Corresponding author: Odalisca Breedy (
Academic editor: L. van Ofwegen
Breedy O, Guzman HM (2015) A revision of the genus
The genus
The eastern Pacific
Other authors revisited this genus.
As it has been addressed before (e.g.
This research represents the first part of the fifth review in a series proposed to evaluate the gorgonian genera historically reported for the shallow eastern Pacific waters. The second part will treat the genus
Museum of Natural History (former BM, British Museum), London, UK
Centro de Investigación en Estructuras Microscópicas, Universidad de Costa Rica
Colección de referencia de Biología Marina Universidad Del Valle, Cali, Colombia
Instituto del Mar de Perú, Lima, Perú
NAZCA Instituto de Investigaciones Marinas, Salinas, Ecuador
Museum of Comparative Zoology, Harvard University, Boston, USA
Scanning Electron Microscopy
Smithsonian Tropical Research Institute, Panamá
Museo de Zoología, Escuela de Biología, Universidad de Costa Rica, Costa Rica
Universidad de Los Andes, Laboratorio de Biología Molecular Marina, Bogotá, Colombia
Colecciones Biológicas, Universidad Peruana Cayetano Heredia, Lima, Perú
Museum of Natural History (former United States National Museum), Smithsonian Institution, Washington, USA
Yale Peabody Museum of Natural History, New Haven, USA
Zoologisches Institut und Zoologisches Museum der Universität Hamburg, Germany
The type specimens used in this study were analysed during visits to museums or acquired on loan from the BM,
For microscopic study, specimens were prepared for
Data on geographical distribution are from our personal collections, Museum catalogues, and published monographs. In some cases there is just one specimen in the collection under a species name, which automatically constitutes the holotype. When needed we designated lectotypes to establish the species identity and avoiding future confusion.
Terminology is according to
Variation is expected in the diameter of stems and branches either in preserved or dry material, due to the preservation history of specimens. Most of the type material is dry and old (more than a century). Some specimens are deteriorated. According to
The polyp sclerites are basically rods and spindles, in most of the cases it was not possible to determine the anthocodial arrangement because of the deterioration of the type material, few type specimens were found preserved in ethanol. However, we did notice that there is no collaret and points arrangements as in other plexaurids. The sclerites are mostly placed longitudinally in irregular arrangements or in some cases in points. Although some variation is expected, the colour of sclerites and colonies is remarkably constant. Some species dye the ethanol a dark purplish colour when preserved. In this genus, colours are mostly hues of brown, and the sclerites do not present much colour variation.
1 | Sclerites in form of spindles and capstans, with tubercular sculpture arranged in whorls, measuring less than 0.3 mm long. Anthocodial sclerites mainly flat rods forming weak or irregular collaret and points arrangements |
|
– | Sclerites in form of spindles highly modified, measuring more than 0.3 mm long. Anthocodial sclerites other than flat rods forming collaret and points or variation of that arrangement |
|
2 | Coenenchyme contains unilateral spinous sclerites, polyps retract into shelf-like or tubular calyces |
|
– | Coenenchyme does not contain unilateral spinous sclerites, polyps retract into prominent or slightly raised dome-shaped polyp mounds |
|
3 | Coenenchymal sclerites mostly spindles, straight, curved, branched, heavily ornamented with complex tubercles, and prickles. Sclerites below the points may be transverse, but small and numerous, not forming distinct collaret |
|
– | Coenenchymal sclerites mainly capstans, radiates and spindles, thin, sharp, with tubercles, some modified as incomplete disks, but not heavily ornamented |
|
Colonies planar or multiplanar, bushy, arborescent, laterally branched, pinnately branched, dichotomous or with long flexible branches without occasional branch anastomosis. Branches and branchlets upward bending almost parallel, and with about the same thickness all along, frequently with slightly enlarged tips. Coenenchyme moderately to very thick (compared to other plexaurids) with a circle of longitudinal canals surrounding the axis and dividing the coenenchyme into a thin inner layer or axial sheath, and a thicker outer layer. Polyps fully retractile within prominent calyces longitudinally and closely placed and at all sides of the branches. Calyces prominent, shelf-like or tubular, with prickly projecting spindles, longitudinally arranged, imbricate or not. Anthocodial sclerites mainly small spindles, in weakly differentiated transverse collaret and points below the tentacles, or just with some sclerites scattered along the neck zone of the polyp. Sclerites of the outer coenenchyme mostly long, unilateral spinous spindles, often massive, sculptured on inner surface by crowded complex tubercles and on outer surface by simple spines or prickles, and in some species with a few more or less prominent coarse, prickly projections. Axial sheath composed of capstans, spindles, or oval forms. Sclerite colours white, various hues of yellow, amber, orange, purple and red. Anthocodials with lower colour hues.
From Cape Hatteras, North Carolina to Brazil, including Bahamas, Greater and Lesser Antilles, and Caribbean islands (
Colony shape and branching patterns are variable among
Holotype:
The holotype is a 20 cm tall and 12 cm wide colony, the branching is lateral, almost in one plane (Fig.
Reported only from the type locality, Bay of Panamá. This species has not been found in our recent surveys along the Pacific coast of Panamá. No data available about the depth range.
This species was first mentioned by
Lectotype (here designated):
The lectotype is an 8.5 cm tall and 4 cm wide incomplete colony, branching is sparingly dichotomous (Fig.
Panamá, Bahía de Caraquéz, Ecuador (
This species was first mentioned by Verrill in1866, together with
Comparative features of the eastern Pacific genus
Species | Colony colour | Colony shape | Length unbranched ends | Diameter of end branchlets | Coenenchyme | Calyx height | Calyx diameter | Calyx arrangement at branchlets | No. calyces/cm | Largest spindles | Axial sheath sclerites length range | Sclerite colours |
---|---|---|---|---|---|---|---|---|---|---|---|---|
|
b | cand | 70 | 7 | T | 2.50 | 2 | cl | 21 | 1.8 (2) | 0.15–0.30 | lb, do, py, w |
|
lb | bu | 20 | 8 | t | 4 | 1.80 | cl | 14 | 1.6 (2.6) | 0.13–0.56 | w, c |
|
lb | bu | - | - | - | - | 1.50 | s | - | 1.2 | - | - |
|
lb-b | bu | 40 | 5 | mt | 2.6 | 1.75 | s | 14 | 1.3 (1.8) | 0.14–0.30 | py, lb, w, c |
|
lb | cand | 70 | 8 | mt | 5 | 0.70 | cl | 26 | 2.0 (2.28) | 0.12–0.46 | w, c |
No type material available for this study, data given from
No information available
calyx arrangement: cl, close; s, sparse
coenenchyme: t, thin, T, thick, mt, moderate thick
colour: b, brown, bi, bicoloured; c, colourless, transparent; do, dark orange; lb, light brown; py, pale yellow; w, whitish
colony shape: bu, bushy, ascending; cand, candelabrum, irregular dichotomous
Plate 3, figs 5–8 (
Holotype figured. According to
Reported for Perú, the type locality.
According to
Lectotype (here designated):
COSTA RICA:
The lectotype is a 14 cm tall and 12 cm wide colony, flabellate, spreading in one plane. It has a sponge attached to the main branches (Fig.
The other material examined is very consistent with the lectotype, variation is basically in the number of branches and size of the colonies. The largest colony measured was a specimen from Perú reaching 35 cm tall and 30 cm wide (Fig.
Panamá: Gulf of Chiriquí, Pearl Islands, 10–20 m. Costa Rica: Nicoya Gulf, Santa Elena Peninsula, Marino Ballena National Park, Golfo Dulce, from 25–40 m. Colombia: Málaga Bay (
This species was described by
The main difference to separate this species from
Holotype:
The holotype is a 17 cm tall and 10 cm wide stout and rigid colony, branching mostly dichotomous (Fig.
Reported only from the type locality, Pearl Islands, Panamá. This species has not been found in our recent surveys along the Pacific coast of Panamá.
Colonies branching mostly in one plane, fan-like, dichotomous, pinnate-like, or unbranched. Branches mostly free or with some anastomosing. Polyp mounds conical, prominent, or slightly raised, scattered or crowded, usually biserial and with two opposed polyp mounds at the tip of the branches. Coenenchyme usually thin. Coenenchymal sclerites mainly capstans, radiates and spindles. Thin, sharp and elongated spindles concentrated in the polyp mounds. Anthocodiae with points arrangements of bar-like rods straight or curved, frequently long. Collaret absent or of a few bar-like rods. Axis horny and flexible. Colour of the colonies red, orange, pink, or white.
Holotype.
(after
Reported for the type locality Point Loma, California.
What remain from the holotype are small pieces of branches: two fragments, 16 mm and 12 mm long, the former with 9 polyps, the latter with 5 (pers. comm. S. Cairns) (Fig.
Proposed genera for species misplaced in the genus
Species | Original author | Author |
Actual status | Proposed status | Distribution | Depth (m) |
---|---|---|---|---|---|---|
|
Riess, 1929 |
|
Tortugas, Kingston, Jamaica, Caribbean sea | 18 | ||
|
Nutting, 1909 | herein |
|
Point Loma, California, Pacific Ocean | 176 | |
|
Thomson & Simpson, 1909 | herein |
|
Andaman sea, Indian Ocean | 83–494 | |
|
Thomson, 1927 | Ofwegen, L.P. van 2014 |
|
Along Monaco, Western Atlantic | 1732 | |
|
Thomson, 1927 | Grasshoff, 1992 |
|
Iles Cap Vert, Western Atlantic | 91 | |
|
Thomson & Simpson, 1909 | herein |
|
Marble Rock, Mergui Archipelago, Andaman sea, Indian Ocean | not given |
Author who transferred the original species to another genera or our new proposed genera for the species.
[based on
Holotype: BM 1933.05.03.094, ethanol preserved, Marble Rock, Mergui Archipelago, Myanmar, Andaman Sea. No more data available.
[see also
From the type locality, Marbel Rock, Mergui Archipelago, Andaman Sea, Indian Ocean. No data available about the depth range.
The two species described in
None available.
[based on
From the type locality, Andaman sea, Indian Ocean, 83–494 m in depth.
We only have a few drawings of sclerites of this species from
Holotype: BM 1933.03.13.024, fragment, ethanol preserved, Campagne 1901, Stn. 1203:
Reported from the scientific campaigns of Prince Albert 1st de Monaco in 1901, Station 1203, along Iles du Cap Vert,
Firstly, we conclude that Verrill’s genus
Our appreciation to Leen van Ofwegen (Naturalis Biodiversity Center, Leiden, The Netherlands), Stephen Cairns (