Systematics and Biodiversity 5 (4): 391?407 Issued 20 November 2007
doi:10.1017/S1477200007002472 Printed in the United Kingdom C? The Natural History Museum
Stephen D. Cairns1?
& Amy Baco2
1Department of Invertebrate
Zoology, Smithsonian
Institution, P. O. Box 37012,
MRC 163, Washington, DC
20013-7012, USA
2Biology Department, MS#33,
Woods Hole, MA 02543, USA
submitted September 2005
accepted May 2007
Review and ?ve new Alaskan species
of the deep-water octocoral Narella
(Octocorallia: Primnoidae)
Abstract Five new species of the octocorallian primnoid genusNarella are described
from seven seamounts in the Gulf of Alaska, the ?rst records of this genus from the
region between Japan and the Gala?pagos, and including the deepest record of the
genus at 4594 m. All species are illustrated with SEM of their sclerites, often with
stereo pairs of the polyps. A list of the 32 valid species of Narella is given. DNA
sequence analysis using the mitochondrial genes ND6 andmsh1 was determined for
all ?ve species and outgroups, but little to no variation was found among the ?ve
morphospecies, consistentwith theknownslowrateofevolutionof themitochondrial
genes in Octocorallia.
Key words Narella, Octocorallia, Primnoidae, Alaska, ND6, msh1, seamounts, new
species
Introduction
The Primnoidae is a diverse family of deep-water octocorals
consisting of 32 genera and approximately 207 species. Al-
though species occur as shallow as 141m, most live at slope
and abyssal depths, the deepest known primnoid being Con-
vexella krampi (Madsen, 1956), which was collected from the
Kermadec Trench at 5850m. The second deepest primnoid
genus is Narella, the deepest of which occurs in the Gulf of
Alaska as reported herein for N. abyssalis from 4594m. Al-
though widespread and diverse, this is the first account of this
genus from the rather large region of the North Pacific north of
a line linking Japan, the Hawaiian Islands, and the Gala?pagos
Islands.
Materials and methods
All specimens reported herein resulted from three NOAA-
sponsored exploratory submersible cruises (see Acknowledge-
ments) made on seamounts in the northern Gulf of Alaska. The
first was in June?July 2002 using the Alvin, the second in July
2004 using the Jason II, and the third in August 2004 using
the Alvin. The second author was present on all three cruises.
Station data are given in Table 1.
Designation of polyp scales follows that of Versluys
(1906) as amplified by Bayer et al. (1983). Also, following
the terminology of Bayer (1951), the region or line that sep-
arates the lateral face of the basal and medial scales from its
dorsal (or abaxial) face is referred to as the basolateral angle
or edge. This edge is often, but not always, curved at a 90?
angle and often bears a ridge or crest.
? Corresponding author. Email: cairnss@si.edu
The opercular scales of Narella have previously been
described (Bayer, 1995, 1997) as occurring as four pairs: an
abaxial pair, two lateral pairs, and the small adaxial pair; how-
ever, close examination of the Alaskan species demonstrates a
different symmetry arrangement. Invariably there is one large,
symmetrical abaxial opercular scale, opposite which is a pair
of small, also symmetrically developed adaxial scales. The
remaining five lateral scales are asymmetrical in shape, each
having a wider adaxial edge, and being an odd number, three
occur on one side of the polyp and only two on the other, giving
the polyp a slightly asymmetrical aspect. This arrangement is
well illustrated by Fig. 9E and by Versluys (1906: fig. 102).
The SEM photomicrographs were taken by the first au-
thor using an AMRAY 1810 scanning electron microscope.
Representatives of 24 of the 32 Narella species were available
at the NMNH for comparison, 14 as type specimens.
DNA sequence analyseswere also used as an independent
source of characters to classify and compare the five new spe-
cies. Mitochondrial cytochrome oxidase I (COI) and 16S are
typically used for these comparisons in invertebrates, but both
of these genes have low variability in octocorals (France et al.,
1996; France & Hoover, 2001, 2002). Instead the mitochon-
drial genes for NADH subunit 6 (ND6) and msh1 homologue
(msh1) were used. msh1 has been shown to be more variable
in octocorals than are COI and 16S (France & Hoover, 2002)
and both msh1 and NADH subunits are currently used in oc-
tocoral phylogenetics (e.g. France & Hoover 2001; Sanchez
et al., 2003; McFadden et al., 2006, Wirshing et al., 2005).
DNA was extracted from corals using Qiagen DNeasy
kits from alcohol-preserved specimens. For PCR amplifica-
tion, primers were used from Sanchez et al. (2003). ND6-
1487F and ND3-2126R were used for ND6 and ND42599F
and Mut3458R were used to amplify the 5? end of msh1. PCR
391
392 Stephen D. Cairns & Amy Baco
Station ?N ?W depth (m) Date Seamount
Alvin
3797 53?53.47? 148?30.66? 2680 26 Jun 2002 Murray
3803?4?5 54?49.51? 152?55.73? 3075 2 Jul 2002 Chirikof
4027?10 53?56.97? 137?23.85? 2377 3 Aug 2004 Denson
4029?16 54?37.18? 136?42.54? 2736 5 Aug 2004 Dickins
4033?23 54?59.24? 140?23.99? 2634 9 Aug 2004 Welker
4033?28 54?59.24? 140?23.99? 2634 9 Aug 2004 Welker
4041?3 56?21.03? 146?22.82? 2818 17 Aug 2004 Giacomini
4041?7 56?21.03? 146?22.82? 2818 17 Aug 2004 Giacomini
4041?9 56?21.03? 146?22.82? 2810 17 Aug 2004 Giacomini
Jason II
91?bb 53?30.752? 163?26.886? 3277 m 18 Jul 2004 Derickson
93?4 53?02.446? 161?11.018? 4594 21 Jul 2003 Derickson
93?11 53?01.811? 161?12.845? 4091 21 Jul 2003 Derickson
93?14 53?01.229? 161?13.451? 3292 21 Jul 2003 Derickson
93?20 53?0.8? 161?14.421? 3465 22 Jul 2003 Derickson
93?29 52?59.032? 161?14.941? 2775 22 Jul 2003 Derickson
93?32 52?0.586? 161?14.406? 3385 22 Jul 2003 Derickson
Table 1 Station data.
conditions for msh1 were 96 ?C for 2min, and 35 cycles of
94 ?C for 1min 30 s, 57 ?C for 30 s, 72 ?C for 1min, followed
by an extension at 72 ?C for 5min. PCR conditions for ND6
were 96 ?C for 2min, and 40 cycles of 94 ?C for 1min 30 s,
45 ?C for 1min, 72 ?C for 1min, followed by an extension at
72 ?C for 5min. An Eppendorf MasterCycler epgradient (Ep-
pendorf) thermocycler was used to carry out PCR amplifica-
tions of 50?l final volume with approximately 50 ng of DNA,
1X PCR Buffer (Promega), 2.5mM MgCl2, 1mM dNTPs,
1?M of each primer and 1.5U Taq polymerase (Promega).
PCR products were purified using QIAquick PCR Puri-
fication Kit (Qiagen). Sequences were run on an ABI 3730.
Sequences were edited and aligned using Sequencher 4.2.2.
Pairwise distances are uncorrected ?p? distances and were cal-
culated in PAUP 4.0b10.
Abbreviations: NMNH=National Museum of Nat-
ural History, Smithsonian Institution, Washington, DC;
PCR= polymerase chain reaction; SEM= scanning electron
microscope; USNM=United States National Museum (now
the NMNH); H:W= height to width ratio of a sclerite.
Systematics
Subclass Octocorallia
Order Gorgonacea
Family Primnoidae Gray, 1858
Genus Narella Gray, 1870
Narella Gray, 1870: 49 [type species: Primnoa regu-
laris Duchassaing & Michelotti, 1860, by monotypy].?
Bayer, 1951: 41?43; 1995: 147?148; 1997: 511.?Cairns &
Bayer, 2003: 618?619; 2004: 7?10 (proposed neotype for
P. regularis).
Stachyodes Wright & Studer in Studer, 1887: 49 [type spe-
cies: Stachyodes regularis Wright & Studer, 1889 (not
Duchassaing & Michelotti, 1860),= Stachyodes studeri
Versluys, 1906, by subsequent monotypy (Wright & Studer,
1889: 54)].?Versluys, 1906: 86?88.?Kinoshita, 1908: 45?
47.?Ku?kenthal, 1919: 452?456; 1924: 308?309.
Calypterinus Wright & Studer, in Studer, 1887: 49 [type spe-
ciesC. allmaniWright&Studer, 1889, by subsequentmono-
typy (Wright & Studer, 1889:53)].
Diagnosis
Colonies of moderate size (up to 50 cm), dichotomously or
pinnately branched in one or more planes, or unbranched.
Polyps face downward in whorls or pairs, only rarely isolated.
Polyps protected by three or four pairs of large abaxial body
wall scales, the adaxial sclerites reduced to one or two pairs
of small buccal scales. Adaxial side of basal scales may meet
forming a complete ring, but they do not fuse as in Calyptro-
phora. Operculum consists of eight triangular scales, eachwith
a longitudinal keel on its inner surface. Coenenchymal scales
usually elongate, bearing longitudinal keels on their outer sur-
face, but in some species they are polygonal and flat. Branch
axis heavily calcified, longitudinally grooved, with a discoidal
basal holdfast.
Remarks
Narella is a species-rich genus, currently with 32 known spe-
cies (see list below), and judging from the large number of
species described herein from a relatively small collection, it
is predicted that many more species will be discovered. The
most significant observations on this genus were published by
Alaskan octocoral species of Narella 393
Figure 1 Narella bayeri, paratype from Jason II 93-14: A, lateral view of two whorls; B, two pairs of adaxial scales; C, stereo abaxial view of a
polyp; D, stereo lateral view of a polyp; E, stereo adaxial view of a polyp. Scale bars for A, C-D= 1.0mm; B= 0.1mm; E= 0.5mm.
Versluys (1906), Kinoshita (1908), Ku?kenthal (1919, 1924),
Deichmann (1936), Bayer (1951, 1995, 1997), and Cairns and
Bayer (2003, 2004). In particular, Versluys (1906) provided
excellent descriptions and illustrations of the Indonesian spe-
cies he described from the Siboga expedition. The genus is
in need of revision worldwide, not least of which is because
almost every species is known only from its type material,
which in most cases consists of just a few specimens. That
is no exception in this paper. Nonetheless, the morphological
characters used by previous authors to distinguish species ap-
pear to be quite distinctive, including the following: branching
mode, polyp size, number of polyps per whorl, number of
whorls per cm, presence or absence of adaxial buccal scales,
ring-like nature of the basal scales, shape of body wall scales
(e.g. presence or absence of a basolateral ridge, spines distal
edge), shape of opercular scales (e.g. H:W of the abaxial oper-
cular), and shape of coenenchymal scales (elongate and keeled
or polygonal and flat). To this we addmolecular sequence data.
Some of these characters may not be consistent at the species
level, as discussed by Cairns and Bayer (2003), but it does
seem that each species has a diagnostic shape of its body wall
sclerites, usually expressed in shape, size and presence or ab-
sence of basolateral ridges. These characters are emphasised
in Table 2.
List of the 32 species of Narella
Species known from Alaska: N. bayeri, N. cristata, N. arbus-
cula, N. abyssalis, N. alaskensis.
Species known from Japan: N. irregularis (Kinoshita, 1907),
N. biannulata (Kinoshita, 1907), N. megalepis (Kinoshita,
1908), N. compressa (Kinoshita, 1908).
Species known from Hawaiian Islands: N. bowersi
(Nutting, 1908), N. ornata Bayer, 1951, N. nuttingi Bayer,
1997.
Species known from western Pacific, primarily Indonesian
region: N. allmani (Wright & Studer, 1889), N. clavata
(Versluys, 1906), N. horrida (Versluys, 1906), N. obscura
(Versluys, 1906), N. orientalis (Versluys, 1906), N. parva
(Versluys, 1906), N. dichotoma (Versluys, 1906) [also
known from Hawaii], N. studeri (Versluys, 1906) [nom.
nov. for S. regularis Wright & Studer, 1889], N. grandiflora
(Ku?kenthal, 1907), N. leilae Bayer, 1951.
Species known from eastern Pacific: N. ambigua (Studer,
1894).
Species known from Antarctica: N. gaussi (Ku?kenthal, 1912).
Species known from southwest Indian Ocean: N. gilchristi
(Thomson, 1911) [= S. capensis Thomson, 1917;= S. gil-
berti nom. nud., Ku?kenthal, 1919].
Species known from Atlantic: N. regularis (Duchassaing &
Michelotti, 1860), N. versluysi (Hickson, 1909), N. bellis-
sima (Ku?kenthal, 1915), N. pauciflora Deichmann, 1936,
N. laxa Deichmann, 1936, N. spectabilis Cairns & Bayer,
2003, N. alvinae Cairns & Bayer, 2003.
Narella bayeri sp. nov.
Figs 1?2
Material examined/Types
Holotype: Jason II-93-11, one large colony, two branches, and
several short branch fragments, all dry except for one branch,
USNM1080446, and one branch at the BM (NHM2005.2346).
Paratypes: Jason II-93-14, one colony and four branches, all
dry except for one branch, SEM stubs 1138?1142, USNM
394
Stephen
D
.Cairns
&
Am
y
Baco
Narella bayeri Narella cristata Narella abyssalis Narella arbuscula Narella alaskensis
Number of pairs and
nature of body wall
scales
3 (occasionally 4),
slightly ?ared
3 (occasionally 4),
?ared
4, not ?ared (compact) 3 (rarely an unpaired
4th) , highly ?ared
3, quite ?ared
Branching Dichotomous, sparse,
uniplanar
Dichotomous, sparse,
uniplanar
Dichotomous, sparse
uniplanar
Dichotomous, sparse,
bushy
Dichotomous, sparse,
uniplanar
Number of whorls/
3 cm; whorl
diameter
11?12, crowded;
3.5mm
8?10, well-spaced;
3.4mm
9, well-spaced;
2.8mm
6?8, crowded!; 6.8mm 10.5?12, crowded;
3?7.5mm
Number of polyps/
whorl
5?6 ?7 2?3 ?4, usu. absent
from one side
2?3 ?4, usu. absent
from one side
6 ?7 4?8 ?9
Polyp length and max.
width (mm)
2.2?3.4? 1.4?1.7 2.1?3.0? 1.3?1.6 1.9?2.4? 1.0?1.3 3.4?4.7? 2.5?3.5 2.7?3.2? 1.3?2.1
Basal body wall
scales; apical
margin
Right angle
basolateral edge
with prominent
ridge; lobate
Right angle
basolateral edge
with prominent
ridge; lobate
Right angle
basolateral edge
with prominent
ridges; straight
Gently rounded
basolateral edge
occas. with tall, but
short ridge,
concave; lobate
Right angle
dorsolateral edge
with low ridge;
prominent sharp
spines
Medial body wall
scales; apical
margin
Right angle
basolateral edge
with distal ridge; not
spinose
Right angle
basolateral edge
with ridge basally;
not spinose
Low ridge at
basolateral edge;
not spinose
Gently rounded
basolateral edge
without ridge; not
spinose
Right angle
dorsolateral edge
with low, continuous
ridge; spined
Abaxial buccal body
wall scales
Striate, rarely ridged Usually ridged Low ridge at
basolateral edge
Not ridged Not ridged
Adaxial buccal body
wall scales
1 pair 1 pair 1?2 pair 1 pair and numerous
smaller scales on
adaxial body wall
1?2 pairs and small
scales on adaxial bw
H:W of large abaxial
opercular scale; keel
1.4; sharp keel 1.4?1.7; rounded keel 2.4?2.7; rounded keel 0.85?1.0; rounded keel 0.9?1.0; sharp keel
Other distinctive
characteristics
occasionally outer
surface of
operculars is keeled
coenenchymal blades
low
basolateral ridges
often bifurcate
distally
Distribution Derickson SM;
3277?4091 m
Derickson SM; 3385 m Derickson SM; 4594 m Derickson and
Giacomini; SM;
2775?3465 m
Gulf of Alaska
Sea-mounts;
2377?3075 m
Table 2 Characteristics of the ?ve Alaskan species of Narella (SM= seamount).
Alaskan octocoral species of Narella 395
Figure 2 Narella bayeri, paratype from Jason II 93-14: A, opercular scales; B, tubercles on inner surface of an opercular; C, abaxial buccal
scales; D, an adaxial buccal scale; E, a medial scale; F, basal scales; G, coenenchymal scales. Scale bars for A, C= 0.5mm;
B= 50 
m; D= 0.1mm; E-F= 1mm; G= 0.5mm.
1080447; Jason II-91-biobox, two branches in alcohol, USNM
1080448.
Type locality
53?01.811 N, 161?12.845?W (Derickson Seamount, Gulf of
Alaska), 4091m.
Description
Colonies are uniplanar and sparsely dichotomously branched,
the largest specimen (the holotype) measuring 29 cm in height
and 20 cm in maximum width, with a basal axis diameter of
4.7mm, although no colonies were collected with their bases
intact. Early branching low in the colony occurs at intervals
396 Stephen D. Cairns & Amy Baco
of every 13?25mm, but terminal branches are quite long, up
to 17 cm in length. The axis is yellow-brown in colour and
woody in texture.
Polyps are arranged in whorls of 5 to 7, 6 being the most
common number, each whorl separated by a distance of 0.5?
1.0mm; 11?12 whorls occur per 3 cm. Polyps are 2.2?3.4mm
in length (including the projecting opercular scales) and 1.4?
1.7mm in greatest width.
Each polyp is protected by three pairs of large abaxial
body wall scales and one pair of small adaxial buccal scales,
although about 10% of the polyps have an extra pair or just
one extra medial scale. The basal scales are the largest, stand-
ing perpendicular to the axis up to 2.1mm in height, the
uppermost region of each basal scale projecting as much as
0.5mm above its connection to the adjacent medial scales
as a vertical, flared rounded lobe. Each basal scale covers a
portion of the abaxial and lateral regions of the polyp, pro-
ducing a right angle basolateral edge that bears a prominent,
sharp ridge for its entire length. The basal scales do not meet
adaxially and thus do not form a closed ring. The medial
scales are roughly rectangular, up to 1.6mm in length, and
also wrap around the abaxial and lateral regions of the polyp,
and, like the basals, bear a prominent ridge at the basolat-
eral edge, but this ridge occurs only for the middle third or
distal two-thirds of the scale. The abaxial buccals are large and
flared, saddle-shaped, up to 1.5mm in length, their basolateral
edges rounded (not at a right angle), rarely bearing a ridge.
There is usually one pair of small (0.30?0.50mm) rectan-
gular adaxial buccal scales. The upper surface of the body
wall scales is granular, the granules often aligned as short
longitudinal ridges on the buccals, and the inner surfaces are
tuberculate.
All opercular scales are elongate-triangular in shape, hav-
ing a keel on the distal half of their inner surface and a concave
upper surface, but occasionally the upper surface will also
bear a short, longitudinal distal ridge. The primary abaxial
opercular scale is the largest of the operculars, up to 1.40mm
in length and 1.0mm in greatest width (H:W= 1.4) and is
symmetrical, with rather broad shoulders to either side of the
keel. The inner- and outer-lateral operculars are smaller (1.2?
1.3mm in length and 0.7?0.8mm wide, H:W= 1.5?1.9) and
are asymmetric, each bearing a shoulder to the adaxial side of
the keel. The two adaxial operculars are by far the smallest
(1.0?1.1mm in length and 0.40mm wide, H:W= 2.7) and are
symmetric in shape. Dissolution of several polyps did not re-
veal any small sclerites that could be attributed to tentacular
sclerites.
Each coenenchymal scale consists of a dense, white,
granular, rod-shaped base from which one or more thin trans-
lucent, smooth blades originate at a right angle, sometimes
several blades originating from one elongate scale. The basal
structure of a coenenchymal scale measures up to 2.1mm
in length, but rarely more than 0.5mm in width, whereas
the erect blades are up to 0.90mm in height. The bases
of the coenenchymal sclerites are coarsely tuberculate, the
surface of the upper blades granular, as are the upper sur-
faces of all the scales. Tubercles measure 15?20 
m in
diameter.
Comparisons
Narella bayeri is distinguished from the other Alaskan species
in Table 2. Among the 32 species in the genus,N. bayeri ismost
similar to N. alvinae Cairns and Bayer, 2003, a species known
only from Bermuda, but at a similar depth to N. bayeri. These
species have similarly shaped and sized polyps, but Narella
bayeri has more whorls per cm, more polyps per whorl, and
bears a ridge on the basolateral edge of the medial scales,
which is lacking in N. alvinae.
Distribution
Derickson Seamount and a seep west of that seamount, Gulf
of Alaska; 3277?4091m.
Etymology
This species is named in honour of Frederick M. Bayer, who
has named five species within this genus and who has devoted
much of his career to the study of deep-sea octocorals (Bayer,
2001).
Narella cristata sp. nov.
Figs 3?4
Material examined/Types
Holotype: Jason II-93-32, 15 branches and branch fragments
(4 dry and 10 in alcohol), SEM stubs 1154?1158, USNM
1080449, and one dry fragment, BM (NHM 2005.2345).
Type locality
52?0.586?N, 16?14.406?W (Derickson Seamount, Gulf of
Alaska), 3385m.
Description
The holotypic type series consists of about nine branches,
several short branch segments, and several detached polyps, all
presumably from the same colony; the largest branch is 8.5 cm
in length with five terminal branches. Branching appears to
be sparsely dichotomous and uniplanar, branching axils being
40??50?. The axis is golden-yellow and stiff, the largest branch
having an axis diameter of only 2mm; the base of attachment
was not collected.
Polyps are arranged in whorls of only 2?4, 3 be-
ing the most common number, one face of the colony
usually devoid of polyps. Whorls are well separated by
1.3?2.2mm; 8?10 whorls occur per 3 cm branch length.
Polyps are 2.1?3.0mm in length and 1.3?1.6mm in greatest
width, the distal end of each polyp directed obliquely
downward toward the branch axis, at least in preserved
specimens.
Each polyp is protected by three pairs of large abaxial
body wall scales and one pair of smaller adaxial buccals,
although rarely a polyp may have an additional medial body
wall scale. The basal scales stand perpendicular to the axis
up to 1.2mm in height, their upper region lobate, projecting
well above the junction with their adjacent medial scales. The
basolateral edge of each basal scale curves at a right angle and
bears a prominent ridge that is continuous across the scale.
The basal scales do not meet adaxially and thus do not form
Alaskan octocoral species of Narella 397
Figure 3 Narella cristata, holotype: A, lateral view of two whorls; B, stereo lateral and abaxial view of two polyps; C-D, stereo lateral and
adaxial views (respectively) of polyps. Scale bars for A and C= 1.0mm; B and D= 0.5mm.
a closed ring. Each medial body wall scale is 0.60?0.85mm
in length, having a curved basolateral edge that bears a low,
continuous ridge, as for the basals, their distal edges are flared
upwards (away from the polyp). The abaxial buccals are larger
(0.9?1.0mm in length), ridged on their curved basolateral
edge, and flared outward. The distal margin of both medials
and abaxial buccals are corrugated and irregular in morpho-
logy (Figs 4C, E). One pair of adaxial buccal scales is easily
seen in adaxial view when a polyp is detached from its whorl,
these adaxial buccal scales being elliptical, 0.25?0.35mm in
greater diameter, and bearing a low central boss; they occur
directly adjacent to the lower edges of the adaxial opercular
scales.
The opercular scales are elongate-triangular in shape,
often with an attenuate tip, the larger symmetrical abaxial
opercular scale up to 1.1mm in length and 0.81 in basal width
(H:W= 1.4?1.7). The lateral operculars are asymmetrical, of
about the same length, but always narrower, as only one side
is developed into a shoulder, resulting in a higher H:W of
1.8?2.3. The symmetrical adaxial operculars are the smallest,
only 0.65?0.75mm in length and 0.25?0.30mm in basal width
(H:W= 2.4?2.7). All operculars are highly concave on their
outer surfaces, tuberculate below, and bear a poorly developed,
rounded keel on the inner distal region.
Coenenchymal scales are fusiform, up to 0.8mm long
and rarely more than 0.15mm in width, each of which bears a
low (usually less than 0.25mm in height) blade or central boss.
Comparisons
Although similar to Narella bayeri, particularly in body scale
architecture, Narella cristata differs from that species in sev-
eral significant characters (Table 2): it has smaller polyps,
fewer polyps per whorl, and fewer whorls per cm. Further-
more, it has much smaller coenenchymal scales and its abaxial
buccal scales are usually ridged and are more flared, as are its
medial scales.
Distribution
Known only from the type locality of Derickson Seamount,
Gulf of Alaska, 3385m.
Etymology
The species name cristata (Latin: cristatus, crested, ridged) is
an allusion to the well-developed basolateral ridges present on
all three pairs of body wall scales.
Narella abyssalis sp. nov.
Figs 5?6
Material examined/Types
Holotype: Jason II-93-4, 20 branches (16 in alcohol, 4 dry),
SEM stubs 1150?1153, USNM 1080450 and two dry frag-
ments, BM (NHM 2005.2343).
Type locality
53?02.446?N, 161?11.088?W (Derickson Seamount, Gulf of
Alaska); 4594m.
Description
The holotypic type series consists of about 20 branches and
many detached polyps, all presumably from the same colony;
the largest branch is 10 cm in length with four terminal
branches; the base was not collected. Branching appears to
be sparse and uniplanar, branching axils being about 15?. The
398 Stephen D. Cairns & Amy Baco
Figure 4 Narella cristata, holotype: A, opercular scales; B, laciniate edge of an opercular scale; C, abaxial buccal scales; D, adaxial buccal
scale; E, medial scales; F, coenenchymal scales showing tall blades; G, ridged basal scales. Scale bares for A, C, E, G= 0.5mm; B, D,
F= 0.1mm.
axis is golden yellow and quite brittle, 1.7mm in maximum
diameter; the basal attachment was not collected.
Polyps are arranged in whorls of only 2?4, 3 being the
most common number; one face of the colony is usually devoid
of polyps, the other three polyps of a whorl being disposed
at right angles to each other. Whorls are well separated by
1.3?2.3mm; invariably 9 whorls occur per 3 cm branch length.
Polyps are compact (not flared), 1.9?2.4mm in length and 1.0?
1.3mm in maximum width.
Each polyp is protected by four pairs of large abaxial body
wall scales and 1?2 pairs of smaller adaxial buccal scales.
The large basal scales stand perpendicular to the axis up to
Alaskan octocoral species of Narella 399
Figure 5 Narella abyssalis, holotype: A, stereo view of part of a whorl; B, stereo lateral view of a polyp; C, stereo lateral view of a polyp; D,
stereo opercular view of a polyp. All scale bars are 1mm.
0.95mm in height, having a straight distal margin that does
not rise far above its junction with the adjacent medial scale.
The basolateral edge of each basal scale curves at a right angle
and bears 1?3 prominent, finely serrate ridges up to 0.3mm in
height. The basal scales do not meet adaxially and thus do not
form a closed ring. The two pairs of medial scales are much
broader than long, each 0.4?0.6mm in length. Like the basal
scales, each lateral scale also curves in a right angle and bears
low ridges on its basolateral edge, which are aligned with those
of the basals. The abaxial buccal scales are slightly larger than
the medials (up to 0.95mm in length) and only slightly flared,
producing a small cowl of about 0.25mm; they also bear low
basolateral ridges. In most cases, the basolateral ridges of the
body wall scales bifurcate near the distal scale margin. Also,
the distal margin of most body wall and coenenchymal scales
are finely serrate. The small adaxial buccal scales occur as
one or two pairs and are elliptical in shape, measuring about
0.3mm in greater diameter; they are highly sculptured with
ridges and granules.
The opercular scales are isosceles triangular in shape,
the larger abaxial operculars symmetrical and up to 1.2mm in
length and 0.45mm in basal width (H:W= 2.4?2.7). The lat-
eral operculars are asymmetrical, often slightly longer than the
abaxials (1.25mm) having a H:W of 2?3.5, and the small sym-
metrical adaxial are rarely over 0.80mm in length (H:W= 3.0?
3.2). All operculars have a sagittally highly concave outer sur-
face, which corresponds to a low blunt rill on the inner surface.
The tips of the abaxial operculars are usually curved downward
over the polyp, and the edges of all operculars are finely ser-
rate. There is a small mound near the base of the outer surface
of most operculars from which granules radiate (Fig. 6A).
Coenenchymal scales are fusiform and similar in shape
to those described for other species, reaching up to 1.3mm in
length and 0.45mm in blade height. Scales with taller blades
are closest to the basal scales and seem to fit into notches of the
basal scales, thus anchoring these large scales and restricting
their lateral movement.
Comparisons
Only two other species of Narella, N. laxa Deichmann, 1936
andN. spectabilisCairns and Bayer, 2003, both from the North
Atlantic, consistently have four pairs of large abaxial bodywall
scales. All three species are found in relatively deepwater, have
few polyps per whorl, and whorls that are fairly well spaced.
A. E. Verrill (see Bayer & Cairns, 2003) suggested that four
pairs of body wall scales would constitute a generic difference,
but he did not publish his results. Later authors did not follow
that suggestion, and the presence of four pairs of body wall
scales in a low percentage of polyps of Narella bayeri, N.
arbuscula and N. cristata that otherwise have only three pairs
of body wall scales argues for a transitional nature of this
character. Narella abyssalis differs from N. laxa in having
smaller polyps and consequently more whorls per cm, smaller
basal scales without an apical spur, more elongate opercular
scales, fewer pairs of adaxial buccal scales, and having low
baso-lateral ridges on all body wall scales. However, Narella
abyssalis is remarkably similar to N. spectabilis, a species
thus far known from only one specimen from the Bahamas,
differing only in having much smaller polyps, smaller adaxial
buccal scales, and in being branched.
Distribution
Known only from the type locality of Derickson Seamount,
4594 m.
400 Stephen D. Cairns & Amy Baco
Figure 6 Narella abyssalis, holotype: A, opercular scales; B, adaxial buccal scales; C, distal inner surface of an opercular scale; D, medial
scales; E, basal scales; F, coenenchymal scales in situ; G, coenenchymal scales showing tall blades. Scale bars for A-B,
D-G= 0.5mm; C= 0.1mm.
Etymology
The species name abyssalis (Latin: abyssus, deep sea, abyss +
alis, an adjectival suffixmeaning pertaining to) is an allusion to
the great depth at which this species was collected, the deepest
of any known Narella.
Narella arbuscula sp. nov.
Figs 7?8
Material examined/Types
Holotype: Jason II-93-29, 20 branches in alcohol and three
branches dry, SEM stubs 1143?1149, USNM 1080451, and
Alaskan octocoral species of Narella 401
Figure 7 Narella arbuscula, holotype: A, abaxial-opercular view of part of a whorl; B, adaxial view of a polyp; C-E, stereo abaxial, lateral, and
adaxial views (respectively) of polyps. All scale bars are 1mm.
one dry fragment, BM (NHM 2005.2344). Paratypes: Jason
II-93-20, one colony in alcohol and one dry branch, USNM
1080452; Alvin 4041-3, one colony in alcohol, USNM
1075465; Alvin 4041-7, one colony in alcohol, USNM
1075466; Alvin 4041-9, four branches in alcohol and one dry,
USNM 1075467.
Type locality
52?59.032?N, 161?14.941?W (Derickson Seamount, Gulf of
Alaska), 2775m.
Description
The holotypic type series consists of 23 branch fragments, all
presumably from the same colony; the largest fragment con-
sists of seven terminal branches and a total height of 10 cm.
Colonies are sparsely and dichotomously branched, appear-
ing to be bushy in shape, the large fragment showing three-
dimensionality. Terminal branches are long, up to 12 cm in
length. No colonies were collected with an intact base. The
axis is golden yellow in colour and rather rigid, 1.8mm in
diameter in the holotype.
Polyps are arranged in whorls of 6?7, but predominantly
6, each whorl directly adjacent to the next (crowded) or sep-
arated by less than 1mm; 6?8 whorls occur per 3 cm. Polyps
are 3.4?4.7mm in length and rather wide, 2.5?3.5mm in the
highly flared basal and buccal regions.
Each polyp is protected by three pairs of large abaxial
body wall scales and one pair of smaller adaxial buccal scales,
as well as several dozen even smaller adaxial scales that are
irregularly arranged and attach directly to the adaxial body
wall. Rarely an additional unpaired medial scale will be
present. The basal scales are quite large, up to 2.9mm in height,
the uppermost lobe of each scale projecting up to 0.90mm
above its connection to the adjacent medial scale. Basal scales
do not have a right angle basolateral edge, but rather gently
curve around the polyp. This region of curvature of the basal
is either smooth or occasionally may bear a tall (up to 1mm)
slender ridge (Figs 7D, 8E) proximally. The basal scales do
not meet adaxially and thus do not form a closed ring. Medial
scales are up to 1.8mm in length, and, like the basals, curve
around the basolateral edge of the polyp and are rarely ridged;
they are also flared outward like the basals. The abaxial buc-
cals are slightly larger (1.7?2.2mm in length), have rounded
basolateral edges, and rounded, highly flared distal margins,
which produce a cowl 0.9?1.0mm in height surrounding the
lower edges of the opercular scales; they are never ridged. The
rounded, almost flattened, nature of the basolateral edges of
the body wall scales and their flared distal margins produce a
?fluted? or open nature to the polyp. There is usually one pair
of smaller, rectangular (0.5?0.8mm in greater width) adaxial
buccal scales, as well as several dozen smaller elliptical to oval
scales (0.2?0.3mm in greater diameter) attached directly to the
otherwise naked/unprotected adaxial side of the body wall.
The opercular scales are triangular in shape, but are quite
broad. The primary (or sagittal) abaxial opercular is the largest
of the operculars, symmetrical, spade-shaped, up to 1.7mm
in length, and usually broader than long (H:W= 0.85?1.0).
It is widest at mid-height and is highly concave along the
sagittal axis. The lateral operculars are narrower (also up to
1.7mm in length, but only about 1mm wide, H:W= 1.5?1.7)
and asymmetric; the adaxial operculars are the smallest (only
1.0?1.1mm in length and about 0.45mm inwidth, H:W= 2.3?
2.4) and are symmetric. Although the outer sagittal axis of all
operculars is concave, the corresponding mound on the inner
402 Stephen D. Cairns & Amy Baco
Figure 8 Narella arbuscula, holotype: A, opercular scales, the smaller ones in the adaxial position; B, abaxial buccal scales; C, adaxial buccal
scales; D, medial scales; E, basal scales; F, lateral view of coenenchymal scales showing tall blades; G, tubercles on inner side of a
coenenchymal scale. Scale bars for A, D= 0.5mm; B, E= 1.0mm; C= 0.1mm; F= 0.5mm; G= 50 
m.
side is not expressed as a sharp keel, but rather as an elongate
rounded ridge.
The coenenchymal scales are similar in size and shape to
those ofNarella bayeri, consisting of a thick basal portion from
which one or two tall (up to 1.3mm), thin, translucent blades
originate perpendicular to the branch axis. Specific tentacular
sclerites were not noted. All sclerites are tuberculate on their
inner surfaces.
Alaskan octocoral species of Narella 403
Figure 9 Narella alaskensis (A-B, E, paratype from 4027-10; C-D, holotype): A, lateral view of two whorls; B, adaxial opercular view; C, stereo
abaxial view of a polyp; D, stereo lateral view of a polyp; E, stereo opercular view. Scale bars for A-C, E= 1.0mm; D= 0.5mm.
Comparisons
Narella arbuscula is distinguished from other congenerics in
the Gulf of Alaska (Table 2) by having almost flat (concave)
and thus highly flared body wall scales that do not have a sharp
basolateral edge, as well as having the largest polyps of any
species. But a similar polyp morphology is shared with at least
two species from Indonesian waters, N. obscura (Versluys,
1906) and N. dichotoma (Versluys, 1906), which are similar to
each other. Narella arbuscula is distinguished from both spe-
cies by having larger polyps, a bushy colony (not uniplanar),
and by having a tall ridge (Figs 7D, 8E) on the proximal
basolateral margin of the basal scales. It is further differenti-
ated fromN. obscura by having elongate, ridged coenenchymal
scales (those of N. obscura are polygonal and not ridged) and
from N. dichotoma in having more polyps per whorl and more
closely spaced whorls.
Distribution
Derickson and Giacomini Seamounts, Gulf of Alaska; 2775?
3465m
Etymology
The species name arbuscula (Latin: arbuscula, small tree)
refers to the three dimensional, bushy shape of the colonies.
Narella alaskensis sp. nov.
Figs 9?10
Material examined/Types
Holotype: Alvin 4033?23, three branches (one dry, one in al-
cohol), SEM stubs 1159?1163, USNM 1075470, and 1 al-
cohol branch at BM (NHM 2005.2347). Paratypes: Alvin
3797, two branch fragments in alcohol, USNM 1080453;
Alvin 3803-4-5, four branch fragments in alcohol, USNM
1080454; Alvin 4033-28, three branch fragments in alco-
hol, USNM 1075471; Alvin 4027-10, eight branches (six
in alcohol, two dry), USNM 1075468; Alvin 4029-16,
three branch fragments in alcohol, one dry, USNM 1075469.
Type locality
54?59.24?N, 140?23.99?W(Welker Seamount, Gulf ofAlaska),
2634m.
Description
The holotype is a branch segment only 11 cm long and 7mm in
branch diameter, showing only one bifurcation; the paratypes
show the species to have a uniplanar colony. Branching appears
to be sparse with long terminals. No specimens were collected
with an intact base. The axis is golden yellow, up to 1.33mm
in diameter in the holotype.
Polyps on large diameter branches are arranged in whorls
of 7?9, but predominantly 8, the polyps of each whorl tightly
packed one against another and contiguous with those of ad-
jacent whorls, showing no intervening branch coenenchyme.
This extremely crowded polyp arrangement results in about
10.5 whorls per 3 cm, and thick cylindrical branches with a
whorl diameter of up to 7.5mm. On smaller-diameter branches
there are only 4 or 5 polyps per whorl and whorls are spaced
about 0.5mm apart. Polyps are 2.7?3.2mm in length and 1.3?
2.1mm in maximum width.
404 Stephen D. Cairns & Amy Baco
Figure 10 Narella alaskensis, holotype: A, opercular scales; B, inner surface of an opercular scale; C, abaxial buccal scales; D, an adaxial
buccal scale; E, adaxial buccal scales in situ; F, medial scales; G, an adaxial body wall scale; H, basal scales; I, coenenchymal scales
showing tall blades. Scale bars for A, C, E-F, H= 0.5mm; B, D, G, I= 0.10mm.
Each polyp is protected by three pairs of large abaxial
bodywall scales, 1?2 pairs of smaller (0.30?0.40mm inwidth),
oval adaxial buccal scales, and a variable number of even
smaller (0.12?0.20mm) scales that are directly attached to the
adaxial body wall. The large basal scales stand perpendicular
to the branch and reach as much as 2.0mm in height, the
distal most 0.6?1.0mm rising well above the junction with
the adjacent medial scales as a sharp to blunt projecting spine
or spur. These spines are often broken off or worn down in
older polyps. The basolateral edge of the basal scale curves at
Alaskan octocoral species of Narella 405
a right angle and sometimes bears a ridge up to 0.25mm in
height near the coenenchyme, but may just as often lack the
ridge. Occasionally there is a short, low longitudinal ridge on
the inner surface of the projecting spur. The basal scales do not
meet adaxially and thus do not form a closed ring. The medial
scales are up to 1.6mm in length, highly flared distally, and
also usually bear a low, smooth ridge at their basolateral edge.
Each medial scale also bears a prominent sharp spine on its
basolateral margin. The large abaxial buccal scales are up to
1.8mm in length, gently curved at the basolateral edge, and do
not bear ridges. Their margins project outward up to 0.5mm
as a cowl surrounding the bases of the opercular scales.
The opercular scales are triangular in shape, the single
large symmetrical abaxial opercular up to 1.1mm in length
and often of the same width (H:W= 0.9?1.0). The five lateral
operculars are asymmetrical in shape, up to 1.0mm in length
and about 0.4?0.6mm in width (H:W= 1.7?2.3); the small
symmetrical pair of adaxial operculars are only 0.6?0.8mm in
length and 0.24?0.34mm in width (H:W= 2.2?2.8). All oper-
culars are highly concave on their outer surfaces and tubercu-
late and distally keeled on their inner surfaces.
Coenenchymal scales are fusiform and similar in shape
to those described for other species, reaching up to 0.9mm in
length and 0.50mm in blade height.
Discussion
Some of the distinguishing characters used in Table 2 vary for
this species depending on the location of the polyp within the
colony. For instance, polyps on terminal branches are some-
what smaller, occurring only 4 or 5 per whorl, the whorls
only slightly spaced from one another, altogether producing
a whorl diameter of only 3?4mm. These polyps also have
well-developed spines on the margins of both basal and me-
dial scales (Fig. 9D). Polyps on larger-diameter (to 7mm)
proximal branches are larger and more crowded, with as
many as 9 polyps per whorl, and the whorls are very closely
spaced. The spines on the basals and medials are present
but somewhat blunt as though through wear over time (Fig.
9C). At first these isolated branches were thought to rep-
resent two closely related species, thus underscoring the
importance of having a complete colony or colonies for
examination.
Comparisons
Narella alaskensis is readily distinguished from other congen-
erics from the Gulf of Alaska (Table 2) by its possession of
prominent spines on the distal margin of the basolateral edge
of the basal and medial scales. But several other species in the
genus have such spination, one of which N. parva (Versluys,
1906), known only from Indonesia at 1300?1595m, has a re-
markably similar polypmorphology.Narella alaskensis differs
from that species in having more polyps per whorl (N. parva
has only 3?4 per whorl) and in having very crowded (contigu-
ous) whorls (those of N. parva are separated by as much as
1mm). Furthermore, it is unlikely that a species would have
such broad distribution and bathymetric ranges, encompassing
the tropical west Pacific and boreal Gulf of Alaska. But, as N.
parva is known from only several poorly preserved specimens,
future comparisons are encouraged between Indonesian and
Alaskan faunas.
Distribution
Chirikof, Murray, Welker, Denson, and Dickins Seamounts,
Gulf of Alaska; 2377?3075m, the most widespread and shal-
lowest of the Narella in the Gulf of Alaska.
Etymology
This species is named for the Gulf of Alaska, being the most
widespread Narella collected from the Gulf thus far.
Genetic data
Seven hundred and eighty-six base pairs of msh1 and 610 bp
of ND6 were compared for these species. Other Alaskan prim-
noids given the preliminary identifications of Calyptrophora
japonica, and Primnoa pacifica var. willeyi, were used as out-
groups. Table 3 summarizes the uncorrected ?p? pairwise dis-
tances between each species for msh1. Only a single haplo-
type was obtained for each species except Narella arbuscula,
which had two haplotypes that differed by one base pair. One
haplotype was represented by individuals collected on Derick-
son Seamount, and the other by individuals from Giacomini
Seamount. Interspecific distance ranged from 0% to 1.39%.
The distance between the species of Narella and the outgroup
species ranged from 1.39% to 3.68%. The two outgroup spe-
cies also had a pairwise distance of 3.68%.
Variation in ND6 was very low, with only three variable
positions within the Alaskan Narella, and an additional two
variable positions when the outgroups were added. Table 4
summarizes the pairwise distances between each species for
ND6. Among the five new Narella species, pairwise distances
ranged from 0 to 0.33%. Comparisons of the Narella species
to the outgroup species ranged from 0.33% to 0.49%. These
results suggest that ND6 is not variable enough to be useful
for intraspecies or intrageneric differences within the octocoral
Family Primnoidae.
Three species N. bayeri, N. cristata and N. abyssalis,
showed no sequence differentiation between species for
either gene. All three species were collected from
Derickson Seamount. It has been suggested that seamounts
may act as hot spots for speciation due to their isolation (e.g.
Parker & Tunnicliffe, 1994; Rogers, 1994; DeForges et al.,
2000), and these three species may represent a relatively re-
cent radiation of Narella on Derickson Seamount. Sanchez
et al. (2003) point out that these genes may provide limited
resolution for recently evolved species, however, it is also
likely that msh1 and ND6 simply have too low variation levels
to distinguish all species within this genus. Five species in
the octocoral genus Corallium (Family Coralliidae), showed
no sequence variation in the first 100 bp of the msh1 gene
(France & Hoover, 2002). Two species of Hawaiian Narella
differed by 2.8% in the same 100 bp region of msh1 (France &
Hoover, 2002). Other published data on this region of the msh1
with multiple species in a single genus include those of Sanc-
hez et al. (2003) and Wirshing et al. (2005), but unfortunately,
they did not include pairwise distances in their results. The only
406
Stephen
D
.Cairns
&
Am
y
Baco
N. bayeri (AK-21,
18, and 17)
N. cristata
(AK-20)
N. abyssalis
(AK-15)
N. arbuscula
haplo 1 (AK 16
and 19)
N. arbuscula
haplo 2 (AK 41,
42, and 109)
N. alaskensis
(AK 36, 37, 38,
WPC 1, 8)
Calyptrophora
japonica
(Ak-94)
Primnoa paci?ca
var. willeyi (AK-107)
Gen Bank # for
Haplotype
N. bayeri (AK-21, 18,
and 17)
? DQ234754
N. cristata (AK-20) 0.0 ? DQ234753
N. abyssalis (AK-15) 0.0 0.0 ? DQ234755
N. arbuscula haplo 1
(AK 16 and 19)
0.507 0.507 0.507 ? DQ234751
N. arbuscula haplo 2
(AK 41, 42, and 109)
0.634 0.634 0.634 0.127 ? DQ234752
N. alaskensis (AK 36,
37, 38, WPC 1, 8)
1.394 1.394 1.394 1.141 1.267 ? DQ234750
Calyptrophora
japonica (Ak-94)
2.788 2.788 2.788 2.788 2.915 3.676 ? DQ234756
Primnoa paci?ca
var. willeyi (AK-107)
1.394 1.394 1.394 1.394 1.521 2.281 3.676 ? DQ234757
Table 3 Uncorrected ?p? pairwise distances (%) for new species of Narella, based on MutS sequences.
N. bayeri (AK-21,
18, and 17)
N. cristata
(AK-20)
N. abyssalis
(AK-15)
N. arbuscula
(AK 16 and 19,
41, 42, and
109)
N. alaskensis
(AK 36, 37, 38,
WPC 1, 8)
Calyptrophora
japonica
(Ak-94)
Primnoa paci?ca var.
willeyi (AK-107)
Gen Bank # for
Haplotype
N. bayeri (AK-21, 18, and 17) ? DQ234762
N. cristata (AK-20) 0.0 ? DQ234761
N. abyssalis (AK-15) 0.0 0.0 ? DQ234763
N. arbuscula (AK 16 and 19,
41, 42, and 109)
0.328 0.328 0.328 ? DQ234760
N. alaskensis (AK 36, 37,
38, WPC 1, 8)
0.328 0.328 0.328 0.164 ? DQ234759
Calyptrophora
japonica (Ak-94)
0.492 0.492 0.492 0.492 0.492 ? DQ234764
Primnoa paci?ca
var. willeyi (AK-107)
0.328 0.328 0.328 0.328 0.328 0.492 ? DQ234765
Table 4 Uncorrected ?p? pairwise distances (%) for new species of Narella, based on ND6 sequences.
Alaskan octocoral species of Narella 407
estimate of the mutation rate for the msh1 gene in octocorals
is 0.2% per million years (LePard, 2003). Although msh1 is
considered the most rapidly evolving mitochondrial protein-
encoding gene for octocorals (France &Hoover, 2002), clearly
there is still a need for a more informative marker at the species
level.
Acknowledgements
The specimens included in this report were collected on two NOAA
Office of Ocean Exploration cruises and a NOAA West Coast and
Polar Programs NURP cruise to seamounts in the Gulf of Alaska
during 2002 and 2004. We would like to acknowledge the following
grant numbers to ARB: UAF 040118 and NA04OAR4600051.
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