Research Article |
Corresponding author: Odalisca Breedy ( odaliscab@gmail.com ) Academic editor: Leen van Ofwegen
© 2016 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 (2016) A revision of the genus Muricea Lamouroux, 1821 (Anthozoa, Octocorallia) in the eastern Pacific. Part II. ZooKeys 581: 1-69. https://doi.org/10.3897/zookeys.581.7910
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The species of the genus Muricea were mainly described from 1846 to 1870. After that very few contributions were published. Although the highest richness of Muricea species is in the eastern Pacific shallow waters, a comprehensive systematic study of the genus does not exist. Recently we started a taxonomic review of the genus in order to validate the status of four species previously included in the genus Eumuricea. Herein we present the second part of the Muricea revision dealing with the species-group characterised by shelf-like calyces instead of tubular-like calyces (the M. squarrosa-group). Original type material was morphologically analysed and illustrated using optical and scanning electron microscopy. Comparative character tables are provided for the genus. The taxonomic status of the species was analysed and established by designating lectotypes, alternatively by recognising a holotype by monotypy. We conclude that the genus Muricea comprises 20 valid species, including the previous four in the M. squarrosa-group. We propose 10 lectotypes, a new combination and three more species groups for the genus based on morphology: the M. fruticosa-group, M. plantaginea-group and M. austera-group.
Alcyonacea , Cnidaria , eastern Pacific, Muricea , plexaurid gorgonian, soft corals, taxonomy
The genus Muricea Lamouroux, 1821 is widely distributed along the eastern Pacific shallow waters (less than 40 m deep), and although it has representatives in the western Atlantic, the highest species richness is attained in the eastern Pacific. We deal with the eastern Pacific Muricea species in the present taxonomic review. The first record appeared as Gorgonia plantaginea Valenciennes, 1846 (Plate 15, Fig.
Recently, we published the first part of the review of Muricea (
The present research represents the second part of the fifth review, in a series of six proposed, aiming to evaluate the genera of gorgonians historically reported for the shallow eastern Pacific waters. Previous reviews dealt with Pacifigorgia Bayer, 1951 (
CASIZ
California
CDRS
CIEMIC
Centro de Investigación en Estructuras Microscópicas,
CRBMco
Colección de referencia de Biología Marina
ICZN International Code of Zoological Nomenclature
INN NAZCA Instituto de Investigaciones Marinas, Salinas, Ecuador
MNHUK
SEM Scanning Electron Microscopy
UCR / MZUCR
Museo de Zoología,
UNIANDES-BIOMMAR
UPCH Colecciones Biológicas, Universidad Peruana Cayetano Heredia, Lima, Perú
NMNM
National
ZMHC
The type specimens and comparative reference material used in this study were analysed during visits to museums or acquired on loan from the CASIZ,
The microscopic study was done at the CIEMIC, the specimens were prepared for SEM following the protocol described in
Data on geographical distribution are from our personal collections, museum catalogues and published monographs. Verrill and the other authors who described Muricea species (Aurivillius, Hickson Deichmann and Valenciennes) did not designate holotypes and in cases, the descriptions could often fit several species. In some cases, only one specimen in the collection was under a species name that automatically constitutes the holotype by monotypy. When the status was unclear we designated lectotypes to establish the identity of the poorly defined species and to avoid future confusion.
The terminology is according to
Comparative character tables are provided for the Muricea species (Tables
Sclerite comparison for the eastern Pacific Muricea species. Measurements given are from the holotypes and paratypes, in mm.
Species | Sclerite colours | Anthocodial sclerite colours | Dominant type of coenenchymal and calycular sclerites | Coenenchymal and calycular spindles maximum size | Anthocodial maximum size |
---|---|---|---|---|---|
M. fruticosa | w, rb, py | yellow, w | uss | 2.0×0.5 | 0.64×0.1 |
M. formosa | w | w | uss | 1.5×0.25 | 0.175×0.075 |
M. aspera | pb, w | w | uss | 1.35×0.35 | 0.25×0.04 |
M. echinata | o, lb | lo, w | uss | 2.4×0.34 | 0.25×0.05 |
M. galapagensis | amb, lo | lo, w | uss | 4.1×0.75 | 0.25×0.06 |
M. plantaginea | rb, amb/w | lo, lb/w | ls | 1×0.2 | 0.25×0.08 |
M. californica | ro,ly,amb | lo | ls | 0.54×0.2 | 0.23×0.06 |
M. mortensenii | w | w | s | 0.7×0.12 | 0.21×0.08 |
M. austera | rb,o,ly | w | uss | 1.5×0.5 | 0.36×0.06 |
M. albida | w | w | uss | 1.4×0.05 | 0.25×0.03 |
M. crassa | rb | lb, w | uss | 2.5×0.7 | 0.22×0.04 |
M. retusa | rp, p,o | o, ly | uss | 1.2×0.6 | 0.4×0.1 |
M. purpurea | dr, ro | ro | ls | 0.7×0.3 | 0.3×0.055 |
M. hebes | py | py | uss | 0.83×0.20 | 0.45×0.1 |
M. nariformis | bo,lb | o | ls | 0.52×0.28 | 0.17×0.035 |
M. robusta | o, bo, lb | o | ls | 0.64×0.26 | 0.15×0.05 |
Comparative features of the eastern Pacific Muricea species. Measurements given are from the holotypes and paratypes, in mm.
Species | Colony colour | Colony shape | Branching pattern | Length of unbranched terminal branchlets | Diameter of end branchlets (mm) | Coenenchyme | Calyx height at branchlets | Calyx arrangement at branchlets |
---|---|---|---|---|---|---|---|---|
M. fruticosa | rb, w, bi | bu | irr | 15–40 | 3–6 | t | 1–1.2 | c |
M. formosa | w | bu | irrlat, dich | 28 | 6–9 | t | 2.8–3 | c |
M. aspera | lb | fla ? | irr, lat | 6–30 | 4–5 | t | 1–2 | c, slightly imbr |
M. echinata | rb | bu | irr, lat | 6–30 | 5–8 | t | 2.8–3 | c |
M. galapagensis | lo | fall | lat | 80 | 1.6–3 | t | 0.6–1 | s |
M. plantaginea | db/w | fla | irr, lat | 10–50 | 2–3 | t | 0.7–1.2 | c, imbr |
M. californica | ro | bu | irr, lat | 0.5–2.8 | 3–3.2 | mt | 1.1–1.9 | c, slightly imbr |
M. mortensenii | py | fla | irr | 2–4 | 2–3 | t | 0.7–1 | c |
M. austera | rb | bu | dich, lat | 50 | 7–8 | T | 1.7–2 | c |
M. albida | w | cand | dich | 11 | 5–7 | T | 0.8–1.8 | c, slightly imbr |
M. crassa | db | bu | dich, lat | 70 | 7–10 | T | 2.7–3 | c |
M. retusa | rp | - | dich | 50 | 7–8 | T | 1–1.5 | c |
M. purpurea | rp | bu | dich | 50–80 | 9–11 | T | 1.5–1.8 | c, slightly imbr |
M. hebes | yb | fing | dich | 32 | 6–9 | T | 1–1.8 | c, slightly imbr |
M. nariformis | bo | fing | dich | 24 | 57 | T | 0.8–1.2 | c |
M. robusta | bo | bu | dich | 70 | 7–8.5 | T | 0.7–1.2 | c |
unilateral spinous spindles: spindle 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.
prickly spindle: warty, irregular spindle with prickly, pointed processes at one tip.
leaf-like spindle: warty spindles with flat, leaf-like processes, terminal or laterally placed.
spinous club-like spindle: club shaped sclerite, with head ornamented by thorny or leafy, sometimes unilaterally placed processes and with warty, thin handle.
The most informative characters for the genus are the branch diameter, the colours (colonies and sclerites) and the type, size and combination of sclerites. The colours, although some variation exist, are very constant and a reliable characteristic (
Presently, the boundaries among octocorals species are based on common characters, basically colony and sclerite shapes and colours that could be influenced, at certain point, by the environment. For that, in determining an octocoral morpho-species the combination of these morphological characteristics results in a more accurate assessment. Furthermore, analysis of several specimens aid in identifying intraspecific variation. Field observation and evaluation of habitat in terms of oceanographic conditions could provide information in the decision making process.
Muricea Lamouroux, (pars.) 1821: 36; Blainville (pars.) 1834: 509; Ehrenberg (pars.) 1834: 134;
Eumuricea (pars.) Verrill, 1869: 449;
Muricea spicifera Lamouroux, 1821, by subsequent designation:
(based on
From Cape Hatteras, North Carolina to Brazil, including Bahamas, Greater and Lesser Antilles, Gulf of México, and Caribbean islands (
Based on calyx morphology,
Muricea fruticosa Verrill, 1869.
Muricea fruticosa Verrill, 1869: 428;
Muricea fruticosa typica
Muricea fruticosa var. miser Verrill, 1869: 430;
Thesea crosslandi Hickson, 1928: 354–356 (syn. n.).
Pseudothesea crosslandi (Hickson, 1928);
Lectotype (here designated).
The lectotype is a large, bushy colony 35 cm tall, and about 45 cm wide. Four main branches, 25–35 mm in diameter, somewhat flattened, arise from an irregular, 52 mm diameter holdfast. The holdfast is spreading and raised about 30 mm above substrate, the specimen is attached to a plaster base for a past years museum display (Fig.
The colony is bicoloured, reddish brown at the tips, fading to a light yellow towards the base (Fig.
There are two colour patterns in the syntype series, a marked bicoloured pattern, with dark reddish tips and whitish to pale yellow stems (Fig.
From México to Perú. Type locality, Pearl Islands, Panamá.
In order to establish the identity of this species, the
COSTA RICA: UCR 482, 486, dry, Punta Conejo, Herradura, Puntarenas, 10 m, J. Cortés, 21 September 1996; UCR 520 (3), dry, Nicoya Gulf, CJ Kalb, 2 March 1967; UCR 576, dry, San Juanillo, Guanacaste, 12.5 m, J. Cortés, 14 June 1991; UCR 588, dry, Pitahaya Beach, Guanacaste, 20–23 m, J. Cortés, 15 June 1991; UCR 837-839, dry, Ballena Bay, Nicoya Gulf, 40 m, R/V Victor Hensen, 2 December 1993. ECUADOR: CASIZ 105032, ethanol preserved, Santa Cruz Island, Nameless Islet, Galápagos Islands, 20 m, P. Humann, no collection date. CDRS 03-76, ethanol preserved, Los Hermanos, Galápagos Islands, 9 m, C. Hickman, 17 January 2003. CDRS 06-33, ethanol preserved, Nameless Island, Galápagos Islands, 9–10 m, C. Hickman, 25 May 2006. IIN 7, 9, dry, Tambip, Salinas, 12–14 m, F. Rivera, P. Martínez, 20 July 2010; IIN 21, dry, Bajo Lunes, Salinas, 18 m, F. Rivera, P. Martínez, 21 July 2010; IIN 34, 69, Gigima, Salinas, 12–14 m, F. Rivera, P. Martínez, 22 July 2010; IIN 88, 89, 123, dry, Los Ahorcados, Machalilla National Park, 10–12 m, F. Rivera, P. Martínez, 25 July 2010. EL SALVADOR: UCR 1938, ethanol preserved, Departamento la Libertad, Playa Mizata, J. Segovia, 27 February 2010. MÉXICO: CASIZ 097734, ethanol preserved, Roca Partida, South side, Revillagigedo Islands, 36 m, R.J. Van Syoc, M/V “Royal Star, Clipperton Island Expedition 1994, 2 May 1994. CASIZ 103387, ethanol preserved, Boca del Tule to Arena Blanca, Baja California Sur, 30 m, W. Lee, J. Moran, J. McCosker, 26–27 April 1976; CASIZ 100843, ethanol preserved, Roca Alejos, Baja California Sur, 18–33, Robert Van Syoc, Cordell Expeditions, 5 November 1990.
Muricea formosa Verrill, 1869: 434–436;
Holotype.
The holotype is a 10 cm long and 6 cm wide colony, branching lateral, in one plane and irregularly dichotomous (Fig.
Colour of the colony is white.
The species has been found living on rocky bottoms, caves and outcrops at 10–13 m in depth. The colonies are mostly growing in one plane, but in some cases they extend in two or three planes (Y. Hooker pers. comm.) (Fig.
Reported from Las Ánimas Islet, Gulf of California by
MÉXICO. M 18, dry, Punta Tiburón, Kino Bay, Sonora, 5.5 m, J.L. Carballo, 11 October 1999. PERÚ. CZA 286, dry, Canoas de Punta Sal, 12 m, Y. Hooker, 2 July 2011. CZA 412–416, 419, 424, dry, Canoas de Punta Sal, 13 m, Y. Hooker, 13 August 2012. CZA 286, Canoas de Punta Sal, 13 m, Y. Hooker, 2 July 2011. CZA 417–418, 420–423, Cabo Blanco, Piura, Y. Hooker, 13 August 2012. CZA 425, dry, El Ñuro, Piura, 10 m, Y. Hooker, 8 August 2012.
Muricea aspera Verrill, 1869: 448–449;
Lectotype.
Paralectotypes. PANAMÁ:
Type series is comprised of fragments, the larger ones are 12 cm tall and 9 cm wide, and 8.5 cm tall and 4.5 cm wide, probably fragments of a larger specimen (
The lectotype is 12 cm tall and 9 cm wide (Fig.
Colour of the colony is light brown.
Only reported for the type locality at extreme low waters (according to
PANAMÁ:
Muricea echinata Verrill, 1866: 328 (pars.);
Muricea echinata var. flabellum Verrill, 1869: 427–428;
not Eunicea echinata Valenciennes, 1855: 13 (nom. nud.);
not Muricea echinata Milne Edwards & Haime, 1857: 143
Lectotype: YPM1565d, dry, Pearl Islands, Panamá, F.H. Bradley, 1866.
Paralectotypes: PANAMÁ:
The lectotype is a colony 8.5 cm long and 7 cm wide, branching lateral and irregular and spreading in almost one plane (Fig.
Colour of the colony is reddish brown.
The examined colonies are up to 10 cm tall and up to 14 cm wide, branching irregular, spreading in almost one plane. Calyces can be up to 3.5 mm long. Colour of the colonies varies to a deeper orange hue.
Muricea echinata was erroneously mentioned by
COSTA RICA:
Muricea (?) galapagensis Deichmann, 1941: 6–9;
Holotype.
The holotype is a 10 cm tall and about 5 mm wide colony, it has one broken branch and according to Deichmann’s illustration (
The colony is light orange (Fig.
Reported only from the type locality, Elizabeth Bay, Albemarle Island, Galapágos Islands, Ecuador.
The Muricea fruticosa group comprises five species: M. fruticosa, M. formosa, M. aspera, M. echinata and M. galapagensis. This species-group is characterised by thin branches, long, prickly calyces and outer coenenchyme and calyces with unilateral spinous spindles (Tables
Gorgonia plantaginea Valenciennes, 1846: pl 15.
not Gorgonia plantaginea Lamarck, 1815: 163 (Antilles).
not Eunicea plantaginea Valenciennes, 1855: 13;
Eunicea tabogensis Duchassaing & Michelotti, 1864: 17.
Muricea appressa Verrill, 1864: 37;
Muricea appressa var. flavescens Verrill, 1869: 446;
Muricea tenella Verrill, 1869: 446–448;
Holotype.
Muricea appressa:
The holotype is a large, flabelliform colony 40 cm tall and 18 cm wide. The colony is in bad shape, the main branches are almost nude and branchlets are bent to the sides (Fig.
Colour of the colony is deep brown.
Verrill´s type series is very consistent in all characters with respect to the Valenciennes’ holotype. For example, the specimen
The species is widespread throughout the central archipelago of the Galápagos (
A widespread distribution, it has been reported from México to Perú, at a depth range from 10–65 m including the oceanic islands, Galápagos (Ecuador) and Revillagigedos (México). Type locality: Mazatlán, México.
The species appears for the first time in Valenciennes’ book (
COSTA RICA: UCR 591, dry, Pitahaya Beach, Guanacaste, 23 m, J. Cortés, 15 June 1991; UCR 634, dry, Las Cocineras, Santa Elena Bay, Guanacaste, at the beach, O. Piedra B., 20 February 1965; UCR 945, dry, Caño Island, 22 m, H. Guzman, 11 February 1984.
MÉXICO: CASIZ 099631, 097735, ethanol preserved, Roca Partida, Revillagigedo Islands, 36 m, R.J. Van Syoc, M/V “Royal Star, Clipperton Island Expedition 1994, 2 May 1994.
Muricea californica Aurivillius, 1931: 111–114;
Lectotype. USA:
The lectotype is a bushy colony 5.6 cm tall and 7.8 cm wide. Two main branches, diameter 4.6–5 mm, arise from an oval holdfast, 1.38 cm in diameter (Fig.
Colour of the colony is reddish orange.
According to
South of point Conception, California to Santa María, Baja California, Mexico (
The species was originally described by
MÉXICO: Geoff1, dry, Baja California, Geoff Shester, 2007. Gorgonia 11, dry, Isla Tiburón, San Carlos Bay, Sonora, 5–25 m, J.L. Carballo, 27 April 2001. PANAMÁ: BM 30.6.17.18 (fragment), (erroneously identified as M. hebes), ethanol preserved, off Panamá, low tide, St. George, Scientific Expedition, Pacific Cruise, C. Crossland, 1923–1924.
Muricea mortensenii Hickson, 1928: 369–371;
Holotype.
(see also
Colour of the colony is pale yellow.
Reported for the type locality, Rey Island, Pearl Islands, Panamá.
The Muricea plantaginea group comprises three species: M. plantaginea, M. mortensenii and M. californica. The group is characterised by thin coenenchyme, thin branches and the lack of unilateral spinous spindles (as defined for the genus). The main components of the calyces and outer coenenchyme are leaf-like spindles with prominent spines and elongated warty spindles. Muricea mortensenii is conspicuously different from the others in the profuse branching, very crowded upwards-curved calyces and soft texture of the colony. Muricea californica and M. plantaginea (typical) have similar colony and sclerites colours. Respect to the colony morphology, M. californica is not as fan-shaped as M. plantaginea. Colonies could develop in various planes. Muricea californica differs also from M. plantaginea in having shorter spindles, no more than 0.55 mm long, and longer, not imbricate calyces (Tables
Muricea austera Verrill, 1869: 430–432;
Lectotype:
The lectotype is a bushy colony, 20 cm long and 23 cm wide, with brittle coenenchyme that is partially lost on some branches and with mostly naked terminal ends (Fig.
Colour of the colony is reddish brown.
The colonies ramify producing a bouquet-like structure (Fig.
It has a widespread distribution, from México to Perú. Type locality: Pearl Islands, Panamá.
Verrill (1868) described this species with specimens from Pearl Islands, Cape San Lucas and La Paz, Baja California. Later,
Verrill did not designate a holotype for the species. The specimen
COSTA RICA: UCR 623–624, 633a, dry, Bajo Isla Chora, Sámara Bay, Guanacaste, 10 m, H. Guzman, 18 March 1984; UCR 779, dry, Peñón de la Bruja, Santa Rosa National Park, 10 m, J. Cortés, 5 March 1994; UCR 836, dry, Cabo Blanco Islet, SW from Cabo Blanco National Reserve, Puntarenas, 12 m, L. Mena, 26 April 1994. ECUADOR: IIN 4b, 5, 6, dry, Tambip, Salinas, 12–14 m, F. Rivera, P. Martínez, 20 July 2010; IIN 23, dry, Bajo Lunes, Salinas, 18 m, F. Rivera, P. Martínez, 21 July 2010; IIN 62, dry, Gigima, Salinas, 12–14 m, F. Rivera, P. Martínez, 22 July 2010; IIN 104, 118, dry, Los Ahorcados, Machalilla National Park, 10–12 m, F. Rivera, P. Martínez, 25 July 2010. EL SALVADOR: UCR 1936, ethanol preserved, Departamento la Libertad, Playa Mizata, J. Segovia, 26 February 2010. MÉXICO: M 11, dry, León echado Island, San Carlos Bay, Sonora, 5–25 m, J.L. Carballo, 6 December 2000; M 29, dry, Lobos Island, Mazatlan Bay, Sinaloa, 5–25 m, J.L. Carballo, 17 November 1998.
Muricea robusta (pars.) Verrill, 1866: 329.
Muricea albida Verrill, 1868b: 412;
Lectotype.
The lectotype is a 15 cm long and 12 cm wide, candelabrum-like colony, branching in one plane and mostly dichotomous (Fig.
Colour of the colony is white.
It is known from the type locality, Pearl Islands, and reported from Taboguilla Island. However, M. albida has not been found during our recent extensive surveys along Pacific Panamá.
The species was described by
PANAMÁ: BM 1946.1.14.43, off Taboguilla, 9 m, C. Crossland, 22 November 1915.
Muricea crassa Verrill, 1869: 425–426;
Lectotype:
The lectotype is a large bushy colony, 40 cm long and 50 cm wide. The holdfast is irregular, and swollen, about 7 cm wide and 4 cm high covering one side of the rock remains to which is attached. The branching is irregularly dichotomous, mostly lateral, in one plane (Fig.
Colour of the colony is dark brown.
The examined colonies are up to 50 cm tall and up to 40 cm wide, but smaller specimens, around 30 cm tall are the majority. The branch tips could reach up to 15 mm in diameter, and the unbranched terminal ends up to 13 cm long. This species is very conspicuous because of the dark colour and strong and prickly branches with wide terminal tips. We have observed M. crassa at various localities in Panamá, from 3–12 m deep around Pearl Islands, Gulf of Panamá and the deepest records are around Coiba Island in the Gulf of Chiriquí, 20–30 m deep. It is not abundant and is sparsely distributed in patches dominated by other species.
It has a widespread distribution from México to Perú. Type locality, Pearl Islands, Panamá.
ECUADOR: IIN 94, 121, dry, Los Ahorcados, Machalilla National Park, 10–12 m, F. Rivera, P. Martínez, 25 July 2010. PANAMÁ:
Muricea retusa Verrill, 1869: 432–434;
Holotype.
The holotype is a 7 cm long and about 6 cm wide colony. It is attached to the holdfast of a large M. fruticosa colony (Fig.
Colour of the colony is reddish purple.
It is only known from the type locality, Pearl Islands, Panamá.
Muricea hebes (pars.) Verrill, 1864: 36.
Muricea purpurea
Muricea purpurea var. nigra Hickson, 1928: 367 syn. n.
Muricea rubra Aurivillius, 1931: 108–109 syn. n.
Lectotype.
Panamá: BM 1946.1.14.44, dry, off Taboguilla Island, 9 m, T. Mortensen, 27 November 1915.
The lectotype is a 22 cm long and 21 cm wide colony with branching in one plane and mostly dichotomous (Fig.
Colour of the colony is reddish purple.
Verrill’s type collection is composed of specimens from Panamá and Acapulco, México. The description of M. purpurea was mostly based on the Panamá specimens (
The colonies are found on rocky substrates, mostly vertically placed or upside down in caves. They also occur in crevices at rocky bottoms and grow straight up. The colonies can extend along the substrate by spreading holdfast up to 30 cm long producing separate branches forming large colonies. When alive, polyps look, or greenish yellow (Fig.
It is a widespread species distributed from México to Perú. The species has been reported for Acapulco, México; Corinto, Nicaragua; Ecuador (
Type locality, Pearl Islands, Panamá.
The species was first mentioned by
There are two other related species M. rubra and M. purpurea var. nigra.
COSTA RICA. UCR 479a, dry, Herradura Beach, 10 m, J Cortés, 2 September 1983; UCR 510, 632a, dry, Sámara Beach, Guanacaste, 12 m, H. Guzman, 18 March 1984; UCR 800, dry, Olocuita Islet, Manuel Antonio National Park, Puntarenas, 8 m, J Cortés, 2 July 1995; UCR 1620, ethanol preserved, Carrillo Beach, Guanacaste, 10 m, J Cortés, 2006; UCR 1693, ethanol preserved, Salinas Bay, Guanacaste, O Breedy, 7 December 2006. ECUADOR: IIN 20, dry, Tambip, Salinas, 12–14 m, F. Rivera, P. Martínez, 20 July 2010; IIN 43, 48, dry, Gigima, Salinas, 12–14 m, F. Rivera, P. Martínez, 22 July 2010; IIN 99, 117, 119, 120, dry, Los Ahorcados, Machalilla National Park, 10–12 m, F. Rivera, P. Martínez, 25 July 2010; IIN 129, dry, Salango Island, Machalilla National Park, 12–25 m, F. Rivera, P. Martínez, 25 July 2010. PANAMÁ:
Muricea hebes (pars.) Verrill, 1864: 36.
Muricea hebes
Lectotype.
The lectotype is 6 cm long and 9 cm wide (Fig.
Colour of the colony is yellowish brown.
Found in México in Pájaros Island and reported for Cabo Pulmo, Gulf of California (according to
The species was created by Verrill in 1864 with specimens from Acapulco, México collected by Agassiz. Later
MÉXICO: M 61, dry, Pájaros Island, Mazatlan Bay, Sinaloa, 5–25 m, J.L., 3 February 1999.
Muricea nariformis Aurivillius, 1931: 109–111.
Holotype.
The holotype consists of two branches, about 6 cm long and 3 cm wide each (Fig.
Unknown.
The species was described by
Muricea robusta Verrill, 1864: 36;
Muricea robusta (pars.)
Lectotype.
The lectotype is a 20 cm long and 10 cm wide colony with partially broken coenenchyme on some branches and with three naked distal axes (Fig.
The revised specimens are consistent with the lectotype.
Found in México in Venado Island by J.L. Carballo and reported for Cape San Lucas (
The species was first mentioned by Verrill in 1864 and formerly described in 1869 with a specimen from Acapulco. We believe that Verrill’s type series are all fragments of the same colony; however, because this is not clear, herein we designated the specimen
COLOMBIA:
The Muricea austera-group comprises eight species: M. austera, M. albida, M. crassa, M. retusa, M. purpurea, M. nariformis, M. hebes and M. robusta. The group is characterised by thick coenenchyme, thick branches and stout colonies that can be bushy, finger-like or candelabrum-like. Species comparisons are in Tables
The second group contains white colonies, including only M. albida. In this group (Table
The third group contains reddish purple colonies, including M. retusa and M. purpurea. These species are similar also in the dichotomous branching, the arrangement of the calyces and the calyx sclerites (in rows) (Tables
We conclude that the genus Muricea in the eastern Pacific comprises 20 valid species that could be separated by their morphological characters into four groups: the M. squarrosa-group represented by four species with tubular calyces (
Our appreciation to Stephen Cairns, Leen van Ofwegen and Gary Williams for critical comments and suggestions that improved our manuscript, and Phil Alderslade for his advice solving taxonomic conflicts. We acknowledge the following people and institutions for their generosity in making available the specimens and information used in this study,