Sea anemones (Cnidaria: Actiniaria, Corallimorpharia, Ceriantharia, Zoanthidea) from marine shallow-water environments in Venezuela: new records and an updated inventory

This study compiles the diversity of sea anemones in different shallow habitats (i.e. rocky shores, coral reefs, mangroves and sandy bottoms) in several locations of Venezuela, including the most important marine reserves of Venezuela: Morrocoy National Park, Mochima National Park, Archipiélago de Los Roques National Park, Laguna de La Restinga National Park, Isla de Cubagua, and Chichiriviche de la Costa. Twenty-six species of sea anemones sensu lato were documented, from which only two actiniarian species are newly recorded in Venezuela. In addition, specimens of the scientific collection of the Museo Oceanológico Hermano Benigno Román, of Isla de Margarita, Venezuela, were examined and its taxonomic status updated. The diversity of sea anemones in Venezuela is updated to 44 species. An illustrated guide of living specimens and short descriptions of the external anatomy of the specimens is included to facilitate sea anemone species identification in the field and to provide a base line for ecological studies that require accurate identification to species level.


Background
Sea anemones are among the cnidarian groups in benthic assemblages of the Caribbean Sea, being found in a wide range of coastal habitats, including seagrass meadows, rocky bottoms, coral rubble, coral reefs, sandy patches, mangrove zones and artificial substrates. Although several inventories of sea anemones species have been made from many coastal and coral reef localities along the Caribbean Basin and Gulf of Mexico (González-Muñoz et al. 2012, many areas remain poorly studied, such as the Caribbean coast of Venezuela (Losada & Pauls 2003). Records of sea anemone species for Venezuela are scattered in studies of different nature, such as taxonomy (Agudo 1987;den Hartog 1980), biological and ecological interactions (Bastidas & Bone 1996;Bitter-Soto 1999;Briggs et al. 1996;Henríquez & Palacios 2008;Losada et al. 1988), feeding behavior (Liñero-Arana & González 2008), and descriptions of crustacean species engaged in symbiotic associations with sea anemones (Knowlton & Keller 1985).
The most extensive reference for sea anemones in Venezuela is the monograph of Agudo (1987), which includes the records and distribution of 47 species along the main coastal and insular localities (Losada & Pauls 2003). However, the taxonomic status of some of the species reported by Agudo (1987) has changed through time (e.g. ). Therefore, an updated status of the sea anemone diversity for the region is necessary.
In this study we examined specimens of sea anemones sensu lato (i.e. orders Actiniaria, Ceriantharia, Corallimorpharia, and Zoanthidea) sampled during November 2012 in several shallow coastal and coral reefs environments along six locations, four of which are marine reserves in Venezuela (Fig. 1). Additionally, we examined the sea anemone specimens previously deposited in the collection of the Museo Oceanológico Hermano Benigno Román (MOHBR), of the Estación de Investigaciones Marinas at the Fundación La Salle de Ciencias Naturales, in Isla de Margarita (Venezuela). We identified 18 species of Actiniaria, one species of Ceriantharia, one species of Corallimorpharia, and six species of Zoanthidea; we confirmed the identification and updated the current taxonomic status of the sea anemone specimens in the collection of the MOHBR, including those from Agudo (1987) vouchered at the MOHBR. We also provided photographs for these 26 species in the lab or in natural habitats to guide identifications. These species do not represent all species of sea anemones found or reported in Venezuela, but they do represent those that are either more abundant or conspicuous. The major aim of this publication is to inventory the most common anemones in Venezuela and to provide an aid distinguishing these species to facilitate further biological and ecological research requiring species-level resolution.

Results
List of species of sea anemone found in the sampled localities We collected and examined 128 specimens belonging to 26 species, 20 genera, 14 families, and four taxonomic orders (Table 1; Appendix 1). Twenty-four of these species have been previously reported for Venezuela; the species Anthopleura pallida Duchassaing & Michelotti, 1864a, and Telmatactis vernonia (Duchassaing & Michelotti, 1864b) are reported here for the first time for the country (Table 1).

Systematics
Order ACTINIARIA Hertwig, 1882 Suborder ENTHEMONAE Rodríguez et al. 2014 Superfamily ACTINIOIDEA Rodríguez et al. 2014 Family ACTINIIDAE Rafinesque, 1815 Actinostella flosculifera (Le Sueur, 1817)    Localities sampled in the present study in bold. (*) records documented in our surveys, (?) doubtful previous records, ( †) species currently considered as invalid, (N) new records for Venezuela. Citation of previous records: (a) Specimens of the MOHBR scientific collection (see Appendix 2), (b) den Hartog (1980), (c) Knowlton & Keller (1985), (d) Riemann-Zürneck (1986), (e) Agudo (1987), (f) Bastidas & Bone (1996), (g) Bitter-Soto (1999)    Short description Pedal disc well developed, 10-30 mm in diameter. Column more or less elongate, 20-80 mm in height, slightly broader than pedal disc, smooth but with small rows of verrucae in its upper part. Pedal disc and column beige or pale orange. Above verrucae and below the margin lies a distinctive collar or marginal ruff formed by 48 rows of small frond-like fused papillae, pale brown or olive-green, and often with dark or pale flashes. Oral disc smooth, short, greyish-white and translucent, but sometimes with white flashes. Oral disc and mouth beige or whitish. Tentacles hexamerously arranged in four cycles (about 48 in specimens examined), inner cycles longer than outer ones, smooth, conical, tapering distally, white and translucent but with small circular white spots in its oral face and along their entire length. For further information on internal anatomy and cnidae see Schlenz & Belém (1992) and González-Muñoz et al. (2012).
Habitat This species inhabits commonly in seagrass meadows, with the column completely burrowed in the sand with only the marginal ruff, oral disc and tentacles protruding into the water column, often between 0.1-5.0 m.
Anemonia sargassensis Hargitt, 1908 (Fig. 2b) Anemonia sargassensis Hargitt, 1908 Anemonia antillensis: Pax, 1924 Anemonia sargassiensis [sic]: Carlgren, 1949 Short description Pedal disc well-developed, 5-20 mm in diameter, reddish or bright to dark-orange. Column cylindrical, short, smooth, reddish to dark-orange. Distal part of column with poorly marked marginal projections forming acrorhagi with holotrichs and basitrichs. Oral disc smooth, 8-25 mm in diameter, wider than column, reddish to dark-orange but with whitish or yellow radial stripes from the periphery of the mouth to the base of the tentacles; mouth reddish or brightorange. Tentacles irregularly arranged in four or five cycles (about 52-76 in specimens examined), inner cycles longer than outer ones, relatively long, smooth, slender, tapering distally, reddish to dark-orange but with the tips yellowish and sometimes with pink flashes. For further information on internal anatomy and cnidae see Fisher (1976), Carlgren & Hedgpeth (1952), and González-Muñoz et al. (2013).
Habitat This species inhabits shallow waters of coral reef lagoons, commonly under rocks or coral rubble, or even among colonies of fire-corals, between 0.5-3.0 m, but it has been also reported on floating Sargassum (Carlgren & Hedgpeth 1952).
Distribution Western Atlantic, from Bermuda and northern coast of the United States to Brazil, along the Caribbean Sea and Gulf of Mexico (González-Muñoz et al. 2012. Agudo (1987) and Carrasquel (2012) previously reported A. sargassensis in Venezuela (Table 1), inhabiting coral reefs, mangroves, and rocky shores. Short description Pedal disc well-developed, 5-10 mm in diameter, grey or whitish. Column cylindrical, stout to elongate, pale to dark-brown. Column with longitudinal rows of verrucae from margin to mid-part; distal end with small and rounded marginal projections forming acrorhagi with holotrichs, basitrichs, microbasic p-mastigophores, and spirocysts. Oral disc narrow, smooth, 3-8 mm in diameter, brownish; mouth slightly clearer than oral disc. Tentacles hexamerously arranged in three or four cycles (about 24-48 in specimens examined), inner cycles longer than outer ones, relatively short, smooth, slender, tapering distally, whitish, translucent, but with white circular spots on its oral face along their entire length, and sometimes with brown flashes. For further information on internal anatomy and cnidae see Daly & den Hartog (2004), and González-Muñoz et al. (2013).
Habitat This species lives commonly attached to coral rubble on sandy shores (González-Muñoz et al. 2013), between 0.5-1.5 m, with grains of sand adhered to the verrucae.
Distribution Western Atlantic, from Bermuda to the Caribbean Sea and Gulf of Mexico (Daly & den Hartog 2004;González-Muñoz et al. 2012; this is the first record for Venezuela found in Boca Seca, Morrocoy National Park (Table 1). Short description Pedal disc well-developed, 12-20 mm in diameter. Column cylindrical, densely covered with longitudinal rows of rounded vesicles from the margin to limbus. Pedal disc and column commonly beige or brown-reddish. Distal end of column with small and rounded marginal projections forming acrorhagi with holotrichs and basitrichs. Oral disc wide, smooth, 20-40 mm in diameter. Oral disc and mouth brown-yellowish, brown-reddish or pale brown. Tentacles hexamerously arranged in five cycles (about 96 in specimens examined), inner cycles longer than outer ones, moderately long, smooth, conical, tapering distally, white or pale-orange. For further information on internal anatomy and cnidae see Carlgren (1952), Carlgren & Hedgpeth (1952), and González-Muñoz et al. (2013).
Habitat This species lives attached to rocks or coral rubble, in shallow waters between 0.5-7.0 m.
Distribution Western Atlantic, from the northern coast of the United States to the Caribbean Sea and Gulf of Mexico (Carlgren & Hedgpeth 1952;González-Muñoz et al. 2012, and in the Caroline Islands, Micronesia (Bosc 1802). Agudo (1987)  Short description Pedal disc well-developed, 7-40 mm in diameter, brownish with bright-orange flashes. Column cylindrical, densely covered with longitudinal rows of rounded vesicles arranged in 24 alternated dark and light bands (dark bands with about five rows of vesicles, light ones with about three). Distal end of column with small and rounded marginal projections forming acrorhagi with holotrichs and basitrichs. Oral disc wide, smooth, 20-50 mm in diameter, olive-green, reddish-brown, dark-red or dark-green; mouth dark-red or yellowish. Tentacles hexamerously arranged in five cycles (about 96 in specimens examined), inner cycles longer than outer ones, moderately long, smooth, conical, tapering distally, olive-green, but with yellowish circular spots on the oral face along its entire length and sometimes with pink or dark-red flashes. For further information on internal anatomy and cnidae see González-Muñoz et al. (2012).
Habitat This species lives attached to rocks, commonly in shallow waters among rocky patches and seagrasses meadows, between 0.5-6.0 m.
Condylactis gigantea (Weinland, 1860) ( Fig. 2f- Short description Pedal disc well-developed, 20-85 mm in diameter. Column cylindrical, smooth, slightly wider than pedal disc. Pedal disc and column scarlet or bright to pale orange. Oral disc wide, smooth, 25-100 mm in diameter; oral disc and mouth yellowish to pale orange with green flashes, or pale green. Tentacles hexamerously arranged in five cycles (about 96 in specimens examined), inner cycles longer than outer ones, long, cylindrical, smooth but striated in appearance, white, greenish or pale brown in color, with tips slightly swollen and blunt, mainly bright green or pink, but sometimes white, bluish or purple. Two color morphs are commonly recognized for this species regarding the color of its tentacles tips: one with pink tips (Fig. 2f ), and another one with bright-green tips (Fig. 2g). For further information on internal anatomy and cnidae see Carlgren (1952), and González-Muñoz et al. (2012).
Habitat This species inhabit shallow waters, inside crevices formed by coral rocks and coral rubble, and is also commonly observed among seagrass meadows and mangroves zones; often between 0.5-30.0 m, but has been reported down to 40 m (Agudo 1987).
Distribution Western Atlantic, from Bermuda to Brazil, along the Caribbean Sea and Gulf of Mexico (González-Muñoz et al. 2012. Agudo (1987) previously reported C. gigantea in Venezuela (Table 1) Short description Pedal disc well-developed, 35-42 mm in diameter. Column cylindrical, elongate but sometimes funnel-shape, 60-100 mm in length and 45-60 in diameter, smooth but with longitudinal rows of vesicles in its distal part. Pedal disc and column whitish or beige, vesicles grey. Central part of oral disc narrow and smooth; mouth often white and with conchula ( Fig. 2h). Tentacles small, vesicle-like, arranged in double radial rows covering almost the entire oral disc, located on endocoelic and exocoelic spaces. Three color morphs regarding tentacles were observed in Venezuela: white tentacles with scattered brown dots (Fig. 2h), white tentacles with pale pink and purple flashes, and tentacles and oral disc completely bright-orange (Fig. 2i). For further information on internal anatomy and cnidae see Corrêa (1973) and González-Muñoz et al. (2013).
Habitat This species inhabits sandy patches, with the pedal disc strongly attached to burrowed rocks, but with the column completely buried in the sand and only the oral disc and tentacles protruding into the water column, often between 1-2 m.
Distribution Western Atlantic, in the Caribbean Sea and Gulf of Mexico to the northern coast of Brazil (Corrêa 1973;González-Muñoz et al. 2013), but also reported in Cape Verde Islands (Wirtz 2009). Agudo (1987) previously reported A. elegans in Venezuela ( Short description Pedal disc well-developed, 10-40 mm in diameter, bright pink or orange. Column cylindrical to plate-like, pinkish or orange in the proximal end and fading into whitish distally, with a flame-like staining pattern; smooth but with longitudinal rows of pink verrucae in its distal part. Oral disc wide, 40-100 mm in diameter, rough, variable in color, often brown with reddish spots, olive-green with darkbrown and white spots, grey with green spots, or white with dark-brown radial stripes from mouth to the base of the marginal tentacles; mouth olive-green, pale-brown, grey or purple. Two types of tentacles: marginal and discal. Marginal tentacles short, conical, tapering distally, numerous (about 360 in specimens examined), with lateral protuberances in the oral face, or smooth, white with longitudinal light brown stripes. Discal tentacles reduced, vesicle-like, arranged in radial rows on the oral disc. Commonly three morphotypes have been recognized for P. crucifer regarding the presence/absence (or an intermediate state) of lateral protuberances on the oral face of the marginal tentacles (the three morphotypes were observed in Venezuela). However, recent morphological and genetic analyses suggest that the three morphotypes should be considered as a single species (González-Muñoz et al. 2015). For further information on internal anatomy and cnidae see Carlgren (1952), and González-Muñoz et al. (2012).
Habitat This species lives attached to rocks and coral rubble, with the column burrowed in the sand and only the oral disc protruding into the water column, between 0.5-20.0 m.
Distribution Western Atlantic, along the Caribbean Sea and the Gulf of Mexico (Carlgren 1952;González-Muñoz et al. 2012. Agudo (1987) previously reported Phymanthus crucifer in Venezuela (Table 1) Short description Pedal disc well-developed, 35-75 mm in diameter. Column short, diabolo-shaped, smooth but with longitudinal rows of small verrucae in its distal part. Pedal disc and column beige, verrucae greenish or light-brown. Oral disc 50-250 mm in diameter, flat to domed, beige or greenish; mouth beige. Tentacles short, digitiform, blunt, arranged in single radial endocoelic rows covering almost the entire oral disc; each endocoelic row with dozens of tentacles but only one tentacle at the margin per exocoelic row (Fig. 2l). Tentacles light brown near their bases and bright-green or with yellowish shades at tips. For further information on internal anatomy and cnidae see Dunn (1981), and González-Muñoz et al. (2012.
Habitat This species lives strongly attached to rocks and coral rubble among sandy and seagrass meadows, often between 0.5-12.0 m.
Distribution Western Atlantic, from Bermuda to Brazil, along the Caribbean Sea and Gulf of Mexico (González-Muñoz et al. 2012. This species has previously been reported in Venezuela by Agudo (1987) and Carrasquel (2012)  Short description Pedal disc well-developed, 10-25 mm in diameter. Column cylindrical to elongate, divided in capitulum and scapus. Capitulum thin, smooth, light to dark-brown, translucent, with white and yellow scattered spots; scapus robust, smooth but corrugate in appearance when contracted, light brown. Mid-part of scapus with four to five rows of cinclides. Oral disc 30-45 mm in diameter, smooth, flat, brownish, translucent, with small white and yellow dots, and sometimes with triangular spots at the base of some tentacles; mouth beige. Tentacles hexamerously arranged in up to five cycles (about 48 to 96 in specimens examined), long, slender, inner cycles longer than outer ones, tapering distally, brown but translucent, with a distinct pattern of white spiral bands giving the appearance of a corkscrew. Acontia can be protruding to the water column through the cinclides or the mouth. For further information on internal anatomy and cnidae see Carlgren (1952), Carlgren & Hedgpeth (1952), González-Muñoz et al. (2012), and Rodríguez et al. (2014).
Habitat This species lives in shallow waters inside crevices of rocks and coral rubble, with only its tentacles above the substrate. It is often observed in caves and crevices at the interface between hard and sandy substratum, commonly between 0.5-15.0 m, but also observed down to 40 m.
Distribution Western Atlantic, from Bermuda to Barbados, along the Caribbean Sea and Gulf of Mexico (González-Muñoz et al. 2012. Agudo (1987) previously reported B. annulata in Venezuela (Table 1) Short description Pedal disc well-developed, 3-10 mm in diameter. Column cylindrical to elongate, only distinctly divisible into capitulum and scapus when contracted. Capitulum smooth, beige to dark brown, translucent; scapus smooth but corrugate in appearance when contracted, beige to dark brown. Column with two to three rows of cinclides in mid-scapus. Oral disc 5-15 mm in diameter, smooth, flat, brownish with white circular spots, or brownish, translucent, with small white, yellow, and bluish scattered dots; mouth brownish and sometimes surrounded by whitish spots. White acontia can be observed protruding through mouth or cinclides. Tentacles hexamerously arranged in four cycles (about 48 in specimens examined), long, smooth, slender, inner cycles longer than outer ones, tapering distally, brown, translucent, with white or bluish scattered dots along its entire length. For further information on internal anatomy and cnidae see Carlgren (1952), Carlgren & Hedgpeth (1952), González-Muñoz et al. (2012), and .
Habitat This species lives attached to rocks, coral rubble, submerged lumber, or mangrove roots, often between 0.5-22.0 m. It is commonly observed forming aggregations as a result of asexual reproduction by pedal laceration (González-Muñoz et al. 2012 Short description Pedal disc well-developed, 3-6 mm in diameter. Column cylindrical, short, smooth, light brown, often with 4-6 branched outgrowths distally (pseudotentacles), which ends are as globular-shaped vesicles. Specimens with two morphs regard of pseudotentacles are often observed: the first one with branched ends bluish with a grey or brown circle in the center (Fig. 3d), and the second one with ends bifurcated, brown with grey or white spots (Fig. 3e). Oral disc 3-5 mm in diameter, smooth, beige and translucent. Tentacles hexamerously arranged in four cycles (about 48 in number), inner cycles longer than outer ones, smooth but striated in appearance, moderately long, slender, tapering distally, greyish or brownish, translucent, with whitish or yellowish tips and scattered bluish dots along its length. For further information on internal anatomy and cnidae see Crowther (2013) and González-Muñoz et al. (2013).
Habitat This species lives attached to hard substratum, inside crevices or holes of rocks or live coral, and commonly observed with only the pseudotentacles protruding into the water column; often between 0.5-6.0 m. Habitat This species lives attached to hard substratum, inside holes or crevices among live corals and rocks, and commonly observed during day with only the pseudotentacles protruding into the water column; often between 0.5-12.0 m, but it has been reported down to 60 m depth (Ocaña et al. 2007).
Family BOLOCEROIDIDAE Carlgren, 1924b Bunodeopsis antilliensis Duerden, 1897 (Fig. 3g Short description Pedal disc well-developed, 4-13 mm in diameter, greyish or olive-green, translucent. Column short, 5-10 mm in diameter and 4-10 mm in height, divided in capitulum and scapus. Capitulum short, narrower than oral disc, smooth, whitish and translucent; scapus wider than oral disc, covered with longitudinal rows of globular vesicles, white to greenish-brown. Oral disc 4-7 mm in diameter, whitish and translucent. Tentacles irregularly arranged in three to four cycles (about 16-46 in specimens examined), inner cycles longer than outer ones, all long, smooth, tapering distally, whitish and translucent, with white dots scattered along their entire length. The tentacles are deciduous and can be completely detached by contraction of a sphincter in the base of each tentacle. For further information on internal anatomy and cnidae see Duerden (1897) and González-Muñoz et al. (2012).
Habitat This species lives attached to seagrass leaves of Syringodium filiforme or Thalassia testudinum, between 0.5-6.0 m.
Distribution Western Atlantic, from Bermuda to Curaçao, and along the Caribbean Sea and Gulf of Mexico (González-Muñoz et al. 2012. Bunodeopsis antilliensis (= B. globulifera) has been reported in Venezuela by Agudo (1987)  Short description Pedal disc well-developed, 10-30 mm in diameter, pale-orange, translucent. Column cylindrical, 12-32 mm height and 16-40 mm diameter, trumped-shaped when expanded and domeshaped when contracted, divided into capitulum and scapus. Capitulum short, smooth, pale-brown to yellowish, translucent; scapus robust, wrinkled-texture, bright to dark-orange, often with white flashes proximally, and with one or two rows of dark-red or brown cinclides proximally. Oral disc slightly wider than column, smooth, pale-brown and translucent, often with radial white stripes that sometimes forming a white ring around the peristome; mouth bright-yellow, often surrounded by a purple ring. Tentacles hexamerously arranged in up to five cycles (about 48-96 in specimens examined), inner cycles longer than outer ones, smooth, slender, moderately short, tapering distally, whitish, greyish or pale-brown, with longitudinal rows of white dots along its entire length. White or brightorange acontia can be protruding through mouth or cinclides. For further information on internal anatomy and cnidae see Gusmão (2010), and González-Muñoz et al. (2013).
Habitat This species is commonly found attached to shells of living hermit crabs, between 0.5-40.0 m.
Distribution Western Atlantic, from the northern coast of the United States to the northern coast of Brazil, and along the Caribbean Sea and Gulf of Mexico (Carlgren & Hedgpeth 1952;González-Muñoz et al. 2012Zamponi et al. 1998). Agudo (1987) and Carrasquel (2012) previously registered C. tricolor in Venezuela (Table 1), inhabiting coral reefs and mangroves.
Family ISOPHELLIIDAE Stephenson, 1935 Telmatactis cricoides (Duchassaing, 1850) ( Fig. 3i scapus robuts, wrinkled-texture, pale-orange. Oral disc slightly wider than column, 41-45 mm in diameter, smooth, whitish pale-brown, with dark-brown and reddish spots irregularly scattered; mouth dark-red. Tentacles hexamerously arranged in five cycles (about 96 in specimens examined), the inner cycles longer than outer ones, smooth, moderately short, cylindrical, with the tips widely swollen and blunt. Tentacle coloration variable, often whitish or pale-brown, with V or W shaped dark-brown or dark-red spots at its bases and with transversal white spots and reddish dots scattered along entire length. For further information on internal anatomy and cnidae see Belém & Schlenz (1989), Cairns et al. (1986), and Wirtz (1996Wirtz ( , 1997. Habitat This species lives attached to hard substrate, under rocks of coral colonies, between 1.5-20.0 m. Short description Pedal disc well-developed, 4-10 mm in diameter, pale-orange, translucent. Column cylindrical, 5-11 mm height and 4-10 mm in diameter, divided into capitulum and scapus. Capitulum short, smooth, and greyish to pale-orange; scapus robuts, wrinkled-texture, pale-orange. Oral disc slightly wider than column, 5-12 mm in diameter, smooth, dark-brown, with white spots, and greenish and purple flashes; mouth brownish. Tentacles hexamerously arranged in five cycles (about 96 in specimens examined), the inner cycles longer than outer ones, smooth, short, cone-shape, tapering distally, pale-brown with purple flashes, and reddish transversal spots along its entire length. For further information on internal anatomy see Duchassaing & Michelotti (1864a) and Cairns et al. (1986).
Habitat This species lives attached to hard substrate, under rocks or coral rubble, between 0.5-1.5 m.
Distribution Western Atlantic, reported in St. Thomas (Duchassaing & Michelotti 1864a), Panamá (Sebens & DeRiemer 1977), and Bermuda (Cairns et al. 1986 Short description Base flat and with irregular contour. Column short, stout, 20-25 mm in diameter and 28-32 mm in height, smooth, trump-shape when expanded and dome-shape when contracted. Base and column scarlet or bright-orange. Oral disc and mouth whitish and translucent. Tentacles long, about 120-160 in specimens examined, inner cycles longer than external ones, cone-shape but with capitated ends forming acrospheras. Tentacles whitish and translucent, acrospheras bright-orange. For further information on internal anatomy and cnidae see den Hartog (1980).
Palythoa grandis (Verrill, 1900) ( Fig. 4b) Protopalythoa grandis Verrill, 1900 Palythoa grandis: Pax, 1910 Short description Colonies of only few polyps that are joined by flat, stoloniferous or extensive encrusting coenenchyme; coenenchyme usually thinner than the free height of the polyps (Ryland & Lancaster 2003). Column trumped-shape, smooth, but rough in appearance due to sandy incrustations, 15-35 mm in height and 10-15 mm in diameter, ochre-yellow or brown. Tentacles in the margin, arranged in two cycles (about 60 in specimens examined), tapering distally, brown. Oral disc broad, convex or umbrella-shape when completely expanded, sometimes with the borders recurved when contracted, 10-19 mm in diameter, dark-brown or olive-greenwith pale-green radial marks, and sometimes with a whitish or beige ring surrounding the slit-like mouth. For further information on internal anatomy see Verrill (1900Verrill ( , 1905 and Varela et al. (2002).
Habitat This species lives attached to rocks and coral rubble, often between 0.5-3.0 m. Sometimes isolated polyps may occur (Verrill 1900). As other species closely related, this species has a tendency to form carpets, often intermingled with species of Zoanthus, which can grow between stolons and polyp clusters (Ryland & Lancaster 2003) (Fig. 4b).
Distribution Western Atlantic, from Bermuda to Venezuela (Acosta et al. 2005;Verrill 1900 Short description Polyps embedded in a lamellar encrusting coenenchyme, forming colonies of several polyps. Column cylindrical, 5-25 mm in height and 5-10 mm in diameter, smooth and often without sandy incrustations, white to beige. Oral disc slightly wider than column, smooth, bluish with paler patterning; mouth slit-like, sometimes with dark triangular spots at each axial extremity. Tentacles in the margin, arranged in two cycles (about 60 in number), tapering distally, bluish or greyish. For further information on internal anatomy and cnidae see Duerden (1898aDuerden ( , 1902, Verrill (1900), Varela et al. (2002), and Reimer et al. (2012).
Remarks External anatomy of these specimens, including column shape and measurements, fits well with those described for Zoanthus pulchellus, except from coloration of the oral disc which is generally described as bright-green, pale-green, or yellow, with light radiating lines and sometimes with pink, brown, or yellow patterning (Duerden 1898a(Duerden , 1902Reimer et al. 2012;Verrill 1900).
Habitat This species lives attached to rocks or coral rubble, mostly in shallow waters, but not intertidal zones, between 0.5-1.0 m.
Habitat This species lives attached to rocks or coral rubble, mostly in shallow waters, but not on intertidal zones, between 1.0-5.0 m.

Discussion
Comparison of previous records of sea anemone species in Venezuela and the findings of the present study Fourty-nine shallow water species of sea anemones have been registered along coastal and coral reefs environments in Venezuela (Table 1) (Agudo 1987;Bastidas & Bone 1996;Bitter-Soto 1999;Carrasquel 2012;den Hartog 1980;Knowlton & Keller 1985;Liñero-Arana & González 2008). In addition, Riemann-Zürneck (1986) documented the record of the actiniarian Monactis vestita (Gravier, 1918) in the Venezuela basin, between 4000-5000 m (see Table 1). Agudo (1987) recorded 27 species of the order Actiniaria, 4 of Ceriantharia, 3 of Corallimorpharia, and 13 of Zoanthidea along the central coast and islands in Venezuela (Table 1). In the present study, we found 24 of the 47 sea anemones species documented by Agudo (1987) while two actiniarian species, Anthopleura pallida and Telmatactis vernonia, are documented for the first time in Venezuela (see Table 1).
Among the 27 actiniarian species reported by Agudo (1987) that we did not find in our surveys, Actinia bermudensis (McMurrich, 1889a), Anthopleura krebsi (Duchassaing & Michelotti, 1860), Isoaulactinia stelloides (McMurrich, 1889b) (= Bunodactis stelloides), and Homostichanthus duerdeni, are species that have been commonly reported in several localities along the Caribbean Sea or the Gulf of Mexico (González-Muñoz et al. 2012. Thus, it might be expected to find these species in Venezuela under a more extensive survey. On the other hand, the known distribution of other species documented by Agudo (1987), as Bunodosoma caissarum Corrêa in Belém, 1987 andBunodosoma cangicum Belém &Preslercravo, 1973 is restricted to South America, along the Brazilian coast (Fautin, 2013;Zamponi et al. 1998), and thus, the presence of these species in Venezuela is atypical or improbable. The species Bunodosoma kuekenthali Pax, 1910 (= B. kükenthali) has only been reported for Barbados (Lewis 1960;Pax 1910) but its taxonomic status remains unknown due to the lack of recent studies about the species. The species Anthopleura cascaia Corrêa, 1964 is considered as nomen nudum according with the International Code of Zoological Nomenclature (ICZN 1999), due that it was originally described in a thesis study (Corrêa 1964), and no other description has been published to date for the species (Fautin, 2013). Bartholomea werneri Watzl, 1922 was recently synonymized with B. annulata by . Agudo (1987) also documented Scolanthus curacaoensis (Pax, 1924) (= Edwadsia horstii), which has a restricted distribution in the Laguna de Chacopata, (Liñero-Arana & González 2008), and in Curaçao (Pax 1924), and one unidentified species of genus Phymactis Milne-Edwards, 1857 (reported as Phymactis sp). Except for H. duerdeni (one specimen available previously deposited at MOHBR, see Appendix 2), it is difficult to assess or confirm the certainty of the other 10 actiniarian records because voucher specimens were not available.
From the three species of corallimorpharians reported by Agudo (1987), we only documented Corynactis caribbeorum (= Pseudocorynactis caribbeorum). This species was only found in Chichiriviche de la Costa, and has been previously reported for Venezuela by den Hartog (1980), outside the Isla Tortuga and in front to Camuri Chico coast. Discosoma carlgreni (Watzl, 1922) and Rhodactis osculifera (Le Sueur, 1817) (= Discosoma sanctithomae) are the other two species of corallimorpharians reported by Agudo (1987) in Venezuela (Table 1). These two corallimorpharian species, as well as Ricordea florida Duchassaing & Michelotti, 1860, are among the most commonly known species of corallimorpharians found in coral reefs along the Caribbean Sea and Gulf of Mexico (den Hartog 1980;Fautin & Daly 2009). Thus, despite we could not find these corallimorpharians at any of our surveys, it is possible that these species are actually present in Venezuela as they have a wide range of distribution in the Caribbean.
For the cerianthid species, Agudo (1987) documented Ceriantheopsis americanus (Agassiz in Verrill, 1864), Ceriantheomorphe brasiliensis Carlgren, 1931, Pachycerianthus curacaoensis den Hartog, 1977, and Isarachnanthus nocturnus (= Arachnanthus nocturnus) in Venezuela (Table 1). Ceriantheopsis americanus has been reported in the Northwest Atlantic, Caribbean Sea, Uruguay, and China (Fautin 2013;Molodtsova 2009). Ceriantheomorphe brasiliensis has been reported in the United States-Texas and Brazil (Carlgren 1931;Carlgren & Hedgpeth 1952;Molodtsova 2009), andP. curacaoensis in Curaçao (den Hartog 1977). In our surveys we only documented I. nocturnus in Chichiriviche de la Costa, but we have also observed another two species of tube anemones near to Isla de Cubagua that were not collected, and thus remain unidentified. More detailed taxonomic studies about cerianthid species in Venezuela are needed, in order to confirm the number of species for the region.
Out of the 13 zoanthid species previously reported in Venezuela (Agudo 1987;Bastidas & Bone 1996), we only found six (Table 1), one of them reported here as Zoanthus cf. pulchellus. Agudo (1987) also documented Zoanthus proteus Verrill, 1900 which is currently considered as synonym of Z. sociatus according to Reimer et al. (2012), and Palythoa mammillosa (Ellis & Solander, 1786), which has been suggested as synonym of P. caribaeorum (Sebens 1982). Agudo (1987) reported Palythoa variabilis (Duerden, 1898a) which is distributed throughout the Caribbean and the Atlantic coast of South America (Alves-Santos et al. 2015;Reimer et al. 2012), but also has been documented as amphi-atlantic (Reimer et al. 2010). Zoanthus solanderi Le Sueur, 1817, Parazoanthus puertoricense West, 1979, P. catenularis (Duchassaing & Michelotti, 1860, and Epizoanthus cutressi West, 1979, are also commonly included in some inventories of sea anemones of the Caribbean Sea (e.g. Ocaña et al. 2007;Reimer et al. 2012;West 1979), and thus, they are also expected to occur in Venezuela as reported by Agudo (1987). The taxonomic status of zoanthid species, mainly those of the genera Palythoa and Zoanthus, is currently considered as chaotic due to the great intraspecific variability present in several species, and thus the true number of valid species of these genera for the Caribbean is unknown (Reimer et al. 2010(Reimer et al. , 2012. However, recent studies have suggested the utility of molecular markers to distinguish among species (Alves-Santos et al. 2015;Reimer et al. 2012). Further detailed taxonomic studies of shallow water zoanthids in Venezuela are needed in order to facilitate identification to the species level.

Sea anemone specimens of the collection MOHBR
Among the 84 specimens of the collection of the MOHRB, the identification of the specimens labelled as Palythoa variabilis, Isaurus duchassaingi, and Diadumene lineata (= Haliplanella luciae), could not be corroborated due to the poor condition of the preserved specimens. However, these zoanthid species are commonly reported in Caribbean inventories of sea anemones (Acosta et al. 2005;Cairns et al. 1986;Reimer et al. 2012), and D. lineata is known as a cosmopolitan species (Fautin 2013). Thus, we might expect the presence of these two species in Venezuela, but their occurrence needs to be corroborated. The external anatomy of the specimens labelled as Zoanthus pulchellus fits well with those previously described for this species: polyps embedded in a thin lamellar coenenchyma rather than stoloniferous, with the column approximately 4-6 mm in diameter and 4 to over 30 mm in height, and 50-60 short tentacles (Reimer et al. 2012). Thus, although there is no information about the coloration pattern of these preserved polyps, we agree with the previous identification that this specimen belongs to Z. pulchellus.

Conclusions
Based on the current taxonomic status and the general distribution patterns of the 47 species documented by Agudo (1987), we suggest that at least eight of his records are currently invalid or erroneous, and only 39 sea anemone species documented by him could be found in Venezuela, of which 24 species are documented in the present study. Taking into account the previous records of D. lineata, and I. duchassaingi in the collection of the MOHBR, and that of Monactis vestita reported by Riemann (1986), plus the two new records of sea anemones documented in our surveys, this study increases to 44 the number of records of sea anemones that might be found in Venezuela.

Methods
The Caribbean coastline of Venezuela extends approximately 2,695 km, plus an important insular region constituted by 14 archipelagos composed of 300 island and keys (Conde & Carmona-Suárez 2003;Miloslavich et al. 2005). Sampling was conducted in November 2012 in four marine reserves along the Venezuelan coast: Morrocoy National Park (10°53'2.51"N, 68°12'43.66"W), Mochima National Park (10°23'44.43"N, 64°22'2.16"W), Archipiélago de Los Roques National Park (11°46'21.67"N, 66°42'8.67"W), and Laguna de La Restinga National Park (10°58'36.75"N, 64°9'4.40"W) (Fig. 1). Additionally, we include observations and collections made in Isla de Cubagua (10°50'8.40"N,64°8'51.74"W) and Chichiriviche de la Costa (10°33'12.18"N, 67°14'24.43"W) in the central coast of Venezuela (Fig. 1). Sampled habitats included sandy bottoms, rocky shore, seagrass meadows, coral rubble, coral reefs and mangrove roots. Representative specimens of each species were collected by hand, either by snorkeling or scuba diving, and using a hammer and a chisel. Specimens were photographed in situ and habitat characteristics were recorded. Collected samples were transferred to the laboratory and maintained in an aquarium to photograph their color while alive. Specimens were relaxed using menthol crystals in seawater, subsequently fixed in 10 % formalin seawater, and their diagnostic characters were examined for identification. Voucher specimens of each identified species were deposited in the collection of the MOHBR (see Appendix 1). In addition, eighty-four specimens of sea anemones previously deposited by other authors in the collection at MOHBR were examined.
We followed the taxonomic classification implemented in Fautin (2013) with modifications from Rodríguez et al. (2014). Taxa were organized in suborder and family, and listed in alphabetical order. External appearance and measurements described for each species were based in living specimens collected in Venezuela. The synonym list for each species only contains reference to the first citation of the species by a particular name.