Corresponding author: Stephen D. Cairns (
Academic editor: B.W. Hoeksema
Thirteen species of deep-water calcaxonian octocorals belonging to the families
Cairns SD (2018) Deep-Water Octocorals (Cnidaria, Anthozoa) from the Galápagos and Cocos Islands. Part 1: Suborder Calcaxonia. ZooKeys 729: 1–46.
Early in my career (1986) I was privileged to participate in a deep-sea submersible expedition to the Galápagos and Cocos Islands, which was sponsored by SeaPharm, Inc. and
Map of the Galápagos Islands localities from where specimens are reported in this paper.
Although the 1986
Thirteen deep-water octocoral species are reported herein, i.e., those that belong to the suborder
As explained in the Introduction, this study is based on all the deep-water calcaxonian octocoral specimens collected by the
The methodology concerning sclerite preparation for
The following abbreviations are used in the text:
Colonies unbranched, sparsely dichotomously branched, or pinnately branched. Polyps arranged in whorls, the bases of adjacent polyps sometimes fused. Polyps covered by eight longitudinal rows of body wall scales, which completely cover the polyp. At least four and up to 24 marginal and submarginal scales bear elongate slender apical spines; marginal scales do not fold over operculars. Coenenchymal scales similar to those of body wall, arranged in one layer. All scales thin, with a smooth pouter face and tuberculate inner face.
Antarctic, Subantarctic, Antipodes, New Zealand, Clarion-Clipperton Fracture Zone, Galápagos, 744–4235 m deep.
A discussion and key to the six species in this genus are given by
Four specimens (?syntypes) are included in the original description. Their deposition is unknown.
Antarctica (between 60–90°E), 3397 m depth (
Galápagos: east of Fernandina, 3381 m depth. Elsewhere: Antarctica, 3397 m depth.
Although only one incomplete colony is available (Figure
The body wall scales (Figures
Despite the long distance between the Antarctic type locality and the Galápagos, this specimen is identified as
This is the first report of this species subsequent to its original description, and was collected at virtually the same depth.
Colonies uniplanar, pinnately or dichotomously branched. Polyps cylindrical to clavate, arranged in whorls of up to 12, all polyps facing upward. Polyps covered with eight longitudinal rows of body wall scales, the number of scales per row decreasing from ab- to adaxial polyp side. Body wall scales granular, smooth, pitted, or covered with tall ridges (cristate). Inner side of opercular scales convex, covered with a multiply serrate keel.
Indo-Pacific, Atlantic, 37–2472 m deep.
The identification of the species of
1 | Branching typically pinnate |
|
– | Branching dichotomous or quasi-dichotomous |
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2 | Branches strictly opposite |
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– | Branches alternate |
|
3 | Scales in outer lateral body wall rows well developed, i.e., having the same or only slightly fewer than those in abaxial row |
|
– | Scales in outer lateral body wall rows sharply reduced in number, i.e., usually less than half the number as that in abaxial row |
|
4 | Coenenchymal scales elongate and not thick; outer surface of abaxial body wall scales on distal half of polyp highly ridged | * |
– | Coenenchymal scales irregular in shape and quite thick (tessellate); outer surface of body wall scales granular or pitted (not ridged) |
|
5 | Seven to nine scales in abaxial body wall scale rows; body walls scales pitted; polyps 1.4–1.8 mm in length |
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– | Nine to eleven scales in abaxial body wall scale rows; body wall scales granular; polyps 1.0–1.2 mm in length |
|
6 | Operculum low and inconspicuous |
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– | Operculum tall and prominent |
|
7 | Abaxial opercular scales with two to four apical points | * |
– | Abaxial opercular scales with a single point |
|
8 | Eight or more scales in each abaxial body wall row |
|
– | Six to eight scales in each abaxial body wall row |
|
9 | Apex of opercular scales prolonged into a rod-like (cylindrical) point | * |
– | Apex of opercular scales not prolonged in a cylindrical point |
|
10 | Eight to ten scales in abaxial body wall scale rows | * |
– | Eleven to thirteen scales in abaxial body wall scale rows | * |
11 | Polyps about 2 mm tall | * |
– | Polyps 1.0–1.8 mm tall |
|
12 | Coenenchymal scales elongate; inner lateral and adaxial body wall scales present; eastern Pacific |
|
– | Coenenchymal scales polygonal; inner lateral and adaxial body wall scales absent; South Pacific |
|
13 | Scales in outer lateral body wall rows well developed, i.e., having the same or only slightly fewer than those in abaxial row |
|
– | Scales in outer lateral body wall rows sharply reduced in number, i.e., usually less than half the number as that in abaxial row |
|
14 | Five or six scales in each abaxial body wall row |
|
– | Ten scales in each abaxial body wall row |
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– | Twelve or 13 scales in each abaxial body wall row |
|
15 | Three scales in each outer lateral body wall row |
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– | One or two scales in each outer lateral body wall row |
|
16 | Four polyps per whorl |
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– | Two or three polyps per whorl |
|
17 | Five scales in each abaxial body wall row |
|
– | Seven scales in each abaxial body wall row |
|
18 | Two or three polyps per whorl; eight to nine whorls per cm branch length; body wall scales with low marginal ridges |
|
– | Three or four polyps per whorl; 9–11 whorls per cm branch length; body wall scales with distal margins strongly reflexed, exposing high crest-like ridges on inner face |
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19 | Two or three polyps per whorl; seven to eight whorls per cm branch length |
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– | Four to six polyps per whorl; nine to ten whorls per cm branch length |
|
20 | Abaxial body wall scales with strong marginal ridges | * |
– | Abaxial body wall scales with only weak marginal ridges |
|
Galápagos: Tagus Cove between Isabela and Fernandina, north of Española, 308–633 m deep; Cocos Island, 628–656 m deep.
Colonies are uniplanar and taller than broad, the holotype (Figure
Colonies of various species.
There are eight longitudinal rows of body wall scales, decreasing in number from ab- to adaxial polyp side, the body wall sclerite formula being: 10–12:10–12:4–5:2. The distal five or six pairs of abaxial scales (Figure
Colonies of various species.
Polyps and sclerites of
Polyps and sclerites of
Named for the type locality of the species.
As defined by the syntype series, the type-locality extends from northern Baja California (latitude 30°30'30"N) to just north of San Diego (latitude 33°02'15"N), and includes the bathymetric range of 219–2469 m.
Galápagos: between Santa Cruz and Marchena, 605–1008 m deep. Elsewhere: northern Baja California to Monterey Bay (new records), California, 219–2469 m deep.
Colonies are uniplanar and taller than wide, the largest Galápagos specimen (
There are eight longitudinal rows of body wall scales, decreasing in number from ab- to adaxial polyp side, the body wall sclerite formula being: 6–8:1–2:1–2:1–2. The marginal and submarginal abaxial body wall scales (Figure
Polyps and sclerites of
Although this species was originally identified as
This species was clearly named after Kumao Kinoshita, and thus the name is herein changed to reflect a masculine ending.
Colonies uniplanar to slightly bushy (lyrate, dichotomous, polychotomous, biplanar) or unbranched. Polyps arranged in whorls, the polyps facing either upward or downward. Polyps consist of two annular sclerite rings, each composed of two inseparably fused scales; a pair of crescent-shaped infrabasals also present. Articular ridge present between basal and buccal body wall scale. Distal margins of body wall scales often spinose, toothed, or lobate. Operculum composed of eight scales; keels usually present on inner face of opercular scales. Coenenchymal scales elongate and flat, sometimes quite thick (as plates). Small curved tentacular platelets often present.
Tropical and temperate latitudes of Atlantic, Pacific, and Indian Oceans, 227–3531 m deep.
Including the new species described herein, the genus contains 23 species, making it the fifth most species-rich genus within the primnoids.
Syntypes: several branches from which 99% of the polyps are detached, MCZ 4815, and
Galápagos: west of Isabela, off Marchena and San Cristóbal, 509–1545 m deep.
The colony is uniplanar, equally and dichotomously branched; the largest colony (
The fused basal scale (Figures
Polyps and sclerites of
Galápagos: off Santa Cruz, Santiago, and Marchena, 445–509 m depth.
The colony is uniplanar, equally and sparsely dichotomously branched, some end branches up to 12 cm in length. The holotype (Figure
The fused basal scale (Figures
Polyps and sclerites of
Only four species in the genus have downward oriented polyps, the so called
Named in honor of John Reed (
(from
All ocean basins except Arctic, 128–4594 m deep.
Previously (
Branch fragments and detached polyps from
As mentioned in the account of
Galápagos: off Santiago, Santa Cruz, and San Cristóbal), 702–741 m deep. Elsewhere: off Panama, 1463 m depth (herein,
The colony is uniplanar, and dichotomously (laterally) and sparsely branched (Figure
The basal scales (Figure
Polyps and sclerites of
Although discussed by several authors through the years (see synonymy), this is the first subsequent report of this species since its original description.
Colonies dichotomously branched in one plane, sometimes in a lyrate pattern and sometimes as two parallel fans. Polyps arranged in whorls of up to eight, all polyps pointed downward. Each polyp covered with two (rarely three) unfused pairs of body wall scales. Articluar ridge present between basal and buccal (or medial) body wall scales. Distal margins of buccal scales smooth or spinose. A pair of infrabasal scales often present. Coenenchymal scales elongate, granular, and sometimes ridged.
Southwestern Pacific Ocean, Galapagos, Japan, Hawaii, North Atlantic, 150–1480 m deep.
The genus diagnosis is herein expanded to accommodate a species having three pairs of body wall scales, otherwise similar to the genus
Known only from northwest of Española, Galápagos, 547–653 m deep.
The colony is uniplanar, equally and dichotomously branched, the largest colony (the holotype, Figure
The basal scales (Figures
Polyps and sclerites of
Superficially this species resembles the genus
Named “
Colonies usually uniplanar and alternately pinnately branched, dichotomously branched, or bottlebrush in shape. Polyps arranged in alternate biserial fashion (nominate subgenus), crowded on all sides (subgenus
Indo-Pacific, western Atlantic, Subantarctic, 10–3181 m deep.
There are 37 species in the genus arranged in four subgenera, most listed in
Holotype:
Galápagos: east of Santa Cruz and off Roca Redonda, 717–808 m deep. Elsewhere: off Ecuador, 3181 m depth.
The colony is uniplanar to slightly bushy, the Galápagos specimen (
The body wall scales (Figure
Polyps and sclerites of
This is the deepest of the 37 known species in the genus. Only one other species occurs in this subgenus,
The specimens reported herein are only the second report of the species. It fits the re-description of the holotype given by
Colonies uniplanar to slightly bushy; branching lateral and somewhat irregular. Polyps stand perpendicular to branch, arranged independently or in pairs or whorls of up to four. Eight marginal scales present, offset in position from opercular scales; marginal, and sometimes submarginal, scales fluted; nematocyst pads present on distal inner surface of fluted marginals. Body wall scales arranged in four to eight longitudinal rows. Operculum well developed, the distal inner surface of operculars prominently keeled. Coenenchymal scales flat to highly concave, sometimes ridged. Pinnular rodlets sometimes present.
Cosmopolitan, except for eastern Atlantic, the Arctic, and off continental Antarctica, 475–3470 m depth.
Including the new species described herein, there are eight species known in this distinctive genus. Accounts of
Galápagos: off Marchena, Santiago, and Isabela, 446–578 m deep.
The colony is uniplanar, the largest colony (the holotype, Figure
The body wall scales (Figure
Polyps and sclerites of
Named in honor of Shirley Pomponi (formerly of
Branching from main branch sympodial in an ascending spiral, clockwise (R) or counterclockwise (L), usually following a repeated geometric branching formula, producing a bottlebrush colony, or dichotomous in one or more parallel planes. Branchlets repeatedly dichotomously branched, resulting in short terminal segments. Polyps large in relation to branchlets, standing perpendicular to branchlets and usually well separated. Sclerites consist of rods and scales. Axis with a brilliant metallic luster, usually golden or yellow in color, and thus referred to as the golden corals.
Cosmopolitan, including off Antarctica (
In order to manage the relatively large number of species in the genus, now standing at 70,
The four species groups of
Body Wall Rods | Body Wall Scales | |
---|---|---|
Tentacular Rods | Group A: 38 species: Indo-West Pacific, Atlantic; 100–3114 m. 2/5R, 1/4L, 3/8L, 1/3R, 2/5L, 1/3L, dichotomous, irregular, pinnate | Group B: 13 species: Indo-West Pacific, western Atlantic; 250–2271 m. 1/4R, 1/5R, 1/6R, 1/7R, 1/7R, 2/5R, biflabellate |
Tentacular Scales | Group D: 1 species: off Brazil, 1300 m. dichotomous | Group C: 18 species: Indo-West Pacific, North Atlantic, Antarctic; 204–3860 m. 1/3L, 1/4L, 2/5L, 1/4R, flabellate |
The holotype is deposited at the
Galápagos: between Isabela and Santiago; Cocos Island, 628–768 m deep. Elsewhere: Hawaiian Islands, 1758–1937 m deep.
The colony is biplanar (perhaps multiplanar), the largest colony examined (
The body wall scales (Figure
Polyps and sclerites of
Throughout Galápagos from Roca Redondo to Española, 560–816 m deep.
The colony is bottlebrush in shape (Figure
The body wall sclerites (Figure
Polyps and sclerites of
Having rods in its body wall and tentacles places
Named “
Galápagos: Roca Redonda, west of Santa Cruz, 717–806 m deep; Cocos Islands, 614–785 m deep.
The colony is bottlebrush in shape (Figure
The upper body wall sclerites (Figure
Polyps and sclerites of
Having rods in its body wall and tentacles places
Named
Colonies sparsely branched, dichotomously or trichotomously from nodes, resulting in a uniplanar colony; internodes long and hollow. Polyps non-retractile, cylindrical, armed with stout needles placed longitudinally in body wall. Spiny pharyngeal rodlets present.
Northeast Atlantic (including Mediterranean), New England Seamounts, eastern Pacific from California to Alaska, Hawaii, Galápagos, 400–2593 m deep (
Including the species described below, there are currently six species in the genus, three of which occur in the Pacific. The genus was most recently keyed and discussed by
Using one or two mitochondrial genes and six species (most of them undescribed),
Known only form the type locality.
The colony is uniplanar, the largest specimen (the holotype, Figure
The polyps are uniserially placed (Figure
Polyps and sclerites of
Named “
I am grateful to John Reed for the gift of specimens collected on the three
Station | Latitude / Longitude | Depth (m) | Date | |
---|---|---|---|---|
|
||||
1912 |
|
806 | Nov 14 1986 | |
1915 |
|
650 | Nov 15 1986 | |
1916 |
|
545–562 | Nov 16 1986 | |
1922 |
|
475–578 | Nov 19 1986 | |
1927 |
|
708–782 | Nov 21 1986 | |
1929 |
|
804 | Nov 23 1986 | |
1930 |
|
333–478 | Nov 23 1986 | |
1931 |
|
441–528 | Nov 24 1986 | |
1933 |
|
663–788 | Nov 25 1986 | |
1934 |
|
252–308 | Nov 25 1986 | |
1935 |
|
391–488 | Nov 26 1986 | |
1938 |
|
614–785 | Nov 30 1986 | |
1942 |
|
606–628 | Dec 2 1986 | |
3902 |
|
554 | Oct 17 1995 | |
3907 |
|
530–576 | Oct 19 1995 | |
3912 |
|
489 | Oct 21 1995 | |
3925 |
|
768 | Oct 28 1995 | |
3930 |
|
450 | Oct 30 1995 | |
|
||||
3108 |
|
1545 | Jul 21 1998 | |
USFWS Albatross | ||||
2818 |
|
717 | Apr 15 1888 | |
3399 |
|
3182 | Mar 24 1891 | |
3403 |
|
700 | Mar 28 1891 | |
3404 |
|
704 | Mar 28 1891 | |
3406 |
|
1008 | Apr 3 1891 | |
3410 |
|
805 | Apr 3 1891 | |
4153 |
|
1760–1837 | Aug 5 1902 | |
4357 |
|
245–284 | Mar 15 1904 | |
4530 |
|
1381–1752 | May 27 1904 | |
4537 |
|
1575–1942 | May 31 1904 | |
R/V Gilliss | ||||
21 |
|
1463 | Jan 18 1972 | |
E/V Nautilus | ||||
NA64–77–01 |
|
3381 | Jul 3 2015 | |
NA64–125–01 |
|
446 | Jul 6 2015 | |
NA64–126–01 |
|
445 | Jul 6 2015 |