 

179°21.62′E, 768 m, 7 Apr 2000, 1 dry colony, USNM 

1006315 (exAB01-54); Jason II 2103-14-2, 1 fragment, 

USNM 1134916, and 1 dry colony, AB01-54 and AB10-

0090; Sea Storm 116, 1 large dry colony and small colony 

in alcohol, USNM 1010273. Type Locality: 52°45′55′′N, 

179°19′42′′W (southern Bowers Bank), 154 m.

Material Examined. Types.

Distribution. Aleutian Islands: off Rat Islands: Kiska, Amchitka, and Semisopochnoi Islands, and 

southern Bowers Bank, 94–768 m.

Etymology. Named cristata (Latin for 

crested or ridged), in allusion to the prominently ridged 

outer surfaces of the body wall scales and the ornate keels 

on the inner sides of the marginal scales.

Thouarella trilineata, new species

Figures 1b, 4a–j

Description. Colonies uniplanar, consisting 

of relatively few main branches, from which branchlets 

originate usually in three rows giving the main branches a 

bushy (bottlebrush) appearance (Figure 1b). Branchlets up 

to 6 cm in length and not further subdivided, arranged in 

an alternating pinnate fashion on main branches. Largest 

colony (the holotype) 23 cm in height and 14 cm wide, 

consisting of only 2 main branches; basal branch axis 

4.5 mm in diameter. Axis pale yellow.

Polyps occurring on main stems and on all sides of 

branchlets in a crowded manner (Figure 4a), resulting in 

35–40 polyps/cm. Polyps 1.4–1.7 mm in length and are 

cylindrical to slightly flared distally, curving upward along 

branch axis. Modified brooding polyps not apparent, perhaps because eggs are fairly small.

Polyps covered with 6-8 longitudinal rows of body 

wall scales: 5–6 scales in the 2 abaxial rows, 4–6 in 2 

outer lateral rows, 3–5 in 2 inner lateral rows, and only 

6–10 randomly placed sclerites on adaxial side below adaxial marginals, hardly deserving of the name of row and 

not covering entire adaxial side (Figure 4b, c). Opercular 

scales triangular (Figure 4f), 0.35–0.52 mm in length (L:W 

= 2.0–2.5), and somewhat distally attenuate, such that the 

8 operculars, when in closed position, do not form an entire operculum, allowing some space between opercular 

scales; adaxial operculars smaller than others (Figure 4c). 

Opercular scales curved inward, resulting in a low operculum (Figure 4b). Outer surface of operculars highly ridged, 

the finely serrate ridges originating from a central basal 

point; proximal inner surface tuberculate whereas distal 

inner surface bears multiple finely serrate ridges. Marginal 

scales triangular (pointed) to spinose (Figure 4g), except 

for the 2 adaxial marginals, which are rounded distally; 

otherwise, marginals about same height as operculars 

(0.42–0.55 mm) but have a broader base and thus a lower 

L:W of 1.0–1.5. Like opercular scales and all body wall 

scales, the outer surface of a marginal is highly ridged with 

deep valleys between the ridges; the distal inner surface 

bears multiple serrate keels. Submarginal scales (Figure 

4h) similar to marginals but slightly less tall and thus having a smaller L:W of 1.0–1.25, and usually lack distinct 

keels. Remaining body scales (Figure 4i) progressively 

smaller toward base of polyp; those in lateral rows somewhat asymmetrical but all having tall ridges on their outer 

surface. Adaxial scales below marginals small (0.14–0.20 

mm in diameter) and elliptical, not formed into rows, and 

exposing some of the adaxial side of polyp. All body wall 

scales rather thick, especially along longitudinal axis.

Coenenchymal scales (Figure 4j) irregular in shape 

and flat, with a granular outer surface and tuberculate 

inner surface; scales up to 0.60 mm in length.

Remarks. Small ophiuroids (disc diameter = 

5–7 mm), the same color as the coenenchyme, are common commensals on this species.

Although similar in many ways to Thouarella cristata, 

T. trilineata differs in having smaller polyps, nonspinose 

adaxial marginals, reduced adaxial marginals such that 

part of the adaxial polyp face is bare, even more highly 

ridged body wall scales that extend to the base of polyp, 

and a less flared polyp shape.

Types and Type locality. Holotype: Dominator 1997-01-39, 1 large colony in alcohol, SEM stubs 

C1425-1429, USNM 1010175. Paratypes: Alb-3330, 1 

colony, USNM 49342; Alb-3331, 1 colony, USNM 49342; 

Delta 6199-5E-5, 1 branch, USNM 1135540, and 1 dry 

colony, AB10-0033; Delta 5993-7C-5, 1 dry fragment, 

USNM 1135538; Dominator 1997-01-49, 1 dry and 1 alcohol preserved colony, USNM 1009952 and 1009949, 

respectively; Jason II 2103-7-1A, fragment, USNM 

1135539; Let’s Go 861-50, 1 branch, USNM 1135541; 

Pacific Knight 941-38, 1 colony, USNM 100798; Sea 

Storm 148, 1 colony, USNM 1009943; Starlight 84-1-7, 

1 colony, USNM 100753; Vesteraalen 941-36, 2 colonies, 
n u m b e r 6 3 4 • 7

USNM 100754 and 100746. Type Locality: 53°01′11′′N, 

170°18′49′′W (northwest of Carlisle Island, Islands of 

Four Mountains), 202 m.

Material Examined. Types.

Distribution. Aleutian Islands from Amchitka Pass to Unalaska, 97–642 m.

Etymology. Named trilineata (Latin for 

“three lines”) for the trilinear arrangement of the branchlets from the main branches.

Genus Plumarella Gray, 1870

Plumarella Gray, 1870:36.—Cairns and Bayer, 2009:39 (revision).

Diagnosis. Colonies usually uniplanar and 

alternately pinnately branched but may also be dichotomously branched and bottlebrush in shape. Polyps arranged 

in alternate biserial fashion on branchlets or crowded on 

all sides of branchlets in no order. Polyps covered by 8 

longitudinal rows of body wall scales, the adaxial rows 

sometimes reduced in number and size. Marginal scales arranged in one circlet of scales, which do not fold over the 

operculum; marginals variable in shape, their distal edges 

being spinose, pointed, or straight; inner surface of marginals may be smooth, ridged, or spiny but never keeled.

Discussion. The 35 species currently assigned to 

genus are listed in Table 2. The genus was recently revised 

by Cairns and Bayer (2009).

The dry specimen of P. flabellata reported by Nutting 

(1912:63), Heifetz (2002), Wing and Barnard (2004:24), 

Heifetz et al. (2005), and Stone and Shotwell (2007) based 

on specimens from Alb-4784 (Attu Island, Near Islands, 

Aleutians), USNM 91438, was too damaged to properly 

identify.

The Alaskan listing of P. longispina by Wing and Barnard (2004), Heifetz et al. (2005), and Stone and Shotwell 

(2007:107) could not be confirmed.

The species listed as Amphilaphis A, B, and C by 

Heifetz (2002), Heifetz et al. (2005), and Stone and Shotwell (2007:106) may be the species of Plumarella (or 

Thouarella) discussed below. Similarly, the Alaskan listing 

of Plumarella sp. 1 or A by Heifetz (2002) and Stone and 

Shotwell (2007), respectively, cannot be confirmed.

Distribution. Indo-Pacific, western Atlantic, Subantarctic; 10–3,182 m. 

Subgenus Plumarella (Plumarella) Gray, 1870

Diagnosis. Plumarella in which polyps occur 

on alternating sides of branchlets.

Discussion. See discussion of subgenus 

Dicholaphis.

Distribution. As for genus.

Plumarella (P.) spicata Nutting, 1912

Figures 1f, 5a–h

Plumarella spicata Nutting, 1912:64–65, pl. 8: figs. 2, 2a, pl. 18: fig. 6. 

—Kükenthal, 1919:350; 1924:261.—Wing and Barnard, 2004:24 

(listed).—Heifetz et al., 2005:132 (listed).—Stone and Shotwell, 

2007:107 (listed).—Cairns and Bayer, 2009:29 (listed).

Plumarella spinosa.—Nutting, 1912:63 (in part: Alb-4781).—Heifetz, 

2002:26 (listed).—Wing and Barnard, 2004:24 (listed).—Heifetz et al., 

2005:132 (listed).—Stone and Shotwell, 2007:107 (listed).

Description. Colonies uniplanar, consisting 

of numerous branches arranged in a quasi-dichotomous 

manner, from which branchlets diverge in a quasidichotomous to loosely alternate pinnate fashion (Figure 

1f), branchlets on same side of a branch being 8–15 mm 

apart. Colonies delicate and flimsy, quite flexible; adjacent 

branchlets not held rigidly parallel to one another. Largest 

colony (holotype fragment) 14 cm tall and 7 cm in width, 

with a basal branch diameter of only 1.2 mm. Branch axis 

straw yellow in color.

Polyps occurring on main branches and branchlets 

in an alternating biserial arrangement in plane of colony 

flabellum, although occasional polyp diverging out of the 

flabellar plane; polyps sparse, about 12–14/cm branchlet 

length. Polyps of moderate size, 1.4–1.6 mm in length, 

slightly flared distally; polyps oriented perpendicular to 

branchlets, although distal ends curved slightly upward.

Polyps covered with 8 longitudinal rows of body wall 

scales: 5 or 6 scales in each abaxial row, 5–6 in outer lateral rows, 4–6 in inner lateral rows, and 4–6 small scales in 

reduced adaxial rows (Figure 5c). Opercular scales (Figure 

5e) isosceles triangular in shape, elongate, and pointed, 

0.32–0.70 mm in length (L:W = 2.7–3.7), the adaxial 

operculars being shortest and often having rounded tips. 

Outer surface of operculars smooth to slightly granular 

proximally; inner surface tuberculate proximally, smooth 

medially, and ridged distally. All 8 marginals (Figure 5c, d, 

f) prominently spinose, consisting of a rectangular (wider 

than tall) to crescent-shaped basal part that is clearly demarked from the distal spine, the spine approaching 70% 

length of scale. Marginal scales up to 0.80 mm in length, 

the adaxial marginals being smaller, and have an L:W 

of 2.1–2.8. Outer surface of basal portion of marginals 

smooth to granular, inner surface tuberculate; distal spine 
8 • smithsonian contributions to zoology

TABLE 2. List of the 35 species of Plumarella Gray, 1870, arranged by subgenus and indicating location data. A dash (–) indicates 

species not found.

Species Distribution Depth (m)

Subgenus Plumarella (Plumarella)

P. penna (Lamarck, 1815) (type species) Australia 10–60+

P. laevis Thomson and Mackinnon, 1911 Australia 90–115

P. dentata Thomson and Russell, 1910 Providence, Indian Ocean 229

P. flabellata Versluys, 1906 Japan unknown

P. spinosa typica Kinoshita, 1907 Japan 237–330

P. s. brevispina Kükenthal, 1919 Japan 50

P. alba Kinoshita, 1908 Japan 550

P. gracilis Kinoshita, 1908 Japan unknown

P. longispina Kinoshita, 1908 Japan 604

P. acuminata Kinoshita, 1908 Japan 330

P. dofleini Kükenthal and Gorzawsky, 1908 Japan 80–250

P. d. var. boninensis Aurivillius, 1931 Bonin Island 128

P. lata Kükenthal and Gorzawsky, 1908 Japan 200–300

P. rigida Kükenthal and Gorzawsky, 1908 Japan 180–600

P. adhaerans Nutting, 1912 Japan 180

P. recta (Nutting, 1912), n. comb. Japan 869–924

=P. alternata (Nutting, 1912) n. comb. Japan 934

=P. attenuata Kükenthal, 1924:301 (misspelling of alternata) – –

P. spicata Nutting, 1912 Aleutian Islands 712–1,912

P. circumoperculum Cairns, 2010 Hawaiian Islands 432–1,373

P. abietina (Studer, 1894), n. comb. Off Ecuador 3,182

P. delicatissima Wright and Studer, 1889 Patagonia 256

P. undulata (Zapata-Guardiola and López-González, 2010) South Georgia 306–343

P. bayeri (Zapata-Guardiola and López-González, 2010) South Georgia 306–343

P. diadema (Cairns, 2006) Brazil, South Georgia 278–1,000

=T. sardana Zapata-Guardiola and López-González, 2010 – –

P. pourtalesii (Verrill, 1883) Western Atlantic 231–950

P. p. forma robusta Deichmann, 1936 – –

P. p. forma obtusa Cairns and Bayer, 2004 – –

P. pellucida Cairns and Bayer, 2004 North Carolina 549–1,160

P. laxiramosa Cairns and Bayer, 2004 North Carolina 348–625

P. dichotoma Cairns and Bayer, 2004 South Carolina to Florida 494–1,065

P. aurea (Deichmann, 1936) Florida 574

P. aculeata Cairns and Bayer, 2004 Florida, Bahamas 400–900

Subgenus Plumarella (Dicholaphis)

P. delicata (Kinoshita, 1907) Japan 731

P. profunda, new species Aleutian Islands 2,514

P. hapala, new species Aleutian Islands 120–384

P. aleutiana, new species Aleutian Islands 79–517

P. superba (Nutting, 1912), n. comb. Aleutian Islands 40–1,258

P. nuttingi, new species Aleutian Islands 492–888

P. echinata, new species Aleutian Islands 150–779

P. robusta, new species Aleutian Islands 712–1,746
n u m b e r 6 3 4 • 9

circular in cross section and covered with small spines that 

are often aligned in discontinuous rows (not keeled, Figure 

5f). Abaxial submarginals often highly spinose like marginals but otherwise remaining body wall scales (Figure 

5g) elliptical to crescent shaped with a rounded, pectinate 

distal edge, only 0.18–0.29 mm in height, and having L:W 

ratios of 0.66–1.3. Their inner surface is tuberculate, but 

their outer surface is spiny with short ridges at their distal 

edges that project as short points. Distal edges of operculars and marginals finely serrate, whereas edges of body 

wall and coenenchymal scales pectinate. Tentacular rodlets absent.

Coenenchymal scales (Figure 5h) small (0.10–0.19 

mm diameter) elliptical scales, coarsely granular above 

and tuberculate below.

Remarks. Among the 27 species in the nominate subgenus (Table 2), only four have dichotomous 

branching: P. spicata, P. bayeri (Zapata-Guardiola and 

López-González, 2010), P. dichotoma Cairns and Bayer, 

2004, and P. aurea (Deichmann, 1936). Plumarella spicata differs from the other three in having spinose marginal scales that are covered with small spines. Plumarella 

spicata is also the only Pacific dichotomously branching 

species and the deepest dwelling of all in the subgenus.

Similar to Acanthoprimnoa in having coarsely granular coenenchymals and pectinate scale edges, P. spicata

differs in having tubercles on its inner scale surfaces.

Type and Type Locality. The holotype, preserved in alcohol, is now broken into at least 15 fragments, 

also SEM stubs B720 and C1437-1439, USNM 30050. 

Type locality: Alb-4780, 52°01′N, 174°39′E (Passage between Near Islands and Rat Islands, Aleutians), 1912 m.

Material Examined. Type; Alb-4771, colony, USNM 80926 (reported by Nutting, 1912); Alb-4781, 

1 branch, USNM 30058 (reported by Nutting, 1912, as 

P. spinosa); Jason II 2100-3-1, 1 dry colony, AB10-0075; 

Jason II 2100-4-8, 1 colony, USNM 1134070; Jason II

2101-7-9, 1 colony, USNM 1134071; Jason II 2104-5-2, 

1 dry branch, USNM 1134073, and 3 dry colonies, AB10-

0091; Jason II 2104-5-3, 1 branch, USNM 1134072. 

Distribution. Aleutian Islands from Near Islands to Adak Canyon, including Bowers Bank, 712–1912 m.

Plumarella (Dicholaphis) Kinoshita, 1907,

new subgenus rank

Dicholaphis Kinoshita, 1907:230-231; 1908b:24-27.—Cairns and Bayer, 

2009:26.

Primnodendron Nutting, 1912:70.

Diagnosis. Plumarella in which polyps occur 

on all sides of branchlets.

Discussion. In our revision of the primnoid 

genera (Cairns and Bayer, 2009), Dicholaphis was kept as 

a separate genus, distinguished from Plumarella because 

of the isolated nature of its polyps, not arranged in alternate biserial fashion as in all other Plumarella species (see 

Cairns and Bayer, 2009:21, key). The discovery of a clade 

of seven Aleutian species having a similar polyp placement 

but being otherwise similar to Plumarella, has caused a 

reevaluation of the genus, leading to incorporating Dicholaphis as a subgenus of Plumarella. The primary difference of this subgenus is that the polyps are arranged on 

all sides of the branchlets, not in alternating biserial fashion. Nutting (1912:70) also established a new genus for 

this primnoid ground plan, Primnodendron (type species, 

P. superbum), which he differentiated from Plumarella

by its “bearing secondary branching from the terminal 

twigs,” but P. superbum fits nicely within the Dicholaphis

subgenus.

The placement of Primnodendron as a junior synonym 

of Thouarella by Cairns and Bayer (2009) was a mistake. 

In retrospect, the genera Plumarella and Thouarella are 

much more difficult to distinguish than one might expect, 

despite the seemingly distinctive characters used by Cairns 

and Bayer (2009:20–21). Indeed, it is often difficult to determine if the marginal scales fold over the base of the 

opercular scales or not, the primary differentiating character used by Cairns and Bayer (2009) to distinguish the two 

genera. A more reliable character to differentiate the two 

genera is the nature of the marginal scales. In Thouarella

the inner surface of the marginal scales are keeled, the 

keels gliding along the outer concave surface of the underlying opercular scales in a tongue-and-groove fashion, 

allowing the marginals to fold and slide over the opercular scales. In Plumarella, the inner marginal surfaces are 

smooth, spiny, or lightly ridged (never keeled) and do not 

fold over or in any way articulate with the outer surface 

of the opercular scales. Thus the marginals of Thouarella

fold and those of Plumarella are fixed. Another difference 

is that in Thouarella the eight marginals tend to form two 

quartets of four scales, whereas the eight marginals of Plumarella are usually in one circlet. This redefining of the 

two genera has led to the reassignment of several species 

in both genera (see Table 2).

The species in this subgenus are arranged herein in 

order of increasing length of their marginal scales.

Distribution. Aleutian Islands and off 

Japan, 40–2514 m.
1 0 • smithsonian contributions to zoology

Plumarella (D.) profunda, new species

Figures 1e, 6a–f

Description. Colonies uniplanar, consisting 

of relatively few main branches from which branchlets 

originate in a loose, alternating, pinnate manner (Figure 

1e), the distance between adjacent branchlets on one side 

of a branch 6–12 mm. Branchlets up to 9 cm in length, 

the entire colony being fairly flexible. Holotype 18 cm in 

height and 11 cm in width, with a basal axis diameter of 

1.8 mm. Branch axis and polyps pale yellow.

Polyps occurring on all sides of the branches and 

branchlets; however, occasional branchlets in an alternate 

biserial arrangement. Polyps 1.5–1.9 mm in length, standing perpendicular to branchlets but often slightly turned 

upward at their tip; polyps widely spaced, only 10–14/cm 

on a branchlet.

Polyps covered with 8 longitudinal rows of body wall 

scales: about 5 in each abaxial and outer lateral row, 4 

in inner lateral rows, and a variable number of smaller 

scales in the reduced adaxial rows. Opercular scales (Figure 6c) isosceles triangular in shape, brittle, and pointed: 

0.47–0.86 mm in length (L:W = 2.5–4.1). Outer opercular surface smooth to mildly granular and longitudinally 

concave near tip; inner surface tuberculate proximally and 

covered with low longitudinal ridges distally. There is little difference between marginal and remaining body wall 

scales, all being roughly rectangular, with a straight (not 

pointed or spinose), slightly serrate distal edge (Figure 6d). 

Body wall scales 0.32–0.42 mm in length, with an L:W 

of 1.0–1.6. Outer surface of body wall scales smooth to 

mildly granular; inner surface tuberculate. Tentacular rodlets common (Figure 6f): cigar shaped, 0.083–0.105 mm 

in length (L:W = 3.6–5.2).

Coenenchymal scales (Figure 6e) elliptical to circular, 

0.11–0.31 mm in diameter; outer surface bear low granules, inner surface tuberculate.

Remarks. This species is unique within the 

subgenus in having rectangular body wall scales, including 

the marginal scales, and in having such elongate opercular 

scales (L:W up to 4.1). It is also the deepest known species 

within its subgenus.

Types and Type Locality. Holotype: 

Jason II 2098-2-1, dry colony and alcohol fragment, SEM 

stubs C14553-1455, USNM 1134074. Paratype: Jason II

Key to the Species of the Subgenus Plumarella (Dicholaphis)

1. Colonies dichotomously branching; species occurs only off Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. delicata

1′. Colonies pinnately branched or shaped as a bottlebrush; occurs only off Aleutian Islands . . . . . . . . . . . . . . . . . . . . . . . 2

2. Colonies bottlebrush in branching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2′. Colonies pinnately branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

3. Polyps large (3.0–3.6 mm), 9–15 polyps/cm; tentacular rodlets present; coenenchymal membrane often present between 

bases of branchlets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. nuttingi

3′. Polyps small (1.0–1.4 mm), 30–35 polyps/cm; tentacular rods absent; no coenenchymal membrane between branchlets . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. superba

4. Marginal scales (exclusive of adaxials) bear elongate spines consisting of over 60% of length of scale; marginal spine spiny 

or ridged on all surfaces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

4′. Marginal scales (exclusive of adaxials) have a pointed or straight distal edge but never produced into an elongate spine; 

inner surface of marginal spines smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5. Colonies consists of a single branch from which branchlets originate in a pinnate manner; marginal spines finely ridged; 

tentacular rodlets present; species occurs deeper than 800 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. robusta

5′. Colonies consists of several major branches from which branchlets originate in a pinnate manner; marginal spines coarsely 

spiny; tentacular rodlets absent; species occurs at depths shallower than 700 m . . . . . . . . . . . . . . . . . . . . . . . P. echinata

6. Marginal scales rectangular, with a straight distal edge; opercular scales quite elongate, with a L:W over 2.5; species 

occurs deeper than 2500 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. profunda

6′. Marginal scales triangular, with a pointed distal edge; opercular scales triangular but with a L:W less than 2.5; species 

occurs at depths shallower than 500 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 

7. Polyps small (0.9–1.2 mm); body wall scales thick and coarsely serrate; tentacular rodlets absent . . . . . . . . P. aleutiana

7′. Polyps larger (2.0–2.4 mm); body wall scales thin, concave above, and finely serrate; tentacular rodlets present . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. hapala
n u m b e r 6 3 4 • 1 1

2098-2-2, 1 small colony, USNM 1134075. Type locality: 

51°23.404′N, 177°04.549′W (south of Kanaga I., Aleutian Islands), 2514 m.

Material Examined. Types.

Distribution. Known only from type 

locality.

Etymology. Named profunda (Latin for 

deep) for its relatively deep capture depth.

Plumarella (D.) hapala, new species

Figures 1c, 7a–k

?Amphilaphis sp. Wing and Barnard, 2004:24, 41, fig. 11.

Description. Colonies uniplanar, consisting of a small number of branches from which branchlets 

originate in an alternate pinnate manner (Figure 1c), the 

distance between branchlets on one side of a main branch 

ranging from 7 to 11 mm. Holotype 26 cm tall and 22 cm 

wide, with a basal branch diameter of 3.7 mm. Branchlets 

4–5 cm in length, occasionally with secondary branching. 

Branch axis yellow-brown and relatively stiff.

Polyps occurring on all sides of main branches and 

branchlets in a crowded manner (Figure 7a), with 18–25 

polyps/cm toward branchlet tip. Polyps rather large, 2.0–

2.4 mm in length, slightly flared distally, and curved upward 

along branch. Many polyps containing an egg, with each 

egg ~0.9 mm in diameter, and thus considerably swelling 

polyp’s body cavity (Figure 7e). Polyps rather fragile, often 

compressed during collection, and covered with mucous.

Polyps encased with 8 longitudinal rows of body 

wall scales: 6–8 in each abaxial row, 5–7 in outer lateral 

rows, 3–6 in inner lateral rows, and 5–8 smaller scales in 

foreshortened adaxial rows (Figure 7b). Opercular scales 

isosceles triangle in shape (Figure 7f), and 0.55-0.80 mm 

in length (L:W = 2.1–2.5), forming a conical operculum 

over polyp. Outer surface of operculars covered with 

small spines arranged in lines radiating from a common 

proximal point; lower part of inner surface tuberculate 

but distal surface covered with rows of spines as on outer 

side. Six of 8 marginal scales (Figure 7g) pointed (not spinose), the 2 adaxial marginals having a rounded distal 

edge. Pointed marginals up to 0.62 mm in length with an 

L:W of 1.3–1.6; adaxial marginals shorter, with an L:W of 

about 1. Outer surface of marginals smooth to faintly striate whereas proximal inner surface tuberculate and distal 

pointed region smooth and thickened longitudinally but 

not keeled (Figure 7h). Remaining body wall scales (Figure 7i) pointed but progressively shorter toward base of 

polyp, ranging in size from 0.88 to 1.2 mm in length (adaxial scales smaller, only about 0.2 mm in diameter), with 

a L:W of 0.9–1.2. Body wall scales thin, with a relatively 

smooth outer surface, contributing to fragility of polyp; 

distal edges of all polyp scales finely serrate. Tentacular 

rodlets common (Figure 7j), cigar shaped, ranging from 

0.080 to 0.110 mm in length (L:W = 3.5–5.0).

Coenenchymal scales (Figure 7k) elliptical to rectangular, having a smooth outer surface, up to 0.6 mm.

Remarks. Within the subgenus, P. hapala is 

perhaps most similar to P. aleutiana (see Key) but distinctive in having thin, outwardly concave body wall scales 

that contribute to a relatively weak polyp body wall. The 

polyps are also relatively large and often contain an egg, 

which increases the size of the polyp (Figure 7e).

Types and Type Locality. Holotype: 

Ocean Olympic, 1 dry colony, SEM stubs C1449-1452, 

USNM 1006310, and dry branches, AB00-22. Paratypes: Alaska Sea about 52°N, about 175° E, 183 m, 

26 Jan 2000, 1 dry colony, USNM 1006334; Alaskan 

Leader 35-2, 1 dry colony, AB02-23; Delta 5997, 1 dry 

colony, USNM 1011096; Delta 5597-8C-1, 1 large dry 

colony, AB10-0024; Delta 6004-11C-1A, B, fragment, 

USNM 1134095, and 2 dry colonies, AB10-0026; Delta

6005-11B-1, fragment, USNM 1134212, and a dry colony, AB10-0029; Dominator 1997-01-49, 1 dry colony, 

USNM 1134093; Jason II 2102-12-1A, fragment, USNM 

1134094 and AB10-0087; Jason II 2102, 12-1B, fragment, 

USNM 1134213; Vesteraalen 941-186, 4 colonies, USNM 

100796; Western Alaska, 52°11.54′N, 176°07.77′E, 183 

m, 25 Jan 2000, 1 dry colony, AB00-36. Type locality: 

52°11.89′N, 176°17.30′E (off Buldir Reef, west of Kiska 

Island), 20 Oct 2000, 366 m.

Material Examined. Types.

Distribution. Throughout Aleutian Islands 

from western Rat Islands to Islands of Four Mountains, 

120–384 m.

Etymology. Named hapala (Latin for soft to 

the touch, delicate) because the thin body wall scales produce a relatively weak body wall, which is often crushed 

or collapsed during collection.

Plumarella (D.) aleutiana, new species

Figures 1d, 8a–k

?Thouarella sp. Wing and Barnard, 2004:24, 43, fig. 13.

Description. Colonies uniplanar, consisting 

of relatively few main branches from which branchlets 
1 2 • smithsonian contributions to zoology

originate in an alternate pinnate manner, the holotype (the 

largest specimen) colony being 50 cm in height and 36 mm 

in width, consisting of 10 main stems or fronds (Figure 

1d); base of colony 7.5 × 6.5 mm in diameter. Branchlets 

4–9 cm in length, only rarely with secondary branching, 

arranged parallel to one another on each side of a main 

branch every 4–6 mm. Branch axis brown to black in color, 

toward base of large colonies elliptical or flattened in cross 

section (up to 10 mm), the greater axis of ellipse in plane of 

flabellum. Branches and colonies somewhat stiff in tension.

Polyps occurring on main stems and all sides of 

branchlets in a crowded manner, with 20–35 polyps/cm 

toward distal end of a branchlet. Polyps small, 0.9–1.2 

mm in length, and cylindrical to slightly flared distally, 

curving upward along the branches.

Polyps covered with 8 longitudinal rows of body wall 

scales: 5–9 in each abaxial row, 4–6 in outer lateral rows, 

2–5 in inner lateral rows, and 2–4 in adaxial rows (Figure 

1d). Opercular scales (Figure 8e) triangular and curved inward, 0.25–0.41 mm in length (L:W = 1.4–1.9), the adaxial 

operculars always the smallest (Figure 8c); operculum low 

owing to downward curvature of opercular scales. Outer 

surface of operculars quite spiny and even longitudinally 

ridged; inner surface tuberculate proximally and smooth 

distally; edges finely serrate. Marginal scales (Figure 8g, h) 

variable in shape, on some polyps spinose (H:W about 1.6) 

and on other polyps simply triangular (H:W = 1.1–1.3), 

but in all cases, adaxial marginals low and rounded distally 

(H:W about 1.0). Outer surface of marginals bearing low 

ridges and granules; inner surface tuberculate proximally 

and smooth distally. Remaining body wall scales (Figure 

8i) progressively less tall but wider toward base, with H:W 

ratios of 0.5–0.7, and having coarsely serrate distal edge. 

Outer surface of body wall scales usually longitudinally 

ridged but occasionally smooth, polyps of both types directly adjacent to one another. Tentacular rodlets absent.

Coenenchymal sclerites (Figure 8j) irregular in shape 

but usually elongate, up to 0.6 mm in length, their outer surface coarsely granular. Coenenchymal sclerites, which cover 

older large-diameter branches, consisting of much smaller 

spiny spheroids 0.050–0.090 mm in diameter (Figure 8k), 

probably the result of degeneration of once larger sclerites. 

Remarks. Plumarella aluetiana is similar to 

P. superba in many characteristics but differs in colony 

shape (pinnate versus bottlebrush), having coarsely serrate 

body wall scales, opercular scales with a highly spiny outer 

surface and smooth inner surface, and curved operculars, 

resulting in a lower operculum. Also, the outer surface of 

most body wall scales of P. aleutiana is ridged, whereas it 

is smooth in P. superba. 

Types and Type Locality. Holotype: 

Delta 5590, 1 large dry colony, SEM C1421-1424, 

USNM 1011356. Paratypes: Alaska Sea, 52°13′52′′N, 

175°09′32′′E, 183 m, 26 Jan 2000, 1 dry colony, USNM 

1006333, and 1 dry branch, AB00-38; Alb-3319, 1 

branch, USNM 51322; Alb-3500, several large colonies, 

alcohol and dry, USNM 49975; Alb-4779, 1 colony, 

USNM 49579; Dominator 1997-01-49, 1 colony, USNM 

1134746; Let’s Go 861-52, 1 colony, USNM 1134754; 

Let’s Go, 861-57, 3 dry colonies, USNM 1134508; MF

(Harvester) 802, VH80-30, 1 colony, USNM 82125;

MF 833-16, 1 colony, USNM 1134538; Pacific Knight

941-29, 1 branch, USNM 100808; Pacific Knight 941-

73, 1 branch, USNM 100842; Pacific Knight 941-204, 

2 colonies, USNM 100752 and 100847; Patricia Lee, 

51°18′42′′N, 179°30′04′′E, 329 m, 20 Oct 2000, 1 dry colony, USNM 1006302; Vesteraalen 2001-01-5, dry colony 

and alcohol fragment, USNM 1006032, and 1 dry branch, 

AB00-40; Vesteraalen 941-15, 1 colony, USNM 1012501; 

51°43′52′′N, 178o43′31′′W, 256 m, 2 Apr 2000, 1 dry colony, USNM 1006739; 51°32′, 179°15′W, col. K.K.Chew, 

278–289 m, Sep 1968, 5 large dry colonies, USNM 60344 

and 60345; 51°55′05′′N, 176°38′13′′E, 14 Feb 2000, 1 

dry colony, USNM 1006246. Type locality: 51°43′22′′N, 

176°29′10′′W (off Beyer Bay, Adak Island), 85 m.

Material Examined. Types; Nontypes: 

Alaskan Leader 35, 1 dry colony, AB10-0019; Delta

5600, 4 dry colonies, AB02-116; Delta 5989-14B-1, 

fragment, USNM 1134747, and 1 dry colony, AB; Delta

5993-7C-4 and 6, fragments, USNM 1134504, and 1 dry 

branch, AB10-0021, 23; Delta 6007-13C-2, dry fragment, 

USNM 1135852, and 1 dry colony, AB10-0067; Delta

6199-5E-7, fragment, USNM 1134537, and 2 dry colonies, AB; Delta 6202-14D-4, fragment, USNM 1134753, 

and 2 dry colonies, AB; Delta 6208-21D-5, fragment, 

USNM 1134748, and 2 dry branches, AB10-0042; 

Delta 6209-21C-1, 1 dry colony, AB; Delta 6219-3B-2, 

fragment, USNM 1135874, and 1 dry branch, AB10-

0058; Delta 6221-2B-9, fragment, USNM 1134755, and 

1 large dry colony, AB10-0060; Jason II 2095-2-1-2, 1 dry 

colony, AB10-0069; Jason II 2102-6-5, 1 branch, USNM 

1135289 and AB10-0085; Jason II 2102-7-1C, D, E, 3 

fragments, USNM 1134507, 1134539, and 1 dry colony, 

AB10-0086.

Distribution. Common throughout Aleutian Islands from Near Island in the west to Unalaska to 

the east, and Pribilof Canyon, 79–517 m.

Etymology. Being the most commonly collected Plumarella in the Aleutian Islands, the species is 

named for the island group.
n u m b e r 6 3 4 • 1 3

Plumarella (D.) superba (Nutting, 1912),

new combination

Figures 2c, 9a–l

Primnodendron superbum Nutting, 1912:71–72, pl. 9, figs. 2, 2a, pl. 19, 

fig. 4.

Thouarella striata.—Nutting, 1912:69, pl. 10, figs. 2, 2a.— Heifetz, 2002:26 

(listed).—Wing and Barnard, 2004:24 (listed).—Heifetz et al., 2005:133 

(listed).—Stone and Shotwell, 2007:107 (listed).

Thouarella (Amphilaphis) superba.—Kükenthal, 1919:412–413; 1924: 291.

Thouarella superba.—Wing and Barnard, 2004:24, 44, fig. 14.—Heifetz et 

al., 2005:133 (listed).—?Dautova, 2007:299, fig. 2.—Stone and Shotwell, 2007:107 (listed).

Thouarella (Thouarella) superba.—Cairns and Bayer, 2009:28 (listed).

Description. Colonies uniplanar, consisting 

of relatively few main branches from which numerous 

branchlets originate in a modified bottlebrush arrangement (Figure 2c), i.e., short branches (1.5–2 cm in length) 

diverge from both sides of main branches in plane of 

colony, whereas even shorter branches (0.5–1.0 cm) diverge from anterior and posterior faces of flabellum, often 

slightly longer on the anterior face (by definition). The 

larger branchlets and sometimes even the shorter ones 

usually bifurcate or give off several still shorter branchlets. Because of the unequal lengths of the branchlets on 

the main branches, the bottlebrush form is described as 

modified. Branches of colony somewhat rigid and wiry. 

Holotype now in 3 pieces, the largest 14 cm tall and 9 cm 

wide; largest specimen examined (USNM 1011277) 26 cm 

tall and 32 cm wide (Figure 2c), with a basal branch diameter of 8.3 mm. Axis black and woody in texture.

Polyps occurring on main stems and on all sides of 

branchlets in a crowded manner, with ~30–35/cm. Polyps 

small, 1.0–1.3 mm in length (not 2 mm as reported by 

Nutting, 1912), and slightly larger at their distal end (but 

not flared), facing upward. 

Polyps covered with 8 longitudinal rows of body wall 

scales: 5–7 scales in abaxial rows, 4–6 in outer lateral rows, 

3 or 4 in inner lateral rows, and only 1 or 2 in adaxial rows, 

the adaxials being reduced in number and size (Figure 9c, 

k). Opercular scales (Figure 9e) triangular, 0.20–0.38 mm 

in length (L:W = 1.3–2.0), the adaxial operculars usually 

smallest. Outer surface of operculars smooth to mildly spinose; inner surface longitudinally ridged as well as tuberculate. Marginal scales (Figure 9g) triangular (pointed) to 

spinose, the distal spine decreasing in size toward adaxial 

side, where adaxial marginals (and sometimes the inner 

lateral marginals) have only rounded distal edges (Figure 

9h). Abaxial marginal scales 0.33–0.48 mm in length 

(L:W = 1.2–1.8), with a smooth outer surface and a tuberculate inner surface, the distal spine being circular in cross 

section and smooth all around; scales thick. Adaxial marginals only about 0.35 mm in length, with a low L:W of 

about 0.75). Abaxial submarginal scales similar to abaxial 

marginals, bearing distal spines; otherwise, distal margin 

of remaining body wall scales transition from spinose to 

pointed to rounded moving toward base of polyp, and 

L:W ratios vary from 0.8 to 1.1. Adaxial body wall scales 

(Figure 9j) much smaller and elliptical in shape, having 

a greater diameter of 0.11–0.17 mm (Figure 9k). Edges 

of all scales slightly serrate, and outer surface of all body 

wall scales proximal to operculars quite smooth (but see 

below). Tentacular rodlets absent.

Coenenchymal sclerites (Figure 9l) irregular in shape 

but usually elongate, rarely over 0.45 mm in length, their 

outer surface bearing coarse granules. 

Remarks. The species was originally placed 

in a separate genus, Primnodendron, Nutting (1912) 

noting that it had strong affinities to both Plumarella

and Thouarella. Kükenthal (1919, 1924) later placed it 

in Thouarella, probably because it has a bottlebrush arrangement of branchlets, a common arrangement for 

Thouarella. Cairns and Bayer (2009) followed suite, officially, but incorrectly, synonymizing Primnodendron with 

Thouarella. However, emphasizing the characteristic of 

the inner marginal keel of Thouarella (see description of 

subgenus above and Taylor et al., in press), this species 

must be transferred to Plumarella, and to the subgenus Dicholaphis, as discussed above. It is one of only two species 

in the genus to have a bottlebrush arrangement of branchlets (see Key), and is distinguished from P. nuttingi by its 

smaller polyps and lack of a coenenchymal membrane at 

the base of its branchlets.

Although most of the specimens examined have very 

smooth outer surfaces of the body wall scales (Figure 

9a, e, h, j), six lots (USNM 1009951, 100814, 1006323, 

1135292, 1135852, and 1135265) are identical in all respects but have radiating ridges on their body wall scales 

(Figure 9b, f). These specimens were collected somewhat 

deeper (to 888 m) than the other specimens (40–102 m). 

Furthermore, two other specimens (USNM 57527, 57536) 

are also otherwise identical but are pinnately branched and 

were collected from an even greater depth of 882–1258 m. 

A station error cannot be ruled out.

The specimen reported as Thouarella striata by Nutting (1912) from Albatross 4778 could not be found, but 

a small vial of several hundred disarticulated sclerites was 

found labeled as such in Nutting’s handwriting, which are 
1 4 • smithsonian contributions to zoology

consistent with an identification of P. superba. They were 

also collected at the type locality of P. superbum.

Types and Type Locality. The holotype, 

now in three pieces, two small vials of sclerites, and SEM 

stubs C1430-1432, are deposited at the USNM (30691). 

Type locality: Alb-4778: 52°612′N, 179°52′E (Petrel Bank 

north of Semisopochnoi Island), 60–79 m. 

Material Examined. Types. Alb-3501, 

1 branch (pinnate form), USNM 57527; Alb-4777, 1 

colony, SEM stub B280, C1433, USNM 50150, and another colony, USNM 57535; Alb-4778, sclerites only (T. 

striata of Nutting, 1912), USNM 1135267; Alb-4781, 1 

colony (pinnate form), USNM 57536; Delta 5993-7C-4, 

fragment, USNM 1135265, and1 dry colony, AB; Delta

6199-5E-1, fragment, USNM 1134506; Delta 6199-5E-7, 

dry colony, AB10-0068; Delta 6199-5E-8, 1 dry colony, 

AB; Delta 5E-9, fragments, USNM 1135266, and 5 dry 

colonies, AB; Delta 5199-5E-14, 1 dry colony, AB10-

0031; Delta 6199-5E-15, fragment, USNM 1134756; 

Delta 5591, 1 colony, USNM 1011276, and 1 dry colony, 

AB02-144; Delta 5604, 1 colony, USNM 1011277; Delta

6203-18-1, fragment, USNM 1134749 and AB10-0039; 

Delta 6206-18-2, fragments, USNM 1135287; Delta

6209-21C-7, fragments, USNM 1135278 and AB10-0049; 

Dominator-1997-01-49, 4 alcohol-preserved colonies, 6 

dry colonies, SEM stubs C1464-1466, USNM 1009951, 

1135264, respectively; Esperanza PC 7-3-6-3, 1 branch, 

USNM 1116845; Let’s Go 861-79, fragments, USNM 

1135290; MF 833-24, 1 branch, USNM 1135292; Spirit 

of the North, 51o21.66’N, 178o31.99’W, depth unknown, 

20 Oct 2000, 1 dry colony, AB00-68; Vesteraalen 941-

36, 4 dry colonies, USNM 100814; Vesteraalen 941-94, 

5 colonies in alcohol, 1 dry colony, USNM 100845; Vesteraalen 2001-01-5, 1 colony, USNM 1006027; Western 

Alaska, 52°11.54′N, 176°07.77′E, 183 m, 12 Jul 2000, 1 

dry colony, USNM 1006323.

Distribution. Aleutian Islands: from eastern Near Islands to Umnak Island, including Petrel Bank 

and Pribilof Canyon, 40–1258 m. Also, off southeastern 

Sakhalin Island, 29 m (Dautova, 2007).

Plumarella (D.) nuttingi, new species

Figures 2d, 10a–k

Description. Colonies uniplanar, consisting 

of relatively few main branches from which branchlets diverge from all sides in a bottlebrush fashion (Figure 2d); 

however, longer branchlets growing in plane of flabellum 

producing a compressed bottlebrush form. Largest colony 

(holotype) 19 cm in height and width, with a basal branch 

diameter of 4.4 mm; axis pale yellow, polyps white. 

Branchlets up to 7 cm in length, usually not subdivided. 

Basal 6–8 mm of each branchlet often immersed in a thin 

coenenchymal membrane that is continuous with main 

branch; coenenchymal scales present in this membrane 

(Figure 10i, k).

Polyps usually occurring on all sides of branches and 

branchlets, occasionally in an alternate biserial manner. 

Polyps large (3.0–3.6 mm in length), standing perpendicular to branchlets, having a slightly upturned distal end, 

and slightly flared distally. Because of large size of polyps, only 9–14/cm on a branchlet. Polyps fleshy, the tissue 

often preventing the operculars from closing in the preserved condition; operculars also appearing to unite basally in a fleshy internal web.

Polyps covered with 8 longitudinal rows of body wall 

scales: 7 or 8 in each abaxial and outer lateral row, 6 or 

7 in inner lateral rows, and 6 or 7 smaller scales in reduced adaxial rows (Figure 10c). Opercular scales (Figure 

10e) isosceles triangular in shape, with an elongate distal 

spine and thus a high L:W ratio (1.0–1.5 mm in length, 

L:W = 2.9–5.0). Outer surface of operculars longitudinally 

ridged and quite spiny, especially on distal region; inner 

surface tuberculate proximally and ridged distally; edges 

of scales finely serrate. Marginals (Figure 10f, g) shorter 

and squatter, more equilateral triangular in shape, their 

distal margin pointed to spinose, the spine consisting of 

25–40% length of scale. Marginals 0.70–1.1 mm in length 

with an L:W ranging from 1.0 to 1.6. Outer surface of 

marginals relatively smooth, sometimes with a longitudinal ridge; inner surface largely tuberculate, the distal edge 

and spine being smooth or bearing a low ridge but not 

keeled (Figure 10g). Remaining body wall scales (Figure 

10h) thin, elliptical in shape, with finely serrate distal edges 

that flare outward; height of body wall scales 0.45–0.55 

mm, with an L:W of 1.0–1.25. Outer edges of body wall 

scales smooth with low ridges; inner surface tuberculate. 

Tentacular rodlets (Figure 10j) common, 4 to 5 across at 

base of a tentacle, 2–3 across distally. Rodlets cigar shaped 

(Figure 10j,k), 100–0.185 mm in length (L:W = 3.5–6.2). 

Coenenchymal scales (Figure 10i) 0.2–0.4 mm in diameter, elliptical, and flat, with a granular outer surface 

and a tuberculate inner surface.

Remarks. Within the subgenus this species has 

the largest polyps and consequently the fewest polyps per 

centimeter, the longest and most slender opercular scales 

(L:W = 5), and a distinctive coenenchymal membrane that 

often occurs at the bases of branchlets, this membrane 

containing coenenchymal sclerites.
n u m b e r 6 3 4 • 1 5

Types and Type Locality. Holotype: Spirit 

of the North, 1 dry colony and many separate polyps, 

SEM stubs C1444-1448, 1462-1463, USNM 1006317; 

Jason II 2102-6-4, fragment, USNM 1134464, and 1 

dry colony, AB10-0084; Western Viking, 52°07.7′N, 

176°00.11′W, depth unknown, 8 Feb 2000, 1 dry colony, 

USNM 1006331, and 1 dry branch, AB. Type Locality: 

51°59.32′N, 177°41.33′W (off Kanaga Island, Andreanof 

Islands), 492 m, 20 Oct 2000.

Material Examined. Types.

Distribution. Aleutian Islands from Amchitka Pass to Great Sitkin Island, 492–888 m.

Etymology. Named in honor of Charles C. 

Nutting, specifically in recognition of his revision of the 

North Pacific deep-water octocorals (Nutting, 1912).

Plumarella (D.) echinata, new species

Figures 2a, 11a–k

Plumarella spinosa .—Nutting, 1912:63, in part (Alb-4769).

Description. Colonies uniplanar, consisting 

of a moderate number of branches (fronds) from which 

branchlets originate in an alternate pinnate manner, the 

largest colony examined (the holotype) 25 cm in height 

and 28 cm in width, consisting of 13 fronds; basal branch 

diameter 3.8 mm (Figure 2a). Branchlets relatively short, 

usually 2–5 cm in length but up to 6 cm long toward 

base of colony, the distal ends of branchlets from adjacent 

fronds slightly overlapping such that the full plane of the 

colony sieves the passing water current. Branchlets rarely 

secondarily branched, arranged parallel to one another 

on each side of a main branch at intervals of 3–4 mm. 

Branch axis bronze, polyps white. Colony branches fairly 

flexible.

Polyps occurring on main branches and all sides of 

branchlets in a crowded manner (Figure 11a), with 20–30 

polyps/cm toward distal end of a branchlet (Figure 11a). 

Polyps small, 1.1–1.4 mm in length, and slightly flared distally, curving upward along branch.

Polyps covered with 8 longitudinal rows of body wall 

scales: only 3–5 in each abaxial row, 3 or 4 in outer lateral 

rows, 2 or 3 in inner lateral rows, and 2–4 smaller scales 

in adaxial rows (Figure 11c, d, j). Opercular scales (Figure 

11e) isosceles triangle in shape and slightly curved over the 

polyp (Figure 11c), the abaxial and outer lateral operculars 

being so long (up to 0.77 mm, L:W = 2.5–3.9) that they 

do not form a cohesive operculum; instead, steepled above 

other opercular scales (Figure 11c, d). Adaxial operculars 

shorter (0.35 mm) and lying flat over polyp. Outer surface 

of operculars highly spiny and longitudinally ridged; inner 

surface smooth to faintly ridged. Six of 8 marginal scales 

(all but the adaxials) prominently spinose (Figure 11f), 

consisting of a square to diamond-shaped basal region 

clearly demarcated from the distal spine, the latter circular 

in cross section (Figure 11g) and constituting as much as 

2/3 length of scale (Figure 11f). Marginal scales up to 0.90 

mm in length, with an L:W = 2.0–3.1, the larger marginals 

obscuring view of underlying operculum. Adaxial marginals much smaller (about 0.25 mm in diameter), circular to 

square (L:W = 1), with a straight distal edge (Figure 11c, 

d). Outer surface of marginals spiny; inner surface of basal 

portion tuberculate, whereas inner distal spine covered 

with low, discontinuous, spiny ridges (Figure 11g). Submarginals (Figure 11h) bear shorter distal spines, 0.45–

0.50 mm in length (L:W = 1.3–1.5). Remaining body wall 

scales rather thick, (Figure 11i) with pointed to rounded 

distal edges and large granules on outer surface, about 0.4 

mm in length, with L:W ratios ranging from 0.98 to 1.4; 

adaxial body wall scales smaller than others. Outer edges 

of all polyp scales finely serrate. Tentacular rodlets absent.

Coenenchymal sclerites (Figure 11k) irregular in 

shape, usually elongate, up to 0.55 mm, and coarsely 

granular on outer surface.

Remarks. Plumarella echinata is distinctive 

among the species in the subgenus in having elongate marginal spines that are spiny (not ridged or smooth) and in 

having relatively few scales in the body wall rows.

Types and Type Locality. Holotype: 

MF-833-38, 1 colony, SEM stubs C1434-1436, USNM 

1135009. Paratypes: Alaska Sea, 52°N, 175°W, depth 

unknown, 1 dry colony. USNM 1006227; Alb-3480, 

1 branch, USNM 57546; Alb-4769, 1 colony, USNM 

43107; Alb-4771, 1 branch, USNM 57534; Delta 6199-

5E-14, fragment, USNM 1135006; Esperanza PC 7-4-6-3, 

2 dry colonies, AB10-0097; Jason II 2095-2-1-2, fragment, USNM 1135010; Jason II 2095-2-7-3, 1 dry colony, AB10-0068; Jason II 2100-4-2, fragment, USNM 

1135993; Jason II 2104-10-2, fragment, USNM 1135995, 

and dry colony, AB10-0092; Let’s Go 861-57, 4 colonies, 

USNM 1134976; Let’s Go 861-58, 2 dry colonies, USNM 

1134975; MF-833-15, 3 colonies, USNM 1134974; Western Viking, 52°17′N, 176°14′E, 284 m, 1 dry colony, 

USNM 1002569. Type locality: 51°42.9′N, 178°51.5′W 

(Gareloi Island [Delarof Islands], Aleutians), 582 m.

Material Examined. Types.

Distribution. Aleutian Islands from off Buldir Reef (Rat Islands) to Amukta Pass, and Bowers Bank, 

150–1,692 m.
1 6 • smithsonian contributions to zoology

Etymology. Named echinatus (Latin for 

spiny) because not only are the marginal scales spinose, 

but the marginal spines also bear smaller spines.

Plumarella (D.) robusta, new species

Figures 2f, 12a–j

Description. Colonies consisting of one 

main branch, a uniplanar pinnate frond, from which 

branchlets originate in alternating pinnate fashion (Figure 

2f), spaced approximately 3–3.5 mm apart on one side of 

main branch in parallel fashion. Branchlets 7–8 cm long 

toward base of colony, gradually increasing in length toward middle of colony and then decreasing to about 4 cm 

at branch tip; all branchlets simple. Largest colony examined (the holotype) 25 cm tall and 13 cm in width, with 

a basal branch diameter of 1.7 mm. Colony fairly flexible 

and pale yellow in life.

Polyps occurring on main stem and all sides of branchlets in a crowded manner (Figure 12a), with ~20 polyps/

cm near branchlet tips. Polyps moderate in size, 1.4–1.8 

mm in length, cylindrical to slightly flared distally, and 

curving upward along branchlets. 

Polyps covered with 8 longitudinal rows of body wall 

scales: 4 or 5 in each abaxial row, 3 or 4 in outer lateral 

rows, 3 or 4 in inner lateral rows, and several smaller 

scales on adaxial face. Opercular scales (Figure 12e) triangular to somewhat rectangular (some operculars within 

same operculum with a pointed tip, others rounded), 

0.30–0.67 mm in length (L:W = 2.0–2.6), curved inward 

over polyp, and with a concave outer surface. Outer surface of operculars bearing low spiny ridges that radiate 

from a proximal point; inner surface tuberculate proximally but bearing multiple ridges distally (Figure 12e). 

Marginal scales (Figure 12d, f, g) prominently spinose, 

with a rectangular basal portion wider than tall and 

clearly demarcated from distal spine, the latter circular 

in cross section and constituting as much as 60% length 

of scale. Marginal scales up to 0.77 mm in length (L:W 

= 1.4–2.6), the adaxial marginals much smaller (Figure 

12d), only 0.35–0.50 mm in length, but also bearing a 

short distal spine. Outer surface of basal portion of marginals granular, inner surface tuberculate. Distal spine 

circular in cross section and bearing 7 or 8 elongate spiny 

longitudinal ridges, the space between ridges deeply channeled, the ridges occurring uniformly on all sides of spine 

(Figure 12g). Occasional marginal bearing a short (0.1 

mm) spur on proximal edge (Figure 12f). Remaining body 

wall scales (Figure 12i) spinose to pointed (sometimes 

bifid), often irregular in shape (0.31–0.56 mm in length, 

L:W = 0.70–2.2), quite thick and distinctly convex; adaxial body wall scales much smaller. Outer and inner 

surfaces of body wall scales similar to that of marginals. 

Distal edges of all polyp scales finely serrate. Tentacular 

rodlets (Figure 12j) common: cigar shaped, 0.055–0.065 

mm in length (L:W = 4.0–4.5).

Coenenchymal sclerites (Figure 12h) elongate, up to 

0.6 mm (L:W = 1.5–2.5), but plate-like: outer surface covered with spiny ridges radiating from a central point, inner 

surface tuberculate.

Remarks. Although similar to P. echinata in 

marginal spine length, P. robusta differs in having ridged 

marginal spines and in having much thicker body wall 

scales that are markedly convex and commonly spinous 

(also see Key). This species co-occurs with P. spicata at 

two stations.

Types and Type Locality. Holotype: Jason 

II 2104-11-1, a dry colony and a small fragment in alcohol, SEM stubs C1456-1459, USNM 1135992. Paratypes: 

Jason II 2101-4-3, dry colony, USNM 1135884; Jason II

2101-7-7, 1 dry colony and a fragment in alcohol, USNM 

1135991; Jason II 2102-6-2, 1 dry colony and a fragment 

in alcohol, USNM 1135987.

Material Examined. Types.

Distribution. Aleutian Islands from a relatively circumscribed region from Amchitka Pass to Adak 

Canyon, 712–1,061 m.

Etymology. Named robusta (Latin for hard, 

strong, robust), for the robust polyps produced by its thick 

body wall scales.

Genus Fanellia Gray, 1870

Fanellia Gray, 1870: 46.—Bayer, 1982:134–135 (key).—Bayer and Stefani, 

1989:470–471 (key).—Cairns and Bayer, 2009:40–41, fig. 12e–m.—

Cairns, 2010:433.

Diagnosis. Colonies usually uniplanar, 

branchlets arranged dichotomously or in pinnate fashion. 

Polyps arranged in pairs or whorls, the polyps facing upward. Polyps protected by 4–8 rows of thick body wall 

scales, in some cases the adaxial and inner lateral scales 

being absent below the marginals resulting in a naked adaxial side. Outer surface of body wall scales covered with 

prominent tubercles, arranged in ridges. Marginal scales 

not folded over operculars, the latter bearing a longitudinal keel or set of ridges on inner surface. Coenenchymal 

scales consisting of an inner layer of tuberculate spheroids 

covered by thick polygonal scales. 
n u m b e r 6 3 4 • 1 7

Discussion. The eight species known in the 

genus are keyed by Bayer and Stefani (1989). Recent 

works discussing species in this genus are included in the 

synonymy above. 

Distribution. Western and central Pacific 

from New Caledonia to Alaska, 82–1,341 m.

Fanellia compressa (Verrill, 1865)

Figure 13a–d, h

?Prymnoa verticillaris.—Ehrenberg, 1834:357 (not Gorgonia verticillata Pallas, 1766).

Primnoa compressa Verrill, 1865:189.

Fanellia compressa.—Gray, 1870:46.—Bayer, 1982:136–140, figs. 11–14.—

Bayer and Stefani, 1989:471 (key).—Heifetz, 2002:22.-–Not Wing and 

Barnard, 2004:fig. 16 (=F. fraseri).—Stone and Shotwell, 2007:106 

(listed).-–Cairns and Bayer, 2009:30 (listed).

Calligorgia compressa.–Studer, 1878:647.—Heifetz, 2002:26 (listed).

Caligorgia compressa.—Wright and Studer, 1889:80.–Verssluys, 1906:81–

82.—Kükenthal, 1919:379; 1924:276.

Description. Colonies uniplanar, up to 80 cm 

in height and 40 cm in width, with a basal branch diameter 

up to 1.8 mm (USNM 60281). Branching dichotomous, 

occurring every 3–5 cm but largely restricted to proximal 

third of colony, the end branches being straight and up to 

50 cm in length; branches on edge of colony sometimes divided into a series of smaller branches in quick succession 

producing a lyrate arrangement in those regions. Dried tissue pale gray, branch axis metallic bronze.

Polyps occurring in close-spaced whorls on all branch 

surfaces: 8–12 polyps per whorl on distal branches 

(Figure 13c) and up to 25 polyps per whorl on largerdiameter basal branches; 4–6 whorls/cm. Polyps sometimes obliquely arranged in whorls approximating a spiral 

arrangement. Polyps cylindrical, closely spaced within 

whorl (Figure 13c), as well as abutting to polyps in adjacent whorls, with little intervening coenenchyme. Polyps 

1.6–2.0 mm in length.

 Polyps covered with 8 longitudinal rows of body 

walls scales: 7–10 scales in each abaxial row, 5–7 in outer 

lateral rows, 2 or 3 in inner lateral rows, and usually only 

1 pair adaxially. Opercular scales triangular, abaxial operculars up to 0.50 mm in length, adaxial much smaller 

(only about 0.25 mm in length); L:W = 1.4–1.6. Marginal 

body wall scales not significantly different from other 

body wall scales. Distal abaxial body wall scales roughly 

square in shape but toward polyp base becoming progressively wider and ultimately on the seventh to tenth abaxial 

scale away from operculum bearing elongate extensions 

or spurs on their adaxial side some of which extend to 

adaxial polyp face (Figure 13a, b). All outer lateral scales 

roughly the same size, but those in third to sixth position 

also bearing adaxial extensions (Figure 13a,b). Inner lateral and adaxial scales much smaller and without extensions. Although abaxial and outer lateral scales bearing 

elongate adaxial extensions, adaxial side of polyp mostly 

unprotected because of small size and number of inner 

lateral and adaxial scales (Figure 13d). Outer surfaces of 

all body wall and opercular scales covered with tall spiny 

(aculeate), discontinuous ridges separated by deep furrows, characteristic of the genus. 

Coenenchymal scales consisting of closely fitting polygonal plates and numerous tuberculate spheroids, the 

latter 0.060–0.090 mm in diameter (Figure 13h). 

Remarks. Fanellia compressa was well described and illustrated by Bayer (1982), but he neglected 

to include opercular and adaxial polyp views and coenenchymal tuberculate spheroids, which are included herein. 

This report also adds 30 new records for this species, 

which previously had been known from only three specific 

localities, extending its known range east to the Islands of 

Four Mountains.

Fanellia compressa is quite similar to F. fraseri and 

is sympatric for much of its distribution, although it occurs farther west and not as far east as the known distribution of F. fraseri. Fanellia compressa differs in having 

more outer lateral body wall scales (5–7 versus 2–3, respectively), which usually have adaxial extensions (F. fraseri does not have these extensions), larger polyps (1.6–2.0 

versus 1.1–1.5 mm), longer distal branches (up to 50 versus 15 cm), dichotomous branching (versus pinnate), 4–6 

whorls/cm (versus 6–7/cm for F. fraseri), and a tendency to 

be taller than broad, whereas F. fraseri is usually equally 

tall as broad.

Both species carry commensal ophiuroids of the 

same color of the coenenchyme, having a disc diameter 

of 7–8 mm.

Types and Type Locality. The type is presumed to be lost (Cairns and Bayer, 2009). Type locality: 

“Aleutian Islands,” depth unknown.

Material Examined. Alaska Sea, 

52°09′24′′N, 173°44′19′′E, 318 m, 5 Feb 2000, 1 dry branch, 

USNM 1006336, and 1 dry colony, AB00-39; Alaskan Trojan, 51°32′07′′N, 177°56′42′′W, 346 m, 1 dry colony, USNM 

1006152; Delta 5E-2, 1 branch, USNM 1123883, and 1 dry 

colony, AB10-0054; Delta 5E-3, 1 branch, USNM 1123884, 

and 1 dry colony, AB10-0100; Delta 5E-5, 1 branch, USNM 

1123885, and 1 dry colony, AB10-0054; Delta 5615, 
1 8 • smithsonian contributions to zoology

51°53.52′N, 176°11.294′E, 156 m, 22 Jul 2002, 1 large dry 

colony, AB02-134; Delta 6199-5E-9, dry branch, USNM 

1123886, and 1 dry colony, AB10-0110; Delta 6221-2B-1, 

branch, USNM 1123874, and 1 dry colony, AB10-0061; 

Delta 6223-2D-1, 1 branch, USNM 1123382, and dry 

branches, AB10-0062; Dominator 1997-01-49, 3 colonies, USNM 1009944; Dominator 1997-01-78, 1 large 

colony, USNM 1009942; Dominator 2000-01-156, 1 dry 

colony, USNM 1010176; Jason II 2101-1-5, 1 branch, 

USNM 1123878, and 1 dry colony, AB10-0077; Jason II

2101-4-1, 1 branch, USNM 1123879, and 1 dry colony, 

AB10-0078; Jason II 2101-7-6, 1 branch, USNM 1123880, 

and 1 dry colony, AB10-0124; Jason II 2101-10-3, 1 

branch, USNM 1123881, and 1 dry colony, AB10-0081; 

Jason II 2102-14-1, 1 branch, USNM 1136354; Let’s Go861-50, 1 dry branch, USNM 1136353; Let’s Go-861-79, 

1 branch, USNM 1123873; MF 833-3, 1 branch, USNM 

82122; MF 833-47, 2 large colonies, USNM 1123826; 

North Pacific, 51°50.33′N, 174°15.82′W, 443 m, 20 Oct 

2000, 2 dry colonies, USNM 1006337; Pacific Knight

941-204, 1 dry colony, USNM 100810; Sea Storm 116, 

2 dry colonies, 3 branches, 1010256; Starlight 84-1-7, 

1 colony, USNM 100747; Vesteraalen 941-51, 1 dry colony, USNM 100833; Vesteraalen 941-153, 1 large dry 

colony, USNM 100813; Vesteraalen 941-163, 1 large dry 

colony, USNM 100811; Western Viking, 52°17′09′′N, 

176°13′57′′E, 265 m, 25 Jun 2000, 1 dry colony, USNM 

1006321; 52°11.17′N, 176°17.00′E, 220 m, 20 Oct 2000, 

1 dry colony, AB00-22; 53°04′N, 170°12′W, depth unknown, 4 Aug 2002, 1 colony in alcohol, AB02-24; specimens reported by Bayer (1982).

Distribution. Known only from the Aleutian Islands from the Near Islands to Islands of Four 

Mountains, 82–1,061 m, although most records are from 

150 to 300 m.

Fanellia fraseri (Hickson, 1915)

Figure 13e–g, i

?Prymnoa verticillaris.—Ehrenberg, 1834:357 (not Gorgonia verticillata

Pallas, 1766).

Caligorgia fraseri Hickson, 1915:553–554, pl. 1, fig. 2, text fig. 5; 1917:23 

(listed).—Kükenthal, 1924:279.

Fanellia fraseri.—Bayer, 1982:140–144, figs. 15–17.—Bayer and Stefani, 

1989:471 (key).—Heifetz, 2002:22 (listed).—Wing and Barnard, 2004: 

24 (listed).—Heifetz et al., 2005:132, 136.—Stone and Shotwell, 2007: 

106 (listed).—Cairns and Bayer, 2009:30 (listed).

?Fanellia compressa.—Wing and Barnard, 2004:fig. 16.

Description. Colonies uniplanar and approximately equally tall as wide: up to 36 cm in height and 32 cm 

in width, with a basal branch diameter of 9.5 mm (USNM 

82123). Branching loosely alternate pinnate, the main 

branch being straight (not geniculate), branchlets originating 

from the same side of a main branch separated by 6–15 mm. 

Branching throughout colony, resulting in relatively short 

end branches, i.e., usually less than 15 cm. Dried tissue a pale 

gray but pink when alive; branch axis a brownish-yellow.

Polyps occurring in closely spaced whorls on all 

branch surfaces: 5–8 polyps per whorl on distal branches, 

increasing to 12 on larger-diameter branches (Figure 13g); 

6–7 whorls/cm. Polyps slightly clavate and closely spaced 

within and between whorls. Polyps 1.1–1.5 mm in length.

Polyps covered with 6 to 8 longitudinal rows of body 

wall scales: 7–8 scales in each abaxial row, 2–3 in outer 

lateral rows, 1 in inner lateral rows, and 0–1 adjacent to 

adaxial operculars, resulting in a naked adaxial side (Figure 13f). Opercular scales triangular, the abaxial operculars up to 0.45 mm in length, the adaxial operculars much 

smaller (Figure 13f); L:W = 1.4–2.0. Each abaxial body 

wall scale in the third through sixth position from operculum bearing a short adaxial wing-like extension; however, 

no other body wall scales with a similar spur (Figure 13e). 

As with F. compressa, the outer surfaces of all scales covered with spiny ridges.

Coenenchymal scales consisting of tuberculate spheroids (Figure 13i) 0.075–0.120 mm in diameter.

Remarks. As with Fanellia compressa, F. fraseri was fairly well described and illustrated by Bayer 

(1982), but he also neglected to include opercular and 

adaxial polyp views and coenenchymal tuberculate spheroids, which are included herein. Otherwise, see Remarks 

for F. compressa.

Material Examined. Delta 5592, 1 dry 

colony, USNM 1011274; Delta 6199-5E-13, 1 branch, 

USNM 1123888, and 1 dry colony, AB10-0044; Delta

5E-4, 1 branch, USNM 1123889, and 1 dry colony, AB10-

0054; Delta 6206-18-14, 1 branch, USNM 1123890, and 

1 dry colony, AB10-0041; Delta 6209-21C-2, 1 branch, 

USNM 1123891, and 1 dry colony, AB; Delta 6209-21C-4, 

1 branch, USNM 1123892; Jason II 2101-1-1, 1 branch, 

USNM 1123877, and 1 dry colony, AB10-0071; Jason II

2104-2-2, 1 branch, USNM 1123887; MF 802-VH-80-30, 

1 colony, USNM 82123; Starlight 84-1-7, 1 colony, 

USNM 1013158; Vesteraalen 5, 1 dry colony and 1 alcohol branch, USNM 1006138; 53°04′N, 170°12′W, depth 

unknown, 4 Jun 2002, 1 dry colony, AB02-24; specimens 

reported by Bayer (1982).
n u m b e r 6 3 4 • 1 9

Types and Type Locality. Two syntypes, 

both dry, are deposited at the BM (1962.7.20.821-822). 

Type localities: Albatross and Portland Banks (just northeast of Kodiak Island), Gulf of Alaska, 92–183 m.

Distribution. Known only from Alaska 

from Amchitka Pass to the Albatross Bank northeast of 

Kodiak Island, 86–1,341 m, although most records are 

from 100 to 150 m.

Genus Primnoa Lamouroux, 1812

Primnoa Lamouroux, 1812:188.—Cairns and Bayer, 2005:226–228 (genus 

revision and key to species); 2009:41–42, fig. 13h–p.

Diagnosis. Colonies dichotomously branched 

and usually bushy. Polyps closely spaced and randomly 

arranged (not in pairs or whorls) but usually facing downward. Polyps fleshy, each covered with 6 longitudinal 

rows of scales, the adaxial side largely naked; 8 highly 

concave marginal scales, the adaxials often quite small. 

Well-developed operculum present, the operculars keeled 

on inner surface. Tentacular rodlets common.

Discussion. The four species in this genus 

were revised by Cairns and Bayer (2005), which also includes a key to the species.

Distribution. North Pacific, North Atlantic, 

Subantarctic South Pacific, 6–1,020 m.

Primnoa pacifica Kinoshita, 1907

Primnoa pacifica Kinoshita, 1907:232.—Cairns and Bayer, 2005:233–239, 

figs. 1C, 4–6 (synonymy, key, description).—Stone and Shotwell, 

2007:72, 93, 107, in situ fig. 2.23.—Whitmire and Clarke, 2007:152 

(listed).—Cairns and Bayer, 2009:30 (listed).

Primnoa resaedaformis pacifica.—Heifetz et al., 2005:132.

Remarks. The typical form of Primnoa pacifica was recently described and figured by Cairns and 

Bayer (2005) and thus will not be redescribed herein. It 

differs from the other species in the genus by having prominently spined basal abaxial body wall scales and from 

variety willeyi in having thicker, more robust polyps and 

sclerites on the lateral surface of the polyps. 

This account reports 28 new records of the species, 

but only one additional from the Aleutian Islands (USNM 

1115517), the others being from the Bering Sea, and the 

northeast Pacific coast from Alaska to Washington.

Type Locality. Mochiyama, Sagami Bay, 

depth unknown.

Material Examined. Cobb TA-06-1, 3, 4, 

6, 13, 14, 15, 16, and 17, Stephen’s Passage, Alexander 

Archipelago, 12–27 m, 9 branches, USNM 1099633-

1099636, 1099638-1099642; DW 6-1-1, 3 branches, 

USNM 1130914 and 1130920; DW 6-7-1, 1 branch, 

USNM 1130918; DW 7-1-1, 1 branch, USNM 1130922; 

DW 7-2-1, 1 branch, USNM 1130917; DW 7-4-1, 

3 branches, USNM 1130921; Fiber Optic Monitoring 

Cruise-1159, 3 branches, USNM 1122474 and 1122478; 

Esperanza ZC-7-7-7-5 ( Zhemchug Canyon, Bering Sea), 

1 branch, USNM 1116849; Quest GB-06-1to 5, Glacier 

Bay, 9.5-18 m, 25 branches, USNM 1099628-1099632; 

51°38′18′′N, 179°34′37′′W (Amchitka Pass), 395 m, 3 

branches, USNM 1115517; 59°01′N, 136°10′26′′W, 25 m, 

1 branch, USNM 1115518; 50°41′03′′N, 125°59′48′′W, 

20 m, 28 Jun 2008, 1 dry branch, USNM 1120445; specimens listed by Cairns and Bayer (2005);

Distribution. Sea of Japan, Sea of Okhotsk, 

Aleutian Islands (Amchitka Pass), Bering Sea (Zhemchug 

Canyon), northeast Pacific coast from Alaska to La Jolla, 

California, 9–800 m.

Primnoa pacifica var. willeyi Hickson, 1915

Primnoa willeyi Hickson, 1915:551–553, text fig. 3.

Primnoa pacifica var. willeyi.—Cairns and Bayer, 2005:239, figs. 1C–E, 

7–8 (synonymy, key, description).—Stone and Shotwell, 2007:107 

(listed).—Cairns and Bayer, 2009:30 (listed).

Primnoa resaedformis willeyi.—Heifetz et al., 2005:132.

Remarks. Primnoa pacifica var. willeyi was recently described and figured by Cairns and Bayer (2005) 

and thus will not be redescribed herein. It differs from the 

other species in the genus by having prominently spined 

basal abaxial body wall scales and from the typical form 

in having a more slender polyp and lacking sclerites on 

the lateral surface of the polyps (see Cairns and Bayer, 

2005:228 [key], 252 [table]).

This account reports three new records of the species, 

adding a second record to the Aleutian Islands (also from 

the Rat Islands), documenting the record from Dickins 

Seamount reported by Cairns and Bayer (2005), and a 

third very shallow record (27 m) from Stephen’s Passage, 

Alexander Archipelago.

Types and Type Locality. Syntypes deposited at the BM (1962.7.20.188). Type locality: WSW 

Moresby Island, British Colombia, 183 m.

Material Examined. Alvin 4028 

(Dickins Seamount), 1 branch, USNM 1075478; Cobb
2 0 • smithsonian contributions to zoology

TA-06-09 (Stephen’s Passage), 2 branches, USNM 

1099637; 51°50′26′′N, 179°52′41′′E (Rat Islands), 863 m, 

2 branches, USNM 1115516; specimens listed by Cairns 

and Bayer (2005);

Distribution. Most records are from off British Colombia and the adjacent Alexander Archipelago, 

Alaska, as well as adjacent Dickins Seamount (GOA), but 

also known from Semidi Island (off Alaska Peninsula), and 

from two records in the Aleutian Rat Islands, 27–863 m.

Primnoa wingi Cairns and Bayer, 2005

Primnoa wingi Cairns and Bayer, 2005:243–246, figs. 1h–i, 9-10; 2009: 30 

(listed).—Stone, 2006:233 (listed).—Stone and Shotwell, 2007:90, 107 

(listed).—Cairns and Bayer, 2009:30 (listed).

Remarks. Primnoa wingi was recently described and figured by Cairns and Bayer (2005) and thus 

will not be redescribed herein. Keys to the species within the 

genus are also found in Cairns and Bayer (2005). It differs 

from the other species in the genus by having inconspicuous 

body wall scales that are immersed in polyp tissue, spatulate operculars, and large curved tentacular rodlets. 

This account reports six new records of the species, 

extending its known distribution to the west (to the Near 

Islands), to the east (to Chatham Strait, Alexander Archipelago), and far to the north into the Bering Sea (Zhemchug 

Canyon), as well as broadening its known bathymetric range.

Types and Type Locality. The holotype (USNM 1010257) and paratypes are deposited at 

the USNM. Type locality: Sea Storm-150, 52°30′57′′N, 

173°29′35′′W (north of Amlia Island, Andreanof Islands, 

Aleutians), 213–220 m.

Material Examined. Type material; Esperanza ZC-7-2-ROV-1, 1 branch, USNM 1116847; Jason 

II 2104-11-2, 1 dry colony, AB10-0093; Sea Storm 95, 

1, USNM 1123872; 51°39′24′′N, 179°35′02′′W, 712 m, 

2 branches, USNM 1115519; 57°13′52′′N, 134°04′21′′W, 

110 m, 4 branches, USNM 1115520, and AB01-88, 1 colony; 52°38′59′′N, 172°15′30′′E, 382 m, 1 branch, USNM 

1115521; 51°50′26′′N, 179°52′41′′E, 863 m, 5 branches, 

USNM 1115550.

Distribution. Aleutian Islands from Near Islands to Amukta Pass, Chatham Strait, and the Bering Sea 

(Zhemchug Canyon), 110–914 m.

Genus Arthrogorgia Kükenthal, 1908

Arthrogorgia Kükenthal in Kükenthal and Gorzawsky, 1908:625.—Bayer, 

1996:605.—Cairns and Bayer, 2009:43.

Diagnosis. Colonies with pinnate or dichotomous branching. Polyps arranged in whorls, the polyps 

pointing downward. Polyps protected by 8 keeled operculars (sometimes with small accessory operculars on inner 

side of opercular scales), a pair of buccal and basal scales, 

as well as a variable number of infraopercular scales that 

may or may not be visible in abaxial view. Several transverse rows of strongly curved, unpaired infrabasal scales 

also present. Tentacular rodlets common.

Discussion. Four species occur in the genus 

(Cairns and Bayer, 2009), three of them found in the 

Aleutian Islands. The specific identity of Arthrogorgia sp. 

(Heifetz, 2002; Stone, 2006) is unknown.

Distribution. North Pacific; Japan to Aleutian Islands, 163–1,359 m.

Arthrogorgia utinomii Bayer, 1996

Figure 14a–l

Arthrogorgia utinomii Bayer, 1996:619–628, figs. 13–20.—Wing and Barnard, 2004:24, fig. 17.—Heifetz et al., 2005:132.—Stone and Shotwell, 

2007:106 (listed).—Cairns and Bayer, 2009:31 (listed).

Description. Colonies uniplanar, branching 

equal dichotomous, the distal branches up to 30 cm in 

length. Largest colony (Alb-4781) 30 cm tall. A thin web 

of tissue occurring at each branching axil, which contains 

coenenchymal sclerites. Axis brown to bronze in color; 

colony orange in life.

Polyps arranged in whorls of 7–10, directed downward; 7–9 whorls per 3 cm branch length. Polyps 3.0–3.5 

mm in length.

Each polyp protected by 8 opercular scales, a variable number of infraopercular scales, 1 pair of buccal 

scales, 1 pair of basal scales, and 2–4 transverse rows of 

infrabasal scales. Opercular scales (Figure 14d) triangular 

to tongue shaped but with a somewhat blunt distal tip. 

Abaxial operculars up to 2.0 mm in length, but operculars decreasing in length toward adaxial side, the adaxial 

operculars(Figure 14c) only 0.5 mm in length; L:W = 1.5–

2.0. Inner surface longitudinally keeled, corresponding to 

a longitudinal crease on outer side. Infraopercular scales 

(Figure 14e) square to rectangular in shape, 1–2 pairs occurring proximal to each abaxial, inner, and outer lateral 

opercular (Figure 14a, b, e), these scales 0.5–1.0 mm in 

width, and easily visible just anterior to distal edge of buccal scales. On adaxial side of polyp stand a pair of square 

adaxial buccal scales 0.35–0.65 mm in width, and often 

a series of smaller adaxial body wall scales proximal to 
n u m b e r 6 3 4 • 2 1

those (Figure 14c, j). The two large abaxial buccal scales 

(Figure 14f) as much as 2.2 mm in length, terminating in a 

rounded or moderately pointed distal edge. The two basal 

scales (Figure 14h) as much as 3 mm in height (including 

the spine), each with a thick longitudinally ridged spine on 

their anterolateral edges that may constitute up to 45% 

of length of scale. Proximal to basal scales 2–4 transverse 

rows of thin, curved infrabasal scales (Figure 14b, i), each 

layer consisting of about 4 scales (Figure 14b), giving 

greater flexibility to polyp. Outer surface of all body wall 

and coenenchymal scales covered with radiating lines of 

small spines (Figure 14g). 

Outer layer of coenenchyme composed of elongate, 

irregularly shaped sclerites (Figure 14k) up to 0.65 mm in 

length; inner coenenchymal layer composed of tuberculate 

spheroids 0.09–0.18 mm in diameter. Tentacular rodlets 

cigar shaped, 0.085–0.110 mm in length, and about 0.030 

mm in diameter (Figure 14l). 

Yellow eggs common in many polyps, each egg measuring about 0.65 mm in diameter.

Remarks. This species was well described and 

copiously figured by Bayer (1996), but he did not provide 

images of individual sclerites, even though he had compiled a finished plate of sclerites that for unknown reasons 

was not included in his publication. Images of individual 

sclerites are provided herein, as well as an amplified description, and records of additional specimens. 

Arthrogorgia utinomii is very similar to A. kinoshitai, and the species are sympatric for a large part of their 

ranges. Arthrogorgia utinomii differs in having slightly 

shorter (up to 2.2 mm) buccal scales, which reveal the distal edges of the underlying infraopercular scales on the abaxial, outer, and inner lateral positions; the buccal scales of 

A. kinoshitai, however, are slightly longer (up to 2.5 mm) 

and terminate in an extended lobate edge, both of which 

together form a protective cowl that mostly covers the infraopercular scales. Furthermore, infraopercular scales are 

less common and may even be absent in some polyps of 

A. kinoshitai. Otherwise, the two species are quite similar.

Types and Type Locality. The holotype, 

collected at R/V Starlight 84-1-36, is deposited at the 

USNM (97198). The two other specimens listed by Bayer 

(1996) in the original description are not designated as 

types and thus are not considered as paratypes. Type locality: 52°28′N, 172°30′W (not E as reported by Bayer, 

1996), off Seguam Island, 234 m.

Material Examined. Holotype; nontype specimens reported by Bayer (1996); Arctic Dawn, 

51°32.42′N, 177°45.29′W, 402 m, 25 Dec 1999, dry 

branches, USNM 1006335; Dominator-5159, 1 colony, 

USNM 1009950; Jason II 2104-5-1, branch, USNM 

1115598, and 1 dry colony, AB10-0104; North Pacific, 

52°04.36′N, 176°58.01′E, 384 m, 20 Oct 2000, dry 

branches, USNM 1006329; 52°27.633′N, 175°09.417′E, 

depth unknown, 4 Aug 2005, 1 dry colony, AB05-57.

Distribution. Aleutian Islands from the 

Near Islands to off Amuka Island, 163–882 m, although 

most records from 250 to 700 m.

Arthrogorgia kinoshitai Bayer, 1952

Figure 15 a–j

Calyptrophora ijimai.—Broch, 1935:26, figs. 15–16; 1940:12, 20.

Arthrogorgia kinoshitai Bayer, 1952:64–65, pl. 2: figs. 1–8, pl. 3: figs. 1–12; 

1996:606–619, figs. 5–12.—Heifetz, 2002:26 (listed).—Cairns and 

Bayer, 2009:31 (listed).

Arthrogorgia kinoshita.—Heifetz et al., 2005:132, 136.—Wing and Barnard, 

2004:24 (listed).—Stone and Shotwell, 2007:106 (listed).

?Arthrogorgia ijimai.—Stone and Shotwell, 2007:106 (listed).

Description. See Remarks.

Remarks. This species was well described by 

Bayer (1952) and profusely illustrated by him (Bayer, 

1996) using SEM, but SEM-quality images of individual 

sclerites have never been published and are provided 

herein. Four new records are also reported.

The species is so similar to A. utinomii that it would be 

redundant to describe it in full. It is illustrated herein and 

distinguished from A. utinomii in the account of that species. The colony from Jason II 2107-5-1 displays extreme 

webbing of the coenenchyme between its basal branches.

Types and Type Locality. The holotype, 

collected at Alb-4781, is deposited at the USNM (49978)

and consists of one colony preserved in alcohol, SEM stubs 

B2499-2500, 2548, C 1414-17, and two “exploded” polyps, all the sclerites of which have been glued to a slide 

by F. M. Bayer to approximate their relative position in 

the polyp. Type locality: 52°14′30′′N, 174°13′E (west of 

Agattu Island, Near Islands), 882 m.

Material Examined. Holotype (USNM 

49978) and specimens (USNM 100748-9, SEM stubs B2510, 

2125, 2143-47) reported by Bayer (1996); Delta 5993-7C-1, 

1 dry colony, AB10-0020; Jason II 2103-5-1, 1 branch, 

USNM 1115594, and 1 dry colony, AB10-0117; Ocean 

Olympic, 52°11′10′′N, 176°17′00′′E, 220 m, 20 Oct 2000, 

1 dry colony, USNM 1006150; Pacific Knight 941-204, 1 

dry colony, USNM 100812 (from same station reported by 

Bayer, 1996); coll. by Renfro, 51°41′46′′N, 178°20′30′′W, 

280 m, 8 Mar 2000, 1 dry colony, USNM 1006248.
2 2 • smithsonian contributions to zoology

Distribution. Sea of Okhotsk (Broch, 1935), 

western Aleutian Islands from Near Islands to Tanaga 

Pass, 220–1,309 m.

Arthrogorgia otsukai Bayer, 1952

Figure 16 a–j

Arthrogorgia otsukai Bayer, 1952:65–69, pl. 2, figs. 9–12, pl. 3, figs. 13–27; 

1996:606, figs. 3–4.—Heifetz, 2002:26 (listed).—Wing and Barnard, 

2004:24 (listed).—Heifetz et al., 2005:132.—Stone and Shotwell, 

2007:106 (listed).—Cairns and Bayer, 2009:31 (listed).

Description. Colonies uniplanar, branching 

equal dichotomous, the distal branches 4–15 cm in length. 

Largest colony (Jason II 2102-4-1) only 20 cm in height 

and 10 cm width. Axis dark brown.

Polyps arranged in whorls of 5–7, directed downward; 

11–13 whorls per 3 cm branch length. Polyps 2.3–2.75 

mm in length.

Each polyp protected by 8 opercular scales (an occasional accessory opercular scale), a variable number of 

infraopercular scales, 1 pair of buccals and basals, and 

4–5 transverse rows of infrabasal scales. Opercular scales 

triangular (Figure 16d), but with an elongate, cylindrical, 

distal blunt spine having numerous prominent, parallel, 

serrate ridges. Operculars decreasing progressively in size 

toward axial side (Figure 16c), the abaxials as much as 

1.5 mm in length, the adaxials only 0.6–0.7 mm in length; 

L:W = 2.0–2.7. Occasionally, much smaller infraoperculars occurring on inner side of adaxial and inner lateral 

opercular scales (Figure 16c). Infraopercular scales (i.e., 

marginals and submarginals) square to rectangular in 

shape, 0–1 pair occurring proximal to each abaxial opercular (Figure 16a, g), 1–2 pairs adjacent to outer and inner 

lateral operculars, and 4–6 pairs on adaxial face (Figure 

16c). These scales often visibly protruding from beneath 

distal margin of covering buccal scale (Figure16a). Adaxial infraoperculars as much as 0.5 mm in width, others as 

much as 0.8 mm in width. The two large abaxial buccal 

scales (Figure 16b, e) as much as 1.4 mm in length terminating in a gently rounded margin. The two basal scales 

(Figure 16f) rarely more than 1 mm in height, terminating 

in a small distal lobe or nub, not a spine. Infrabasal scales 

(Figure 16a, h) not arranged in pairs but in 4 or 5 transverse rows of 4 or 5 thin, curved scales. Outer surface of 

all scales covered with radiating lines of small spines.

Coenenchymal scales (Figure 16i) thin, flat, polygonal scales as much as 0.5 mm in width. Coenenchymal 

webbing present between basal branches of colony from 

Jason II 2102-4-1. Tentacular rodlets (Figure 16j) 0.10–

0.13 mm long and 0.030 mm in width, usually slightly 

flattened.

Remarks. This species was well described by 

Bayer (1952) and partially illustrated by him (Bayer, 1996) 

using SEM, but SEM-quality images of individual sclerites 

and various polyp views have never been published and 

are provided herein. One new record is reported here; it is 

the only record known having a specific locality.

Arthrogorgia otsukai is distinguished from other Aleutian congeners by lacking spines on its basal scales and in 

having smaller polyps. It also has multiridged opercular 

tips (ridges occurring even on the outer surface), flattened 

tentacular rodlets, and lacks coenenchymal tuberculate 

spheroids.

Types and Type Locality. The types (16 

colonies) are deposited at the USNM (49979) and also 

includes SEM stubs B2561-2, C1419, 1420, and an “exploded” polyp, all the sclerites of which have been glued 

to a slide by F. M. Bayer to approximate their relative 

position in the polyp. Although Bayer (1952) referred to a 

holotype, no particular specimen in the type lot was segregated or specified in the text as the unique type; thus all 

16 colonies are considered to be syntypes, as implied by 

Bayer (1996). Type locality: Somewhere between Aangan 

River, Kamchatka and Bowers bank, depth and Albatross

station unknown.

Material Examined. Syntypes; Jason II

2101-2-1, 1 branch, USNM 1115557, and 1 dry colony, AB10-0111; Jason II 2102-4-1, 1 branch, USNM 

1115597 and AB10-0112; Jason II 2102-4-2, 2 branches, 

USNM 1115595, and 1 dry colony, AB10-0118.

Distribution. Amchitka Pass and south of 

Kanaga Island, Aleutian Islands, 1,359–1,372 m. The type 

locality is too generalized to be of use.

Genus Calyptrophora Gray, 1866

Calyptrophora Gray, 1866:25.—Bayer, 2001:367–368.—Cairns, 2007b: 

527.—Cairns and Bayer, 2009:44–45, figs. 15i–n.—Cairns, 2009:420–

426 (key to species).

Diagnosis. Colonies uniplanar, branched or 

unbranched. Polyps arranged in whorls, facing upward in 

most species, but downward in some. Polyps composed 

of 2 annular sclerite rings (basal and buccal), each composed of 2 inseparably fused scales; crescent-shaped infrabasals also present. Buccal and basal scales often bearing 
n u m b e r 6 3 4 • 2 3

distal cylindrical spines or flattened teeth. Opercular scales 

keeled. Coenenchymal scales granular, sometimes ridged. 

Tentacular rodlets often present.

Discussion. The 16 valid species in the genus 

were listed, keyed, and phylogenetically analyzed by 

Cairns and Bayer (2009).

Distribution. Pacific Ocean and western Atlantic, 229–3,531 m.

Calyptrophora laevispinosa Cairns, 2007

Figure 2e

Calyptrophora laevispinosa Cairns, 2007b:536–537, figs. 1G, 14–15.—

Cairns and Bayer, 2009:31 (listed).

Remarks. This species was recently described 

and illustrated, and thus little can be added to its characterization; however, a picture of a large colony (40 cm in 

height, Figure 2e) is provided and six additional localities 

are provided, extending its known distribution throughout the Gulf of Alaska and to a depth of 3,531 m, the 

greatest depth recorded for a species of Calyptrophora.

The specimens from Alvin 3802 and 4037 are identical to C. laevispinosa, differing only in having ridged 

basal spines. Ironically, this was the character used to key 

various species (Cairns and Bayer, 2009) and to name the 

species (Cairns, 2007b), and thus it may be necessary to 

re-evaluate the consistency of this character among the 

Calyptrophora species or to reevaluate this particular 

specimen, perhaps using molecular analyses.

Galls containing a vermiform parasite are common in 

holotype and USNM 1075477.

Types and Type Locality. The holotype is 

deposited at the USNM (1102462), and a piece is also at 

the CAS (175310). Type locality: 46.683°N, 126.782°W 

(Gorda 1996 Flow off Washington), 3,107 m.

Material Examined. Alvin 3802, 8 

branches, USNM 1082615-1082617; Alvin 4033, 1 

branch, USNM 1075472; Alvin 4037, 1 large colony and 

several branches, USNM 1075473-1075475; Alvin 4041, 

1 dry colony and 1 alcohol preserved branch, USNM 

1075476 and 1075477, respectively; Jason II-93-18, 1 

dry colony, USNM 1081195; Jason II-93-23, 1 dry colony 

and 1 alcohol-preserved branch, USNM 1081196.

Distribution. Gorda Flow off Washington, 

GOA (Derickson, Patton, Giacomini, and Welker Seamounts), 2,672–3,531 m. Not known from the Aleutian 

Islands, rather only from the closely adjacent GOA.

Genus Parastenella Versluys, 1906

Stenella (Parastenella) Versluys, 1906:39, 45.

Candidella (Parastenella).—Bayer, 1956:F222.

Parastenella.—Bayer, 1961:295.—Cairns, 2007a:245–247; 2007b:518 (tabular key).—Cairns and Bayer, 2009:45–46.—Cairns, 2010:434 (key to 

species).

Diagnosis. Colonies flabellate or slightly 

bushy, dichotomously or irregularly branched, usually 

fairly stiff. Polyps occurring in whorls, pairs, or individually, oriented perpendicular to branch. Body wall scales 

arranged in 5–8 longitudinal rows, the 8 marginal scales 

elongate and often (most commonly those on abaxial side) 

extending as a semicircular to tubular flute; each marginal 

flute bearing a concentrated patch of nematocysts (a nematocyst pad) on their distal inner side. Opercular scales 

offset in alignment from marginal scales. Opercular scales 

keeled on inner surface. Tentacular rodlets common.

Discussion. Seven species are known in the 

genus (Cairns and Bayer, 2009; Cairns, 2010), which are 

keyed and discussed in the papers cited above. The genus 

is easily distinguished by having its opercular scales offset 

from its marginals (see below). Cairns (2010) suggested 

that marginal nematocyst pads were characteristic of the 

genus, as he reported them from P. bayeri and P. gymnogaster. They are herein confirmed from P. doederleini and 

P. ramosa, and are retrospectively noted in P. atlantica

(see Cairns, 2007a:fig. 3). Thus they need only be confirmed in P. pacifica and P. spinosa. 

In most primnoids, there are two rows of abaxial 

scales that cover the abaxial side of the polyp, the adjacent abaxial edges of these scales abutting or overlapping along the sagittal axis of the polyp, which is also 

the case in Parastenella. In most primnoid genera a set 

of eight opercular scales are in directly alignment with 

the body wall scales; however, in Parastenella the opercular scales are uniquely offset from the marginal scales 

such that each opercular scale is aligned with the space 

between two adjacent marginals (see Cairns, 2007a:fig. 5, 

and Figure 17c herein) . This results in one large abaxial 

opposing a small adaxial scale and aligned with the sagittal axis, and three couples of lateral opercular scales that 

are paired across the sagittal axis. These three pairs may 

be termed the outer lateral, transverse lateral, and inner 

lateral pairs.

Distribution. Known from all ocean basins 

except eastern Atlantic and off continental Antarctica, 

330–3,470 m.
2 4 • smithsonian contributions to zoology

Parastenella doederleini

(Wright and Studer, 1889)

Figures 2b, 17a–j

Stenella doederleini Wright and Studer, 1889:58, pl. 20, fig. 7.—Not Studer, 

1894:64 (=P. ramosa).—Kükenthal, 1915:152 (key to species); 1919: 

446–447; 1924:304–305 (key to species).—Aurivillius, 1931:290–291.

Stenella (Parastenella) doederleini.—Versluys, 1906:45–47, pl. 1 fig. 3, textfigs. 49–54.—Kinoshita, 1908b:28–30, pl. 2, fig. 12, text figs. 4–7.

Candidella (Parastenella) doederleini.—Bayer, 1956:F222, fig. 159-4a.

Parastenella doederleini.—Bayer, 1981:934, 936, fig. 69.—Cairns, 

2007b:518–519 (tabular key), 523, fig. 4B.—Whitmire and Clarke, 

2007:152 (listed).—Cairns and Bayer, 2009:31, 45–46, fig. 16a–b.—

Cairns, 2010:435 (key to species).

Description. Colonies bushy, dichotomously 

branched, and rather small, the largest Aleutian colony 

only 6 cm in height (Figure 2b), whereas other colonies 

reported to be (Aurivillius, 1931) as much as 25 cm in 

height. Axis dark brown, usually visible through single 

thin, translucent white layer of coenenchymal scales. Polyps occurring singly, in pairs, and occasionally in whorls 

of 3, projecting perpendicular to branch surface, slightly 

clavate, and 2.1–2.5 mm in length. 

Each polyp covered with 8 opercular scales, 8 marginal 

scales, and 5or 6 longitudinal rows of body wall scales. As 

is typical for the genus, 1 pair of opposing symmetrically 

shaped abaxial and adaxial opercular scales separated by 3 

pairs of asymmetrical lateral opercular scales (Figure 17c). 

Whereas length of the operculars (Figure 17e) not varied 

much (0.71–0.92 mm), the basal width increasing progressively from adaxial to abaxial side, such that abaxials L:W 

= 1.7 and adaxials about 2.1. Outer surface of operculars 

deeply longitudinally creased and covered with tiny spines; 

inner surface prominently keeled. The 8 marginal scales 

forming an asymmetrical rosette (Figure 17c) when viewed 

from above, the 5 marginals on abaxial and lateral sides 

bearing elongate flutes, whereas the 3 on adaxial side lacking flutes (Figure 17c). Fluted marginals (Figure 17d) consisting of a broad rectangular base supporting a projecting 

semi-circular to tubular flute, the flutes quite narrow 

(0.10–0.15 mm or 1⁄4 width of the base), 50–55% length 

of the scale, bearing a nematocyst pad in their distal inner 

surface (Figure 17c, j). Nematocysts closely packed, each 

uniformly 0.021 mm in length and 0.003 mm in diameter. 

Nonfluted adaxial marginals with a rounded distal margin. Body wall scales arranged in roughly 6 rows: 2 abaxial 

rows of 5 or 6 scales, 2 short lateral rows of 2 or 3 scales, 

and 2 adaxial rows of 5 or 6 scales, the latter (Figure 17f) 

roughly square (0.35–0.45 mm in width) and completely 

covering adaxial polyp surface. Abaxial body wall scales 

(Figure 17g) proximal to marginals also bearing flutes, but 

progressively smaller toward base of polyp.

Coenenchymal scales (Figure 17h) irregular in shape, 

up to 0.55 mm in length, flat, and often bearing a boss or 

ridge on center of outer surface. Tentacular rodlets (Figure 17i) cigar shaped, 0.070–0.082 mm in length (L:D = 

5.1–5.8).

Remarks. Parastenella doederleini is quite 

similar to P. ramosa as is discussed by Cairns (2007b) by 

narrative and tabular key and by Cairns (2010) in a key to 

the species in the genus. In summary, P. doederleini is distinguished by having smaller, more delicate polyps; more 

slender and shorter marginal and submarginal flutes; and 

by having a tendency to have bosses or ridges on its coenenchymal scales. Their distributions do not quite overlap in that, whereas P. ramosa in known from the Gulf 

of Alaska and the closely adjacent Commander Islands to 

the west, it has not yet been collected from the Aleutian 

Islands; P. doederleini also tends to be found in deeper 

water than P. ramosa. Parastenella doederleini is the type 

of the genus and has been redescribed several times based 

on new material by Versluys (1906), Kinoshita (1908b), 

Kükenthal (1919), and Aurivillius (1931). It is a new record for Alaska and the Aleutian Islands.

The presence of nematocysts pads on the inner surface 

of the marginal scales is confirmed in this species (Figure 

17c,j), as suggested might be the case for all the species 

within the genus (Cairns, 2010).

Types and Type Locality. The holotype is 

deposited at the BM (1889.5.27.37) and a fragment also 

at the ZMA (Coel. 3085, see van Soest, 1979). Type locality: Challenger-237, 34°37′N, 140°32′E (off Yokohama, 

Japan), 3423 m.

Material Examined. Jason II 2098-2-2, 

branch, USNM 111556; Jason II 2100-3-2, branches, 

USNM 1115561; Jason II 2100-3-3, branch, SEM stubs 

C1478-1481, USNM 1115560.

Distribution. Indonesia, Japan, Aleutian Islands (Adak Canyon), 400–3,423 m.

Parastenella ramosa (Studer, 1894)

Stenella ramosa Studer, 1894:64–65.

Stenella doederleini.—Studer, 1894:64.

Stenella (Parastenella) ramosa.—Versluys, 1906:47–48.—Kükenthal, 

1919:445; 1924: 303.
n u m b e r 6 3 4 • 2 5

Parastenella ramosa.—Cairns, 2007b:518–523, figs. 1E, 4, 5.—Cairns and 

Bayer, 2009:31 (listed).

Remarks. Parastenella ramosa has recently 

been described and figured by Cairns (2007b), and thus 

will not be redescribed herein. Keys to the species within 

the genus are found in Cairns (2007b, 2010), and comparison to P. doederleini is made above. This account reports six new records of the species, extending its known 

distribution west to the Commander Islands, Russia, and 

Adak Canyon in the Aleutian Islands.

The presence of nematocysts pads in the inner surface 

of the marginal scales is confirmed in this species, as suggested might be the case for all the species within the genus 

(Cairns, 2010).

Types and Type Locality. Holotype 

lost (see Cairns, 2007b). Type locality: Albatross 3384: 

7°31′30′′N, 79°14′W (Gulf of Panama), 837 m.

Material Examined. Alvin 3798, branch, 

USNM 1082596; Alvin 3799, branch, USNM 1082601; 

Alvin 3806, branches, USNM 1082639, -2633, -2641; 

Alvin 4028, branch, USNM 1075358; Jason II 2099-18-1, 1 

dry colony, AB10-0074; R/V Keldysh (Mir 2) 22, branches, 

USNM 1106689; specimens reported by Cairns (2007b).

Distribution. Gulf of Panama, various seamounts off western USA and Canada (e.g., Rodriguez, 

Davidson, Axial 1998 South Flow, Welker, and Dickins), 

Gulf of Alaska (Giacomini, Pratt, Murray, and Warwick 

Seamounts), Pip Volcano (Commander Islands, off Kamchatka, Russia), Adak Canyon, 665–1,766 m.

Parastenella gymnogaster Cairns, 2007

Parastenella gymnogaster Cairns, 2007b:523–526, figs. 1B, 6–7.—Cairns 

and Bayer, 2009:31 (listed).

Remarks. Parastenella gymnogaster has recently been described and figured by Cairns (2007b) and 

thus will not be redescribed herein. Keys to the species 

within the genus are found in Cairns (2007b, 2010). In 

summary, it differs from other species in having broad, 

short marginal flutes, only six rows of body wall scales 

(the scales of the abaxial rows being quite broad and linearly arranged), ornately ridged coenenchymal scales, and 

a naked adaxial polyp face. This account reports three new 

records of the species from three additional seamounts in 

the Gulf of Alaska, although it is not known from the 

Aleutian Islands. 

Types and Type Locality. The holotype 

and paratypes are deposited at the USNM (holotype = 

1075464). Type locality: Alvin 4033-23, 54°59′01′′N, 

140°23′59′′W (Welker Seamount off British Columbia), 

2,634 m.

Material Examined. Types; Alvin 3803, 

branches, USNM 1082619; Alvin 3804, branches, USNM 

1082620, -2624, -2626, -2628, -2630, -2632; Alvin 4029, 

branches, USNM 1075359-1075361, 1075363.

Distribution. Juan de Fuca Ridge off Oregon and seamounts off Oregon and British Columbia, 

Gulf of Alaska (Derickson, Chirikof, Marchand Seamounts), 1,962–2,869 m.

n u m b e r 6 3 4 • 2 7

FIGURES 1–17

FIGURE 1. Colonies of various species: (a) Thouarella cristata, holotype (USNM 1116600), dry; (b) T. trilineata, holotype 

(USNM 1010175), in alcohol; (c) Plumarella hapala, holotype (USNM 1006310), dry; (d) P. aleutiana, holotype (USNM 

1011356), dry; (e) P. profunda, holotype (USNM 1134074), dry; (f) P. spicata, holotype (USNM 30050), in alcohol. Scale bars: 

a–c, 5 cm; d, 10 cm; e–f, 2.5 cm.
2 8 • smithsonian contributions to zoology

FIGURE 2. Colonies of various species: (a) Plumarella echinata, holotype (USNM 1135009), in alcohol; (b) Parastenella doederleini, Jason IV 2100-3-3, USNM 1115560, in alcohol; (c) Plumarella superba, Delta 5604 (USNM 1011277), dry; (d) P. nuttingi, holotype (USNM 1006317), dry; (e) Calyptrophora laevispinosa, Alvin 4037 (USNM 1075475), in alcohol; (f) Plumarella 

robusta, holotype (USNM 1135992), dry. Scale bars: a,f, 2.5 cm; b, 0.5 cm; c–e, 5 cm.
n u m b e r 6 3 4 • 2 9

FIGURE 3. Polyps and scales of Thouarella cristata from the holotype (USNM 1115600), a–d are stereo views: (a) lateral view 

of branchlet showing randomly arranged polyps; (b–d) abaxial, adaxial, and opercular views of polyps, respectively; (e) opercular scales; (f) marginal scales; (g) abaxial body wall scales; (h) coenenchymal scales; (i) adaxial body wall scales. Scale bars: a, 

1 mm; b–d, 0.5 mm; e–i, 0.25 mm.
3 0 • smithsonian contributions to zoology

FIGURE 4. Polyps and scales of Thouarella trilineata from the holotype (USNM 1010175), a–c are stereo views: (a) lateral view 

of branchlet showing randomly arranged polyps; (b–c) two opercular views of polyp, also showing adaxial polyp face; (d) adaxial side of polyp (m = marginal scales); (e) lateral view of polyp; (f) opercular scales; (g) marginal scales; (h) submarginal scales; 

(i) body wall scales; (j) coenenchymal scales. Scale bars: a, 1 mm; b–e, 0.5 mm; f–j, 0.25 mm.
n u m b e r 6 3 4 • 3 1

FIGURE 5. Polyps and scales of Plumarella spicata from the holotype (USNM 30050), a–d are stereo views: (a) adaxial view of 

polyp; (b) lateral view of polyp; (c) adaxial view of polyp; (d) opercular view of polyp; (e) opercular scales; (f) marginal scales; 

(g) body wall scales; (h) outer surface of coenenchymal scales, in situ. Scale bars: a–d, 0.5 mm; e–g, 0.25 mm; h, 0.10 mm.
3 2 • smithsonian contributions to zoology

FIGURE 6. Polyps and scales of Plumarella profunda from the holotype (USNM 1134074), a–b are stereo views: (a–b), lateral 

and abaxial views of polyps; (c) opercular scales; (d) body wall scales; (e) coenenchymal scales; (f) tentacular rodlets. Scale bars: 

a–b, 0.5 mm; c–e, 0.10 mm; f, 0.025 mm.
n u m b e r 6 3 4 • 3 3

FIGURE 7. Polyps and scales of Plumarella hapala from the holotype (USNM 1006310), a–c are stereo views: (a) lateral view of 

branchlet showing randomly arranged polyps; (b–c) adaxial view of polyps; (d) lateral view of polyp; (e) lateral view of gravid 

polyp; (f) opercular scales; (g) marginal scales; (h) smooth inner surface of a marginal scale, in situ; (i) body wall scales; (j) a 

tentacular rodlet; (k) a coenenchymal scale. Scale bars: a, 1 mm; b–e, 0.5 mm; f–g, i, k, 0.25 mm; h, 0.10 mm; j, 0.025 mm.
3 4 • smithsonian contributions to zoology

FIGURE 8. Polyps and scales of Plumarella aleutiana from the holotype (USNM 1011356), a–d are stereo views: (a–b) lateral 

and abaxial views of polyps; (c–d) adaxial opercular views of two polyps; (e) opercular scales; (f) submarginal scale; (g) marginal 

scale; (h) inner surface of several smooth-tipped marginal scales, in situ; (i) body wall scales; (j) inner surface of coenenchymal 

scales, in situ; (k) coenenchymal tuberculate spheroids. Scale bars: a, c, 1.0 mm; b, d, h, 0.5 mm; e–g, i, 0.10 mm; j, 0.25 mm; 

k, 0.025 mm.
n u m b e r 6 3 4 • 3 5

FIGURE 9. Polyps and scales of Plumarella superba (a, e, g–l, holotype, USNM 30691; b, f, USNM 1009951; c–d, USNM 

50150), a-d are stereo views: (a) abaxial view of a typical polyp with smooth scales; (b) abaxial view of a polyp with ridged 

body wall scales; (c–d) adaxial and opercular views of polyps, respectively; (e) typical opercular scales with smooth outer face; 

(f) variant opercular scales with ridged outer surface; (g) abaxial marginal scales; (h) adaxial marginal scale; (i) submarginal 

scale; (j) smooth body wall scales; (k) adaxial body wall scales; (l) coenenchymal scales. Scale bars: b–d, 0.5 mm; a, e–l, 0.25 mm.
3 6 • smithsonian contributions to zoology

FIGURE 10. Polyps and scales of Plumarella nuttingi from the holotype (USNM 1006317), a–d, k are stereo views: (a, c) lateral 

and adaxial view of polyps, respectively; (b–d) opercular view of polyps; (e) opercular scales; (f) marginal scales; (g) inner face of 

a marginal scale; (h) body wall scales; (i) coenenchymal scales in situ in web; (j) tentacular rodlets; (k) tentacular rodlets, in situ. 

Scale bars: a–d, k, 0.5 mm; e–i, 0.25 mm; j, 0.05 mm.
n u m b e r 6 3 4 • 3 7

FIGURE 11. Polyps and scales of Plumarella echinata from the holotype (USNM 1135009), a–d are stereo views: (a) lateral 

view of branchlet showing randomly arranged polyps; (b) view of outer lateral face of a polyp; (c–d) adaxial and opercular 

views of polyps, respectively (m = marginal scales); (e) opercular scales; (f) marginal scales; (g) spination on a marginal spine; (h) 

submarginal scale; (i) abaxial body wall scales; (j) adaxial body wall scales; (k) coenenchymal scales. Scale bars: a, 1 mm; b–d, 

0.5 mm; e–f, h–k, 0.25 mm; g, 0.05 mm.
3 8 • smithsonian contributions to zoology

FIGURE 12. Polyps and scales of Plumarella robusta from the holotype (USNM 1135992), a–d are stereo views: (a) lateral view 

of branchlet showing randomly arranged polyps; (b) abaxial view of a polyp; (c–d) adaxial opercular view of two polyps (m = 8 

marginal scales); (e) opercular scales; (f) marginal scales; (g) spination on a marginal spine; (h) coenenchymal scales; (i) body wall 

scales; (j) tentacular rodlets. Scale bars: a, 1 mm; b–f, h–i, 0.25 mm; g, 0.05 mm; j, 0.025 mm.
n u m b e r 6 3 4 • 3 9

FIGURE 13. Polyps and sclerite of Fanellia compressa (a, c–d, h, USNM 57543; b, USNM 57542), a–d are stereo views: (a–b) 

oblique view of two polyps showing outer lateral body wall scales with lateral extensions (ol = outer lateral scales); (c) opercular 

view of a polyp whorl; (d) adaxial view of a polyp (il = inner lateral scales, ol = outer lateral scales, ad = adaxial scales); (h) coenenchymal tuberculate spheroid. Polyps and sclerite of Fanellia fraseri (e–g, i, USNM 51284), e–g are stereo views: (e) adaxial 

and lateral views of polyps; (f) adaxial opercular view of a polyp; (g) opercular view of a polyp whorl; (i) coenenchymal tuberculate spheroid. Scale bars: a, d–f, 0.25 mm; b–c, g, 0.5 mm; h, 0.01 mm; i, 0.025 mm.
4 0 • smithsonian contributions to zoology

FIGURE 14. Polyps and scales of Arthrogorgia utinomii (a, f–h, paratype, USNM 80829; b–e, i–k, paratype, USNM 58168; 

l, holotype), a–c are stereo views: (a–b) lateral view of two polyps showing infraopercular scales; (c) adaxial view of a polyp; 

(d) opercular scales; (e) infraopercular scale; (f) buccal scales; (g) spination on outer surface of body wall scale; (h) basal scales; 

(i) infrabasal scales; (j) adaxial body wall scale; (k) coenenchymal scales; (l) tentacular rodlets. Scale bars: a–b, 1 mm; c, f, 

0.5 mm; d–e, h–k, 0.25 mm; g, 0.01 mm; l, 0.025 mm.
n u m b e r 6 3 4 • 4 1

FIGURE 15. Polyps and scales of Arthrogorgia kinoshitai (a, c, Pacific Knight 941-204, USNM 100748; b, d–j, holotype, 

USNM 49978), a–c are stereo views: (a) lateral view of a polyp showing infrabasal and accessory infraopercular scales; (b) opercular view of a polyp whorl; (c) adaxial view of a polyp showing small adaxial body wall scales and other accessory infraopercular scales; (d) buccal scales; (e) infraopercular scale; (f) opercular scales; (g) basal scales; (h) coenenchymal scales; (i) infrabasal 

scale; (j) tentacular rodlet. Scale bars: a, 1 mm; b, 2 mm; c–d, g, f, 0.5 mm; e, h–i, 0.25 mm; j, 0.01 mm.
4 2 • smithsonian contributions to zoology

FIGURE 16. Polyps and scales of Arthrogorgia otsukai from a syntype colony (USNM 49979), a–c are stereo views: (a) lateral 

view of polyp showing both infrabasal and infraopercular scales; (b) abaxial buccal view of a polyp; (c) adaxial polyp view; 

(d) opercular scales; (e) outer surface of a buccal scale; (f) basal scales; (g) outer lateral infraopercular scales; (h) infrabasal scales; 

(i) coenenchymal scale; (j) tentacular rodlets. Scale bars: a–c, 0.5 mm; d–i, 0.25 mm; j, 0.025 mm.
n u m b e r 6 3 4 • 4 3

FIGURE 17. Polyps and scales of Parastenella doederleini from Delta J2100-3-3 (USNM 1115560), a–c are stereo pairs: 

(a) outer lateral view of a polyp; (b) adaxial view of a polyp; (c) opercular view of a polyp (adaxial side on right); (d) fluted marginal scales; (e) opercular scales; (f) adaxial body wall scales; (g) abaxial body wall scale; (h) coenenchymal scales; (i) tentacular 

rodlets; (j) clustered nematocysts of a nematocyst pad on inner side of a marginal flute. Scale bars: a–c, 0.5 mm; d, 0.1 mm; e–h, 

0.25 mm; i, 0.025 mm; j, 0.05 mm. 

Details on all the stations where specimens for this study were recorded are 

provided in Table A.1.

Appendix: Station List
4 6 • smithsonian contributions to zoology

TABLE A.1. List of data stations, their locations, and date data were recorded at each. A question mark (?) indicates lack of information.

Research vessel 

and station no. °N Latitude ° Longitude Depth (m) Date

Alaskan Leader

35 53°04′ 170°12′W ? 4 Jun 2002

35-2 53°03.7′ 170°12′W 182–630 4 Jun 2002

Albatross (Alb)

3319 53°40′30′′ 167°30′00′′W 108 18 Jun 1890

3330 54°00′45′′ 166°53′50′′W 642 21 Jun 1890

3331 54°01′40′′ 166°48′50′′W 840 21 Jun 1890

3480 52°06′00′′ 171°45′00′′W 518 8 Jul 1893

3500 56°02′00′′ 169°30′00′′W 221 17 Jul 1893

3501 55°51′00′′ 169°18′00′′W 1,258 17 Jul 1893

4769 54°30′40′′ 179°14′00′′E 433–446 3 Jun 1906

4771 54°30′00′′ 179°17′00′′E 779 4 Jun 1906

4777 52°11′00′′ 179°49′00′′E 79–95 5 Jun 1906

4778 52°12′00′′ 179°52′00′′E 60–79 5 Jun 1906

4779 52°11′00′′ 179°57′00′′W 99–102 5 Jun 1906

4781 52°14′30′′ 174°13′00′′E 882 7 Jun 1906

Alvin

3798 53°53′34′′ 148°31′56′′W 855 27 Jun 2002

3799 53°59′32′′ 148°30′14′′W 1,308 28 Jun 2002

3802 54°31′50′′ 150°18′13′′W 1,778–1,993 1 Jul 2002

3803 54°48′31′′ 152°55′44′′W 2,869 2 Jul 2002

3804 54°56′50′′ 151°19′11′′W 2,417 5 Jul 2002

3806 48°05′21′′ 132°50′38′′W 815 10 Jul 2002

4028 54°33.17′ 136°50.64′W 755 4 Jun 2004

4029 54°37.18′ 136°42.54′W 2,704–2,789 5 Jun 2004

4033 54°59.24′ 140°23.99′W 2,635–2,665 9 Jun 2004

4037 56°22.18′ 142°25.62′W 2,783 13 Jun 2004

4041 56°21.03′ 146°22.82′W 2,803 17 Jun 2004

Cobb, John

TA-6-1 and 9 57°53′39′′ 133°20′12′′W 27 22 Mar 2006

DW (Deep Worker)

6-1-1 54°37.55′ 130°57.0′W 247 19 Jun 2009

6-7-1 52°31.36′ 131°23.16′W 351 20 Jun 2009

7-1-1 54°37.84′ 130°57.05′W 242 19 Jun 2009

7-2-1 51°45.26′ 130°44.24′W 500 12 Jun 2009

7-4-1 52°31.32′ 131°22.33′W 320 16 Jun 2009

Delta

5E-2-5, 7, 9 51°37.322′ 177°14.35′W 155 ?

5590 51°43′22′′ 176°29′10′′W 85 13 Jul 2002

5591 51°47′29′′ 176°15′34′′W 79 13 Jul 2002

5592 51°47′29′′ 176°15′34′′W 165 13 Jul 2002

5597 52°45′18′′ 179°19′47′′W 145 15 Jul 2002

5997-8C-1 51°21.822′ 179°29.07′W 185 ?

5600 52°32.7351′ 177°33.7975′E 87 15 Jul 2002

5604 52°13′27′′ 179°53.17′W 40 16 Jul 2002

5989-14-B-1 51°57.094′ 176°50.3431′W 134 1 Jul 2003

5993-7C-1,4,5,6 51°24.082′ 178°34.479′W 100 3 Jul 2003
n u m b e r 6 3 4 • 4 7

TABLE A.1. continued

Research vessel 

and station no. °N Latitude ° Longitude Depth (m) Date

6004-11C-1,6 52°45.913′ 179°19.701′W 154 6 Jul 2003

6005-11B-1 51°43.986′ 179°19.188′W 120 6 Jul 2003

6007-13C-2 51°55.214′ 177°28.281′W 150 7 Jul 2003

6199-5E-1,5,7,8,9,13,14,15 51°37.117′ 177°13.853′W 135–175 26 Jun 2004

6202-14D-4 51°57.715′ 176°50.004′W 165 27 Jun 2004

6203-18-1 51°52.226′ 176°15.996′W 138 28 Jun 2004

6206-18-2,14 51°51.97′ 176°15.13′W 98–100 29 Jun 2004

6007-13C-2 51°55.214′ 177°28.281′W 150 ?

6208-21D-5 51°51.97′ 176°15.13′W 100 30 Jun 2004

6209-21C-1,2 51°43.178′ 176°17.944′W 100 30 Jun 2004

6209-21C-4 51°43.582′ 176°17.540′W 95 30 Jun 2004

6209-21C-7 51°43.983′ 176°17.021′W 52 30 Jun 2004

6219-3B-2 51°52.374′ 175°15.695′W 154 5 Jul 2004

6221-2B-1 51°54.117′ 173°52.462′W 142 6 Jul 2004

6221-2B-9 51°54.819′ 173°53.139′W 119 6 Jul 2004

6221-2D-1 51°56.744′ 173°55.306′W 82 6 Jul 2004

Dominator

1997-1-39 53°01′11′′ 170°18′49′′W 202 19 Jun 1997

1997-1-49 52°22.45′ 170°34.25′W 231–277 21 Jun 1997

1997-1-78 51°58′40′′ 172°35′24′′W 173 28 Jun 1997

2000-1-156 51°51′01′′ 178°25′57′′E 388–395 1 Jul 2000

5159 52°26′02′′ 179°56′29′′E 170 21 Jul 2000

Esperanza

PC 7-3-6-3 56°09′37′′ 168°48′40′′W 310 30 Jul 2007

PC 7-4-6-3 55°58.98′ 170°00.79′W 300 1 Aug 2007

ZC 7-2-ROV-1 58°23′27′′ 174°59′22′′W 914 5 Jun 2007

ZC 7-7-7-5 58°36.195′ 174°45.195′W 520 8 Sep 2007

Fiber Optic

1159 48°07′47′′ 125°05′41′′W 272–279 10 Jul 2008

Jason II

93-18 53°00′58′′ 161°13′42′′W 3,531 21 Jul 2004

93-23 53°00′44′′ 161°14′25′′W 3,442 22 Jul 2004

2095-2-1-2 51°43.311′ 173°46.879′W 2,828 25 Jul 2004

2095-2-7-3 51°48.693′ 173°49.965′W 843 25 Jul 2004

2098-2-1, 2 51°23.404′ 177°04.549′W 2,514 29 Jul 2004

2099-18-1 51°30.164′ 177°02.332′W 1,429 ?

2100-3-1,2,3 51°31.582′ 177°05.506′W 1,746 1 Aug 2004

2100-4-2, 8 51°31.607′ 177°05.568′W 1,692 1 Aug 2004

2101-1-1, 5 51°28.954′ 177°53.438′W 1,341 2 Aug 2004

2101-2-1 51°28.996′ 177°53.530′W 1,332 2 Aug 2004

2101-4-1, 2, 3 51°30.059′ 177°55.461′W 1,061 2 Aug 2004

2101-7-6,7,9 51°30.56′ 177°55.357′W 824 2 Aug 2004

2101-10-3 51°31.488′ 177°57.513′W 490 2 Aug 2004

2102-4-1, 2 51°17.521′ 179°32.532′W 1,359 3 Aug 2004

2102-6-2 4,5 51°18.549′ 179°31.693′W 888 3 Aug 2004

2102-7-1C,D,E 51°19.624′ 179°30.771′W 248 3 Aug 2004

2102-12-1A,B 51°19.886′ 179°30.599′W 192 3 Aug 2004

continued
4 8 • smithsonian contributions to zoology

TABLE A.1. continued

Research vessel 

and station no. °N Latitude ° Longitude Depth (m) Date

2102-14-1 51°19.969′ 179°30.194′W 176 3 Aug 2004

2103-5-1 51°47.85′ 179°57.255′E 1,309 4 Aug 2004

2103-7-1A 51°47.898′ 179°57.169′W 1,267 4 Aug 2004

2103-14-2 51°50.972′ 179°51.191′E 746 4 Aug 2004

2104-2-2 51°43.825′ 179°35.034′W 1,009 5 Aug 2004

2104-5-1,2,3 51°43.302′ 179°34.928′W 712 5 Aug 2004

2104-10-2 51°39.403′ 179°35.041′W 713 5 Aug 2004

2104-11-1,2 51°39.397′ 179°35.028′W 712 5 Aug 2004

Keldysh (Mir)

22 55°13′12′′ 167°14′04′′E 1,200 31 Jul 1990

Let’s Go

861-50 51°16.6′ 179°14.6′W 97 14 Jun 1986

861-52 51°14.46′ 179°11.84′E 241 17 Jun 1986

861-57 51°29.85′ 178°39.59′E 154 18 Jun 1986

861-58 51°35.88′ 178°17.01′E 371 18 Jun 1986

861-79 52°03.9′ 177°13.84′E 93 28 Jun 1986

Miller Freeman (MF)

802-VH-80-30 52°52′34′′ 168°47′43′′W 86 5 Jun 1980

833-3 51°43.0′ 177°20.8′E 567 28 Jul 1983

833-15 51°29.8′ 178°39.0′E 185 30 Jul 1983

833-16 51°25.8′ 178°50.5′E 205 30 Jul 1983

833-24 51°17.8′ 179°24.8′E 229 31 Jul 1983

833-38 51°42.9′ 178°51.5′W 582 3 Jun 1983

833-47 51°56.6′ 176°52.8′W 201 5 Jun 1983

Pacific Knight

941-29 53.06917° 169.93°W 145 10 Jun 1994

941-38 52.4685° 170.56°W 319 6 Jun 1994

941-73 52.34783° 174.57°W 112 19 Jun 1994

941-204 53.0965° 171.7°E 455 31 Jul 1994

Sea Storm

95 51°55′29′′ 178°14′59′′E 234 5 Jul 2002

116 52°04′10′′ 177°14′25′′E 87–94 11 Jul 2002

148 52°28′16′′ 173°34′31′′W 194–201 21 Jul 2002

150 52°30′47′′ 173°29′35′′W 213–220 21 Jul 2002

Starlight

84-1-7 52°36′ 172°20′W 238 12 Jul 1984

84-1-36 52°28′ 172°30′W 234 13 Jul 1984

Vesteraalen

941-15 54°04′46′′ 166°23′43′′W 79 6 Jun 1994

941-36 52°56′ 169°31′W 227 10 Jun 1994

941-51 53° 171°W 126 13 Jun 1994

941-94 52° 174°W 91 26 Jul 1994

941-153 52°10′ 179°43′E 94 11 Jul 1994

941-163 51°37′ 178°25′W 155 18 Jul 1994

941-186 52° 176°W 254 24 Jul 1994

2001-1-5 52°40′43′′ 169°07′01′′W 102 21 May 2001
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Index

Acanthoprimnoa, 9

Plumarella (Plumarella) spicata compared to, 9

Alaskan Leader, 11, 12, 46

Alaskan Trojan, 17

Alaska Sea, 11, 12, 15, 17

Albatross (Alb), 3, 5, 6, 7, 9, 12, 13, 14, 15, 20, 21, 22, 25, 46

Aleutian Islands

coral diversity in, 1–2

history of primnoid octocorals from, 3–4

Alvin, 19, 23, 25, 28, 46

Amphilaphis, 7, 11, 13

Arctic Dawn, 21

Arthrogorgia, 3, 20

diagnosis, 20

distribution, 20

Arthrogorgia ijimai, 21 

Arthrogorgia kinoshitai, 2, 21–22, 41

A. utinomii compared to, 21

description, 21

distribution, 2, 22

Arthrogorgia otsukai, 2, 22, 42

description, 22

distribution 2, 22

Arthrogorgia utinomii, 2, 20–21, 40

A. kinoshitai compared to, 21

description, 20–21

distribution, 2, 21

Caligorgia compressa, 17

Caligorgia fraseri, 3, 18

Calyptrophora, 22–23

diagnosis, 22–23

distribution, 23

Calyptrophora ijimai, 21

Calyptrophora laevispinosa, 2, 23, 28

distribution, 2, 23

Candidella (Parastenella), 23

Candidella (Parastenella) doederleini, 24
5 4 • smithsonian contributions to zoology

Challenger, 24

Cobb, John, 19, 46

coral diversity, 1–2

Deep Worker (DW), 3, 19, 46

Delta, 3, 4, 6, 11, 12, 14, 15, 17, 18, 

21, 28, 43, 46–47

Dicholaphis, 6, 9, 13

Diplocalyptra. See Thouarella 

(Diplocalyptra)

Dominator, 6, 11, 12, 14, 18, 21, 47

Epithouarella. See Thouarella 

(Epithouarella)

Esperanza, 14, 15, 19, 20, 47

Euthouarella. See Thouarella 

(Euthouarella)

Fanellia, 3, 16

diagnosis, 16

distribution, 17

Fanellia compressa, 2, 17, 18, 39

description, 17

distribution, 2, 18

F. fraseri compared to, 17

Fanellia fraseri, 2, 3, 18, 39

description, 18

distribution, 2, 19

F. compressa compared to, 17, 18

Fiber Optic, 19, 47

Gorgonia verticillata, 17, 18

Jason II, 3, 6, 9, 10, 11, 12, 15, 16, 15, 

20, 21, 22, 23, 24, 25, 47–48

Jason IV, 4, 28

Keldysh (Mir), 25, 48

Let’s Go, 6, 12, 14, 15, 18, 48

Miller Freeman (MF), 3, 12, 14, 15, 

18, 48

Narella, 4

Narella abyssalis, 2

Narella alaskensis, 2

Narella arbuscula, 2

Narella bayeri, 2

Narella cristata, 2

North Pacific, 18, 21

Ocean Olympic, 11, 21

Pacific Knight, 6, 12, 18, 21, 41, 48

Parastenella, 23

diagnosis, 23 

distribution, 23

Parastenella atlantica, 23

Parastenella bayeri, 23

Parastenella doederleini, 2, 4, 23, 24, 

25, 28, 43

description, 24

distribution, 2, 24

P. ramosa compared to, 24

Parastenella gymnogaster, 2, 4, 23, 25

distribution, 2, 25 

Parastenella pacifica, 23

Parastenella ramosa, 2, 4, 23, 24–25

distribution, 2, 25 

P. doederleini compared to, 24

Parastenella spinosa, 23

Parathouarella, 4

Patricia Lee, 12

Plumarella, 3, 5, 7, 9, 12, 13

diagnosis, 7

distribution, 7

Thouarella compared to, 9

Plumarella (Dicholaphis), 8, 9

diagnosis, 9

distribution, 9

key to, 10

Plumarella (Dicholaphis) aleutiana, 2, 8, 

10, 11–12, 27, 34

description, 11–12

distribution, 2, 8, 12

key to, 10

Plumarella (D.) superba compared 

to, 12

Plumarella (Dicholaphis) delicata, 8

key to, 10

Plumarella (Dicholaphis) echinata, 2, 8, 

15–16, 28, 37

description, 15

distribution, 2, 8, 10, 15

key to, 10

Plumarella (Dicholaphis) hapala, 2, 8, 

10, 11, 27, 33

description, 11

distribution, 2, 8, 11

key to, 10

Plumarella (Dicholaphis) nuttingi, 2, 8, 

13, 14–15, 28, 30

description, 14

distribution, 2, 8, 15

key to, 10

Plumarella (D.) superba compared 

to, 13

Plumarella (Dicholaphis) profunda, 2, 

8, 10–11, 27, 32

description, 10

distribution, 2, 8, 11

key to, 10

Plumarella (Dicholaphis) robusta, 2, 8, 

16, 28, 38

description, 16

distribution, 2, 8, 16

key to, 10

Plumarella (Dicholaphis) superba, 2, 8, 

12, 13–14, 28, 35

description, 13

distribution, 2, 8, 14

key to, 10

Plumarella (D.) aleutiana compared 

to, 12

Plumarella (D.) nuttingi compared 

to, 13

Plumarella (Plumarella), 7, 8

diagnosis, 7

distribution, 7

Plumarella (Plumarella) abietina, 8

Plumarella (Plumarella) aculeata, 8

Plumarella (Plumarella) acuminata, 8

Plumarella (Plumarella) adhaerans, 8

Plumarella (Plumarella) alba, 8

Plumarella (Plumarella) alternata, 8

Plumarella (Plumarella) attenuata, 8

Plumarella (Plumarella) aurea, 8, 9

Plumarella (Plumarella) bayeri, 8, 9

Plumarella (Plumarella) circumoperculum, 8

Plumarella (Plumarella) delicatissima, 8

Plumarella (Plumarella) dentata, 8

Plumarella (Plumarella) diadema, 8

Plumarella (Plumarella) dichotoma, 

8, 9

Plumarella (Plumarella) dofleini, 8

Plumarella (Plumarella) dofleini var. 

boninensis, 8

Plumarella (Plumarella) flabellata, 8

Plumarella (Plumarella) gracilis, 8

Plumarella (Plumarella) laevis, 8

Plumarella (Plumarella) lata, 8

Plumarella (Plumarella) laxiramosa, 8

Plumarella (Plumarella) longispina, 8

Plumarella (Plumarella) pellucida, 8

Plumarella (Plumarella) penna, 8

Plumarella (Plumarella) pourtalesii, 8

Plumarella (Plumarella) pourtalesii

forma obtusa, 8

Plumarella (Plumarella) pourtalesii

forma robusta, 8

Plumarella (Plumarella) recta, 8

Plumarella (Plumarella) rigida, 8

Plumarella (Plumarella) spicata, 2, 7, 8, 

9, 16, 27, 31

Acanthoprimnoa compared to, 9

description, 7, 9

distribution, 2, 9

Plumarella (Plumarella) spinosa 

brevispina, 8

Plumarella (Plumarella) spinosa 

typica, 8

Plumarella (Plumarella) undulata, 8
n u m b e r 6 3 4 • 5 5

Plumarella flabellata, 3, 7

Plumarella longispina, 7

Plumarella spicata, 7

Plumarella spinosa, 7, 15

Primnoa, 3, 19

diagnosis, 19

distribution, 19

Primnoa compressa, 3, 17 

Primnoa pacifica, 2, 3, 19

distribution, 2, 19

Primnoa pacifica var. willeyi, 2, 19–20

distribution, 2, 20

Primnoa resaedaformis pacifica, 19

Primnoa resaedaformis willeyi, 19

Primnoa willeyi, 19

Primnoa wingi, 2, 3, 20

distribution, 2, 20

Primnodendron, 9, 13

Primnodendron superbum, 3, 9, 13, 14

Prymnoa verticillaris, 3, 17, 18

Quest, 19

Rhopalonema, 4

Rurik Expedition, 3

Samalga Pass, 3

Sea Storm, 6, 18, 20, 48

Spirit of the North, 14, 15

Starlight, 6, 18, 21, 48

Stenella (Parastenella), 23

Stenella (Parastenella) doederleini, 24

Stenella (Parastenella) ramosa, 24

Stenella doederleini, 24

Stenella ramosa, 24

Thouarella, 3, 4–5, 7, 9, 11, 13

diagnosis, 4–5

distribution, 5

Plumarella compared to, 9

Thouarella (Amphilaphis) superba, 13

Thouarella (Diplocalyptra), 4

Thouarella (Epithouarella), 4

Thouarella (Euthouarella), 4 

Thouarella (Parathouarella), 4

Thouarella (Thouarella), 4

Thouarella (Thouarella) superba, 13 

Thouarella cristata, 2, 5–6, 27, 29

description, 5

distribution, 2, 6

Thouarella hilgendorfi, 3, 5

Thouarella sardana, 8

Thouarella striata, 13

Thouarella superba, 13

Thouarella trilineata, 2, 6–7, 27, 30

description, 6

distribution, 2, 7

U.S. North Pacific Exploring 

Expedition, 3

Vesteraalen, 6, 11, 12, 14, 18, 48

Western Alaska, 11, 14

Western Viking, 15, 18
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