Research Article |
Corresponding author: Odalisca Breedy ( odaliscab@gmail.com ) Academic editor: Leen van Ofwegen
© 2015 Odalisca Breedy, Hector M. Guzman.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Breedy O, Guzman HM (2015) A revision of the genus Muricea Lamouroux, 1821 (Anthozoa, Octocorallia) in the eastern Pacific. Part I: Eumuricea Verrill, 1869 revisited. ZooKeys 537: 1-32. https://doi.org/10.3897/zookeys.537.6025
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Muricea is an amphi-American genus. Verrill proposed dividing the species from the Pacific Ocean into three genera and established the genus Eumuricea for five eastern Pacific species with tubular calyces. Eumuricea is basically characterized by colonies with elongate, cylindrical calyces with truncate margins and star-like opercula, and the occurrence of unilateral spinous spindles. According to these characteristics, Eumuricea does not show enough difference from Muricea to be treated as a separate genus. Original type material of Eumuricea was morphologically analysed and illustrated using optical and scanning electron microscopy. We conclude that the eastern Pacific species should be placed in the genus Muricea and form a group characterised by tubular calyces that comprises four species at present, M. acervata, M. hispida, M. squarrosa, and M. tubigera and a dubious species M. horrida. Lectotypes were designated for M. squarrosa and M. hispida to establish their taxonomic status. The genus Eumuricea has also been misunderstood by former authors who erroneously assigned species to it. For these species we propose new combinations: Swiftia pusilla, Astrogorgia splendens and A. ramosa.
Alcyonacea, Astrogorgia, Cnidaria, eastern Pacific, Eumuricea, Muricea, Leptogorgia, plexaurid gorgonian, soft corals, Swiftia, taxonomy
Muricea Lamouroux, 1821 is an amphi-American genus with representatives in the western Atlantic and the eastern Pacific (
The genus Muricella includes species with rather thin coenenchyme, filled with long spindles, low subconical calyces arising from between the large spindles, usually standing at right angles from the axis, and covered with much smaller and shorter spindles (
The eastern Pacific Eumuricea and Muricea were separated, according to
Eumuricea was proposed by
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 Muricea Lamouroux, 1821 sensu stricto. Previous reviews dealt with Pacifigorgia Bayer, 1951 (
MNHUK
CIEMIC Centro de Investigación en Estructuras Microscópicas,
CRBMco Colección de referencia de Biología Marina
INN NAZCA Instituto de Investigaciones Marinas, Salinas, Ecuador
SEM Scanning Electron Microscopy
UCR Museo de Zoología, Escuela de Biología,
UNIANDES-BIOMMAR
UPCH Colecciones Biológicas, Universidad Peruana Cayetano Heredia, Lima, Perú
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 SEM following the standard protocol described in
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
branched spindle: a spindle with some of the processes much elongated and branchlike, often crooked.
calyx: cylindrical or wartlike projecting anthostele. In Muricea, it is mostly formed by the same type as the outer coenenchyme sclerites. They are arranged as a fringe around the border of the calyx with the sharp or spiny processes projecting outwards.
calyx shelf-like: calyx with prolonged lower border, polyp opens upwards.
calyx tubular: tube-like calyx with mostly even, truncate borders, polyp opens straight and distally.
club: monoaxial sclerite enlarged at one end, the head, and tapered at the other end, the handle. According to the modification of the head they are classified in leaf, thorn, wart or torch. Muricea, the heads are mostly ornamented by sharp, thorn-like or spine-like processes, and the handles are warty, or of an almost smooth surface, curved or straight.
unilateral spinous spindles: sclerites with asymmetrically arranged warts on the surface. Inner side with low composite warts, close together or very crowded, their processes often anastomose. Outer side with fewer cone-shaped tubercles, long projecting in some cases. Ends of the sclerites blunt, acute or both.
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 | Leptogorgia |
– | 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 |
2 | Coenenchyme contains unilateral spinous sclerites, polyps retract into shelf-like or tubular calyces | Muricea |
– | Coenenchyme does not contain unilateral spinous sclerites, polyps retract into prominent or slightly raised dome-shaped polyp mounds | 3 |
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 | Astrogorgia |
– | Coenenchymal sclerites mainly capstans, radiates and spindles, thin, sharp, with tubercles, some modified as incomplete disks, but not heavily ornamented | Swiftia |
Muricea Lamouroux, (pars) 1821: 36; Blainville (pars)
Eumuricea (pars)
Eumuricea (Muricea)
Muricea spicifera Lamouroux, 1821, by subsequent designation: Milne Edwards and Haime 1850. [M. spicifera was later synonymised with Muricea muricata (Pallas, 1766) apud
Diagnosis (based on
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 Muricea species. The shape of calyces shelf-like or tubular, and related features as being imbricate or sparse show many intermediate forms. In the tubular-calyces species group the apical branches show a closer arrangement of calyces and smaller projecting angles in respect to the branch than at the lower branches. Therefore, the strongest character that separates Muricea from other genera is the type of sclerites.
Muricea acervata Verrill, 1866: 327–328; Rossi 1955;
Muricea (Eumuricea) acervata Verrill, 1869a: 419–421.
Eumuricea acervata Kükenthal, 1924: 143.
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
Muricea hispida Verrill, 1866: 328;
Muricea (Eumuricea) hispida Verrill, 1869a: 422–423.
Eumuricea hispida Kükenthal, 1924: 151–152;
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 M. acervata with a minimal description. They both were properly described in 1869a. Muricea hispida was described from two specimen fragments from Panamá. Muricea hispida is similar to M. squarrosa and M. tubigera. These three species have long tubular calyces, similar colour and shape of the colonies. The main difference that separates them is the calyx length M. tubigera with the largest and M. squarrosa with the shortest (Table
Comparative features of the eastern Pacific genus Muricea Lamouroux, 1821. Diameter of the branches is including calyces; size of the sclerites and other measurements are based on type material examined in this study. () Represents Verrill max size of sclerites. All measurements are given in mm.
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 |
---|---|---|---|---|---|---|---|---|---|---|---|---|
M. acervata | b | cand | 70 | 7 | T | 2.50 | 2 | cl | 21 | 1.8 (2) | 0.15–0.30 | lb, do, py, w |
M. hispida | 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 | - | - |
M. squarrosa | lb-b | bu | 40 | 5 | mt | 2.6 | 1.75 | s | 14 | 1.3 (1.8) | 0.14–0.30 | py, lb, w, c |
M. tubigera | lb | cand | 70 | 8 | mt | 5 | 0.70 | cl | 26 | 2.0 (2.28) | 0.12–0.46 | w, c |
Muricea horrida Möbius, 1861: 11–12;
Muricea (Eumuricea) horrida Verrill, 1869a: 423.
Eumuricea horrida Kükenthal, 1924: 151.
Plate 3, figs 5–8 (
Holotype figured. According to
(after
Reported for Perú, the type locality.
According to
Muricea (Eumuricea) squarrosa Verrill, 1869a: 423–424.
Eumuricea squarrosa Kükenthal, 1924: 159.
Muricea squarrosa Harden, 1979: 159–160.
Lectotype (here designated):
COSTA RICA: UCR 587, dry, Pitaya Beach, Guanacaste, Pacific coast, Costa Rica, 20–23 m, J. Cortés, 16 June, 1991; UCR 1742, ethanol preserved, Bajo Negro, Marino Ballena National Park, 25 m, O. Breedy,13 April 2008; UCR 2261, ethanol preserved, Isla Larga Oeste, Manuel Antonio National Park, 19 m, O. Breedy and H. Guzman, 6 February 2012; UCR 2262, ethanol preserved, Isla Larga, Manuel Antonio National Park, 25 m, O. Breedy and H. Guzman, 7 February 2012; UCR 2396, ethanol preserved, Marino Ballena National Park, 25 m, O. Breedy, 27 April 2002; UCR 2410; 2414, ethanol preserved, La Danta, Santa Elena Bay, 35 m, O. Breedy and Minor Lara, 10 August 2014; UCR 2418–2419, ethanol preserved, Bajo Mixta, Golfo Dulce, 21 m, O. Breedy and H. Guzman, 7 February 2009 ECUADOR: IIN 25, dry, Bajo Lunes, Reserva de Producción Faunística Marino Costera Puntilla de Santa Elena, 18 m, P. Martínez, F. Rivera, R. Nabot and O. Breedy, 21 July 2010; IIN 47, dry, Gigima, Reserva de Producción Faunística Marino Costera Puntilla de Santa Elena, 14 m, P. Martínez, F. Rivera, R. Nabot and O. Breedy, 22 July 2010. PANAMÁ:
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 M. hispida and M. tubigera is that the calyces are shorter and more distantly placed. Other differences were discussed above (under M. hispida).
Muricea (Eumuricea) tubigera Verrill, 1869a: 421–422.
Eumuricea tubigera Kükenthal, 1924: 150.
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.
Gorgonia exserta Ellis & Solander, 1786, by monotypy.
Eumuricea pusilla Nutting, 1909: 718–719;
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 Eumuricea Verrill, 1869.
Species | Original author | Author |
Actual status | Proposed status | Distribution | Depth (m) |
---|---|---|---|---|---|---|
E. atlantica | Riess, 1929 | Muricea Lamouroux | Tortugas, Kingston, Jamaica, Caribbean sea | 18 | ||
E. pusilla | Nutting, 1909 | herein | Eumuricea | Swiftia Duchassaing & Michelotti, 1864 | Point Loma, California, Pacific Ocean | 176 |
E. ramosa | Thomson & Simpson, 1909 | herein | Eumuricea |
Astrogorgia Verrill, 1868 |
Andaman sea, Indian Ocean | 83–494 |
E. rigida | Thomson, 1927 | Ofwegen, L.P. van 2014 | Thesea | Along Monaco, Western Atlantic | 1732 | |
E. rugosa | Thomson, 1927 | Grasshoff, 1992 | Leptogorgia ruberrima | Iles Cap Vert, Western Atlantic | 91 | |
E. splendens | Thomson & Simpson, 1909 | herein | Eumuricea | Astrogorgia Verrill, 1868 | Marble Rock, Mergui Archipelago, Andaman sea, Indian Ocean | not given |
Astrogorgia Verrill, 1868b: 414;
Muricella Kükenthal, 1924: 169.
Acanthomuricea Fabricius & Alderslade, 2001: 212.
Astrogorgia sinensis Verrill, 1868b by monotypy.
[based on
Eumuricea splendens Thomson & Simpson, 1909: 258–259.
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 Eumuricea by
Eumuricea ramosa Thomson & Simpson, 1909: 260–261.
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
Gorgonia ruberrima W. Koch, 1886: 14–18.
Eumuricea rugosa
Leptogorgia monodi Stiasny 1937: 309.
Leptogorgia ruberrima Stiasny, 1940: 361; Grasshoff 1988: 111;
Holotype: BM 1933.03.13.024, fragment, ethanol preserved, Campagne 1901, Stn. 1203: 15°54’ N, 22°54'45"E, Iles du Cap Vert, 91 m, 18 August 1901.
(see also
Reported from the scientific campaigns of Prince Albert 1st de Monaco in 1901, Station 1203, along Iles du Cap Vert, 15°54’ N, 22°54'45"E, Western Atlantic.
Firstly, we conclude that Verrill’s genus Eumuricea was misinterpreted by former authors who erroneously assigned this genus to a diverse group of species. Secondly, that Eumuricea corresponds to a group of Muricea with tubular calyces as Verrill proposed, but because all other characteristics are found in Muricea, we do not consider the shape of the calyx as a sole character to separate this genus, since there are at least three more types of calyx structure within Muricea. And thirdly, the Muricea group with tubular calyces comprises four valid species M. acervata, M. hispida, M. squarrosa and M. tubigera, and a dubious one, M. horrid, reported for the eastern Pacific. It is intriguing that Muricea squarrosa is the only species in the genus that recently has been collected at various localities along the eastern Pacific. Although extensive surveys have been conducted along the Pacific coast of Panama and Peru (main type localities), the other species have not been found. Presently, we do not have data to support the idea of local extinction of species, but it is very likely that changing oceanographic conditions could have affected octocoral diversity. Perhaps more survey effort along the eastern Pacific would give further information about this genus. Presently, we recommend taking into account the status of the species of Eumuricea for biodiversity records and assessments.
Our appreciation to Leen van Ofwegen (Naturalis Biodiversity Center, Leiden, The Netherlands), Stephen Cairns (