Francisco Kelmo ¹ and Rita Vargas ²

1 Benthic Ecology Research Group, 606 Davy Building, University of Plymouth, Drake Circus, Plymouth, Devon, PL48AA. United Kingdom. E-mail: F.Kelmo@plymouth.ac.uk

2 Museo de Zoología, Escuela de Biología, Universidad de Costa Rica, 2060 San José, Costa Rica. E-mail: ritav@biologia.ucr.ac.cr

Received 16-IX-2000.       Corrected 04-VI-2001.       Accepted 30-V-2002.

Abstract: This paper is the first taxonomic account of the hydroid orders Anthoathecatae and Leptothecatae from the Caribbean and Pacific coast of Costa Rica. All specimens are currently stored at the reference collection of the Museo de Zoología, Escuela de Biología, Universidad de Costa Rica. Sixteen hydroid species are recorded: Eudendrium carneum, Pennaria disticha, Acryptolaria longitheca, Plumularia floridana, Halopteris polymorpha, Aglaophenia dubia, Aglaophenia latecarinata, Lytocarpia tridentata, Macrorhynchia phillipina, Macrorhynchia sp., Clytia gracilis, Cnidoscyphus marginatus, Thyroscyphus ramosus, Dynamena disticha, Sertularella diaphana, and Tridentata distans. An extensive synonymy has been given for each species. A simplified taxonomic key is included, and illustrations and descriptions are provided for each species.

Key words: Anthoathecatae, Lepthothecatae, hydroids, taxonomy, Costa Rica.

]]> The hydroid fauna of Costa Rica is little studied and therefore poorly known. Studies on species from the Pacific coast of the country include those of Fraser (1938, 1948) and Hertlein (1963), but no study exists recording the hydroids from the Caribbean coast. The most recent report on the biodiversity of cnidarians from Costa Rica (Cortés 1997) is a list of species previously reported.

The scarcity of reliable taxonomic characters in various hydroid taxa, together with a scarcity of data on the biology of these animals, including the life cycles (Calder 1988, Kelmo and Santa-Isabel 1998), hampers hydrozoan taxonomy. This discourages research into the group, and therefore impedes the advancement of knowledge.

The purpose of this report is to provide the first taxonomic account of the Anthothecate and Leptohecatae hydroids currently stored at the reference collection of the Museo de Zoología, Escuela de Biología, Universidad de Costa Rica (UCR), San Pedro, San José, Costa Rica. Here we present a detailed synonymy, a simplified identification key, descriptions and illustrations for each studied species covering both Pacific and Atlantic coasts.

Colonies were examined using stereoscopic and light microscopy, and the taxonomic identifications were based on the specific literature noted in the text of this paper. All descriptions provided herein are from the examined material, with accompanying illustrations constructed using brightfield microscopy.

The listed synonymy essentially follows that of Calder (1988, 1991, 1997a), Marques (1992), Migotto (1993), and Kelmo and Santa-Isabel (1998), and virtually all of these lists have been verified by examination of the original references. In most of cases, only one significant record has been cited to document occurrences worldwide. Hydroid classification has been extensively modified by Petersen (1979, 1990), Werner (1984), Bouillon (1985), and Calder (1988, 1991, 1997a). However, in the interests of simplicity, the classification adopted here is that followed by Calder (1988, 1991, 1997a, 1997b).

Nematocysts were examined by compressing pieces of tissue, or entire individuals, between slide and coverslip (Calder 1991). The specimens were treated with 10% solution of sodium hypoclorite for 5-10 seconds, and rinsed twice in distilled water prior to slide preparation (Kelmo and Santa-Isabel 1998). Nematocyst categories were identified according to the classification of Weill (1934). At least 10 nematocysts of each type were measured from each species in order to determine size ranges.

A total of 243 colonies were analyzed, consisting of nine families and sixteen species of anthoathecate and leptothecate hydroids from both the Pacific and Caribbean coast of Costa Rica (Table 1).
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Order Antoathecatae Cornelius, 1992
Super Family Eudendrioidea L.Agassiz, 1862
Family Eudendriidae L. Agassiz, 1862

Eudendrium carneum Clarke, 1882
Fig. 1)

Eudendrium ramosum McCrady, 1859; A.Agassiz, 1865; Congdon, 1906; Fraser, 1912; Bennitt, 1922 [ Not Eudendrium ramosum (Linnaeus, 1758)]. Eudendrium carneum Clarke, 1882; Calder, 1988; Marques, 1992; Kelmo and Santa-Isabel, 1998. Eudendrium cunninghami Kirkpatrick, 1910.

Material examined: eleven colonies, with male and female gonophores, Puerto Limón, Limón, Caribbean coast, 9°59’40’’ N, 83°01’30’’ W, 12 November 1993 (UCR-845).

Description: branched and bushy colonies, up to 12 cm, arising from creeping hydrorhizal stolons. Hydrocaulus upright, polysiphonic, more or less alternately branched; primary branches also polysiphonic and irregularly branched; secondary branches polysiphonic basally and irregularly or alternately branched. Perisarc thick, annulated or wrinkled at bases of branches and hydranth pedicels, with occasional annulations elsewhere. Hydranths urn-shaped, about 0.82 mm long from proximal end to base of hypostome, 0.63 mm wide, with a shallow perisarc groove; hypostome very large, flared to knob-shaped. Tentacles solid, filiform, 28-32 in number, disposed in one whorl. Gonophores fixed sporosacs, developing on hydranth distal to perisarc groove. Female gonophores on reduced hydranths with partially atrophied tentacles; spadix bifid, curving over egg. Male gonophores with up to five chambers each, borne on atrophied hydranths. Cnidome comprising heterotrichous anisorhizas 20.2-22.8 µm x 9.3-11.4 µm, and by heterotrichous microbasic euryteles 8.2-9.4 µm x 3.5-4.0 µm.

Known range: Bermuda (Congdon 1906, 1907; Bennitt 1922; Summers 1972; Calder 1986, 1988); western Atlantic (Fraser 1944); eastern Atlantic (Kirkpatrick 1910); Indian Ocean (Millard 1975); eastern Pacific, Gulf of California, north of Clarion Island (Fraser 1948); Brazil (Marques 1992, Kelmo and Santa-Isabel 1998).

Family Halocordylidae Stechow, 1921

Pennaria disticha (Goldfuss, 1820)
(Fig. 2)

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Pennaria disticha Goldfuss, 1820; Plumularia pennaria Blainville, 1830; Pennaria cavolini Ehrenberg, 1834; Chiaje, 1841 in  Calder, 1988. Anisocalyx pinnarium Costa, 1842. Globiceps tiarella Ayres, 1854. Eucoryne elegans Leidy, 1855. Pennaria tiarella McCrady, 1859. Pennaria gibosa L.Agassiz, 1860. Halocordyle tiarella Allman, 1872. Pennaria symmetrica Clarke, 1879. Globiceps globator Haeckel, 1879. Pennaria inornata Brooks, 1883. Pennaria australis Bale, 1884. Pennaria adamsia van Lendenfeld, 1885b. Pennaria pennaria Marktanner-Turneretscher, 1890. Halocordyle australis Bale, 1894. Halocordyle cooperi Warren, 1906. Pennaria pacifica Clarke, 1907. Pennaria australis var. cooperi Warren, 1908. Pennaria australis Warren, 1908. Pennaria disticha var. australis Ritchie, 1910. Pennaria wilsoni Bale, 1913. Halocordyle disticha Stechow, 1923a. Halocordyle wilsoni Stechow 1923a. Corydendrium splendidum Boone, 1938. Halocordyle disticha var. australis Vervoort, 1941. Halocordyle fragilis Vannuci, 1951. Halocordyle pennaria var. australis Mammem,1963. Pennaria americana García-Corrales and Aguirre, 1985 [Nomem Nudum according to Calder (1988)]. Penaría (Halocordyle) tiarella García-Corrales and Aguirre, 1985. Pennaria europea García-Corrales and Aguirre, 1985 [Nomem Nudum according to Calder (1988)]; Pennaria (Halocordyle) disticha  García-Corrales and Aguirre, 1985. Pennaria symetrica García-Corrales and Aguirre, 1985; Halocordyle disticha Calder, 1988; Kelmo and Santa-Isabel, 1998; Pennaria disticha Silveira and Migotto, 1992.

Material examined: four colonies without gonophores or medusae buds, artificial reef, Bahía Huevos, Guanacaste, Pacific coast, 10°38’40’’ N, 85°41’30’’ W, 28 March 1984 (UCR-895). Eight colonies without gonophores or medusae buds, Manzanillo, Limón, 9º 38’10’’ N, 82º 38’10’’ W, Caribbean coast, 19 September 1992 (UCR-690).

Description: erect colonies, arising from a creeping and branched hydrorhiza; growth monopodial with terminal hydranths. Hydrocaulus monosiphonic, reaching about 0.4 mm wide, zigzag to nearly straight, annulated basally, divided at more or less regular intervals by one or more well-developed annulations; internodes 0.62-4.0 mm long, each typically supporting a branch distally. Perisarc thick, black or dark-brown. Branches up to 28 mm long, annulated basally, given off alternately from opposite sides of hydrocaulus, curved outwards, divided into internodes. Internodes 1.6-4.2 mm long, marked by distinct to rather faint annulations proximally and distally; each internode giving rise to a ramulus from both its upper surface and its distal end. Ramuli unbranched, annulated basally or throughout entire length, each terminating in a hydranth. Hydranths clavate to pear-shaped, up to 1.8 mm long, 0.35 mm wide; with a whorl of about 10 to 18 long, filiform or faintly knobbed tentacles aborally; a varied number of short, capitate tentacles in one or more regular or irregular verticils medially; and a whorl of about five capitate tentacles orally. Hypostome dome shaped. Cnidome comprising desmonemes 4.4-5.3 µm x 3.2 -3.8 µm; basitrichous haplonemes 5.6-9.5 µm x 2.5- 3.7 µm; heterotrichous microbasic euryteles 10.6-13.5 µm x 6.0-6.9 µm, and four sizes of stenoteles: (i) very small, 5.5-6.5 µm x 4.2- 4.6 µm; (ii) small, 7.3-7.8 µm x 5.4-5.8 µm; (iii) medium, 14.1-17.5 µm x 10.0-12.2 µm, and (iv) large, 28.0-40.0 µm x 16.4-20.5 µm.

Known range: Bermuda (Verrill 1900, Congdon 1907, Bennitt 1922, Weill 1937, Cowden 1964, 1965a, b; Summers and Haynes 1969; Summers 1970, 1972; Lesh-Laurie 1976, Calder 1986, 1988); weastern Atlantic (Fraser 1944); eastern Atlantic (Brinckmann-Voss 1970); Indian Ocean (Millard 1975); western Pacific (Yamada 1959); eastern Pacific, from San Pedro, California; San Francisco Bay, east of Panamá City; off La Libertad, Ecuador (Fraser 1948); Brazil (Migotto 1993, Kelmo and Santa-Isabel 1998).

Order Leptothecatae Cornelius, 1992
Superfamily Lafoeoidea A.Agassiz, 1865
Family Lafoeidae A.Agassiz, 1865
Subfamily Lafoeinae A. Agassiz, 1862

Cryptolaria longitheca Allman, 1877; Clarke, 1879; Fewkes, 1881; Bedot, 1912; Bedot, 1916; Bedot, 1918; Bedot, 1925; Cryptolaria crassicaulis Driesch, 1889; Ritchie, 1911; Leloup, 1932; Stranks, 1993. Lafoea (Cryptolaria) longitheca Bonnevie, 1899; Billard, 1906. Cryptoaria crassicaulis Pictet and Bedot, 1900; Browne, 1907; Jaderholm, 1907; Stechow, 1913a, b; Jaderholm, 1919. Cryptolaria crassicaulis var. dimorpha Ritchie, 1911; Jarvis, 1922. Oswaldaria crassicaulis Stechow, 1921; Stechow, 1923b; Leloup, 1940. Oswaldaria longitheca Stechow, 1923b; Acryptolaria longitheca Fraser, 1943, 1944; Deevey, 1954; Vervoort, 1968; Vervoort, 1972; Cairns et al., 1991; Calder, 1996; Calder and Vervoort, 1998. Acryptolaria crassicaulis Yamada, 1959; Rees and White, 1966; Millard, 1967; Leloup, 1974; Gravier-Bonnet, 1979; Vervoort, 1985; Ramil and Vervoort, 1992; Altuna Prados, 1994; Blanco, 1994a,b; Altuna Prados, 1995; Bouillon et al., 1995; Mendel and López Gonzalez, 1996. Acryptolaria crassicaulis var dimorpha Smaldon et al., 1976; Gravier-Bonnet, 1979.

Material examined: one colony without gonothecae, northwestern Isla del Caño, Puntarenas, Pacific coast, 8°46’08’’ N, 84°17’04’’ W, 990 m, 25 January 1994 (UCR-899).

Description: colony erect and flabellate, 1.2 cm high, arising from a creeping hydrorhiza. Hydrocaulus geniculate, polysiphonic; nodes indistinct; branches in one plane, resembling hydrocaulus, and not secondarily branched. Perisarc fairly thick. Hydrothecae very long, slender, total length excluding renovations is about 1793 µm, occurring on hydrocaulus and branches, given off alternately in one plane, each adnate to axial tube for most of its length, curving outwards and slightly frontwards, becoming free distally; narrowest at the base, gradually widening distally, free part nearly cilyndrical. Hydrothecal walls smooth; perisarc of abcauline wall thicker basally than distally. Margin entire, slightly flaring, usually with 3 or 4 renovations; orifice round, diameter about 296 µm. Operculum, diaphragm, desmocytes and nematothecae absent. Cnidome represented by very small microbasic mastigophores, 3.5–4.2 µm x 1.1–1.4 µm.

Known range: western Atlantic, mid-Atlantic Ridge, Australia (Allman 1877, 1888 as Cryptolaria crassicaulis); Clarke 1879, Vervoort 1972, Calder 1996, Calder 1997b, Calder and Vervoort 1998.

Family Thyroscyphidae Stechow, 1920

Thyroscyphus marginatus (Allman, 1877) ]]> (Fig. 4)

Obelia marginata Allman, 1877; Campanularia insignis Allman, 1888 [not Campanularia insignis Fewkes, 1881 = Campanularia macroscypha Allman, 1877]; Obelia (Lytoscyphus) marginata Ritchie, 1909; Lytoscyphus insignis Ritchie, 1909; Lytocysphus marginata Ritchie, 1909; Lytoscyphus marginatus Billard, 1910; Campanularia marginata Nutting, 1915; Leptocysphus marginatus Jaderholm, 1920; Cnidoscyphus marginatus Splettsosser, 1929; Thyroscyphus marginatus Calder, 1983, 1990.

Material examined: twelve colonies without gonophores, Manzanillo, Limón, Caribbean coast, 9º 37’55’’ N, 82 o 39’10’’ W, 19 September 1992 (UCR-688). Nineteen colonies without gonophores, Punta Piuta, Limón, Caribbean coast, 10°00’35’’ N, 83°02’15’’ W, subtidal zone, 1-2 m, 13 November 1983 (UCR-847).

Description: colonies erect, up to 20 cm high, arising from a creeping hydrorhiza. Hydrocaulus monosiphonic, up to 1 mm wide, straight basally, becoming slender and distinctively geniculate distally, divided into internodes at regular intervals by slightly, distinct or indistinct, oblique nodes. Perisarc thick basally and thinner distally. Internodes with prominent apophysis distally; apophyses alternating from side to side, each supporting a hydrotheca. Hydrocladia typically alternate, unbranched or alternately branched; hydrocladia divided by oblique nodes into internodes resembling, but more slender than, internodes of hydrocaulus, and with basal internode somewhat longer than others. Hydrothecae pedicellate, cone-shaped, 1018-1090 µm long from apophysis to margin; adcauline wall convex, abcauline wall nearly straight; diaphragm a ring of perisarc, often somewhat thicker on adcauline than abcauline wall; margin entire, occasionally renovated, with a ring like thickening; hydrothecal operture round, 486- 680 µm wide; operculum shed. Hydranth attached to hydrothecal wall by an annular fold distal to diaphragm. Abcauline diverticulum not observed. Gonophores not observed. Cnidome comprising microbasic mastigophores of three distinct sizes: (i) very large, 31- 34 x 12.4-16 µm; (ii) large, 27-29 x 9.8- 10.6 µm; and (iii) small, 3.5-3.9 x 1.1-1.8 µm.

Known range: Bermudas (Allman 1888, Congdon 1907, Ritchie 1909, as Campanularia insignis), (Jaderholm 1920 as Leptoscyphus marginatus), (Bennit 1922, Fraser 1944 as Campanularia marginata), Calder 1986, 1990; western Atlantic Calder 1983; eastern Atlantic Vervoort 1959.

Thyroscyphus ramosus (Allman, 1877) ]]> (Fig. 5)

Thyroscyphus ramosus (Allman, 1877); (Splettstösser, 1929); (Vervoort, 1959); (van Gemerden-Hoovegeen, 1965).

Material examined: thirteen colonies, without gonophores, Punta Moín, Limón, Caribbean coast, 10°00’05’’ N, 83°05’W, 9 m, 12 November 1983 (UCR-842). Nine colonies without gonophores, north-east coast of Isla Uvita, Limón, Caribbean coast, 9°59’40’’ N, 83°00’50’’ W, 11 November 1993 (UCR-844). Eight colonies without gonophores, Refugio Nacional de Vida Silvestre Gandoca-Manzanillo, Limón, Caribbean coast, 9°37’55’’ N, 82°39’10’’ W. 20 m, 4 August 1995 (UCR-849).

Description: colonies erect, up to 25 cm high, arising from a creeping hydrorhiza. Hydrocaulus monosiphonic, up to 1.2 mm wide, divided into internodes, by oblique nodes. Internodes of 1180-2968 µm long and 250-772 µm in diameter, inclined alternately in opposite directions. Nodes vary from 300 to 1043 µm in diameter. Irregularly branched. Perisarc of moderate thickness. Internodes with an apophysis distally placed, supporting a pedicellate hydrotheca. Pedicel short, with 2 to 5 annulations. Hydrothecae alternately placed; abcaulinar wall slightly convex. Apophysis margin of hydrothecae about 1198 to 1802 µm long, and diaphragm margin vary from 800 to 1343 µm long. Diaphragm oblique, virtually thicker on adcauline wall. Margin with four teeth of moderate prominence. Operculum with four conspicuous valves. Gonothecae not observed. Cnidome comprising microbasic mastigophores of two distinct sizes: (i) very large, 22.6–25.2 x 10.3–11.8 µm and (ii) small, 8.1–9.6 x 3.0–4.1 µm.

Known range: Caribbean Sea (Van Germerden-Hoogeveen 1965); western Atlantic (Vervoort 1959); southern Atlantic (Allman 1888, Migotto 1993).

Dynamena disticha (Bosc, 1802) ]]> (Fig. 6)

Sertolara pumila Cavolini, 1785; Sertularia disticha Bosc, 1802; Dynamena disticha Lamouroux, 1812; Dynamena cornicina  McCrady, 1859; Sertularia cornicina Verrill, 1874; Sertularia exigua Allman, 1877; Sertularia complexa Clarke, 1879; Sertularia mayeri Nutting, 1904; Sertularia pourtalesi Nutting, 1904; Sertularia densa Stechow, 1919; Dynamena densa Stechow, 1920; Tridentata complexa Stechow, 1922; Tridentata cornicina Stechow, 1922; Tridentata disticha Stechow, 1922; Tridentata exigua  Stechow, 1922; Tridentata pourtalesi Stechow, 1922; Dynamena mayeri Stechow, 1925; Dynamena disticha form dense Picard, 1951a; Dynamena disticha form souriei Picard, 1951b; Dynamena cavolini Riedl, 1959 [Justified emendation]; Dynamena disticha Calder, 1990; Migotto, 1993; Souza, 1997.

Examined material: thirty six colonies with gonothecae, northeast part of Isla Uvita, Limón, Caribbean coast, 9°59’40’’ N, 83°00’50’’ W, 11 November 1993 (UCR-844).

Description: colonies erect, up to 1.2 cm high, arising from a creeping hydrorhiza. Hydrocaulus monosiphonic, straight, unbranched, divided into internodes by usually distinct oblique nodes, or by distinct hinge-joints; basal part of hydrocaulus with one or more athecate internodes 1.6 to 2.1mm long, each with one pair of frontally placed hydrothecae. Hydrothecal pairs contiguous for a varying distance in front of internode; distal half parallel with axis of internode; distal half curving outwards with orifice facing nearly perpendicular to internodal axis. Abcauline wall concave, 267-474 µm long; contiguous portion of adcauline wall straight, 129-360 µm long; free part of adcauline wall convex, 202- 350 µm long. Hydrothecal base about 112 µm wide, with triangular perisarcal projections extending proximally; hydrothecal orifice oval, 98-136 µm wide from adcauline to abcauline wall; margin with two large pointed lateral teeth and a smaller marginal adcauline tooth. Operculum consisting of a large abcauline valve and a smaller tent-shaped adcauline valve. Hydrothecae usually with internal teeth. Gonophores fixed sporosacs. Gonothecae oval-shaped laterally, round-shaped in cross section, usually with six, rounded, tranverse ridges. Gonotechae about 1mm long from base to orifice, 0.62 mm wide, placed by short pedicels into hydrorhiza. Submarginal teeth present. Cnidome comprising microbasic mastigophores of two distinct sizes: (i) large, 21.0–26.2 x 8.9–10.02 µm, and (ii) small, 6.8–8.1 x 2.0–2.8 µm.

Known range: Bermudas (Bennitt 1922 as Sertularia cornicina; Calder 1986, 1990), western Atlantic (García et al 1980), Indian Ocean (Mammen 1965), western Pacific (Park and Rho 1986 as Dynamena cornicina), eastern Pacific (Fraser 1937), Brazil (Migotto 1993, Souza 1997).

 Sertularella diaphana (Allman, 1885)
(Fig. 7)

Examined material: five colonies without gonothecae, Refugio Nacional de Vida Silvestre Gandoca-Manzanillo, northern Punta Mona, Limón, Caribbean coast, 9°37’45’’ N, 82°37’07’’ W, 9 m, 4 August 1995 (UCR-846).

Description: colonies erect, up to 16 mm high, arising from a creeping hydrorhiza. Hydrocaulus usually polysiphonic, rarely monosiphonic, slightly geniculate, divided into internodes at regular intervals by oblique nodes sloping alternately left and right. Perisarc thick. Internodes with pairs of alternate hydrothecae; apophyses short, given off alternately from opposite sides of hydrocaulus; each apophysis separated from hydrocladium by a distinct transverse node. Hydrocladia straight to slightly geniculate, up to 9.3 mm long, divided into internodes of varied length by oblique nodes. Hydrothecae alternate, on opposite sides of hydrocladia, almost entirely adnate, each with axis oblique to that of internode; adcauline wall convex, 310-349 µm long; abcauline wall slightly concave, occasionally with a faint bulge basally; basal diameter 105- 151 µm. Hydrothecal orifice nearly circular, 201-220 µm wide from adcauline to abcauline tooth; margin with four small teeth. Operculum with four valves. Hydranths with an abcauline diverticulum. Gonophores not observed. Cnidome comprising microbasic mastigophores of two distinct categories: (i) large, 23.2–2 7.4 x 9.3–11.2 µm, and (ii) small, 6.6–8.4 x 2.6–3.1 µm.

Known range: Bermudas (Congdon 1907 as Sertularella speciosa, Calder 1990), western Atlantic (Vervoort 1968 as Sertularella speciosa), Indian Ocean (Millard 1975), western Pacific (Billard 1925a), Brazil (Souza 1997).

Tridentata distans (Lamouroux, 1816)
(Fig. 8)

Dynamena distans Lamouroux, 1816; Sertularia lamourousii Deshayes and Milne-Edwards, 1836; Sertularia gracilis Hassall, 1848; Sertularia (Dynamena) distans Busk, 1852; Dynamena gracilis Marktanner-Turneretscher, 1890; Sertularia stookeyi  Nutting, 1904; Stechow, 1922. Sertularia distans Billard, 1906; Sertularia distans gracilis Billard, 1912; Tridentata gracilis  Stechow, 1920; Tridentata (Sertularia) gracilis Stechow, 1920; Tridentata heterodonta Stechow, 1922. Tridentata distans Hiroito, 1969; Calder, 1990; Souza, 1997. Sertularia distans Migotto, 1993.

Examined material: twenty five colonies with gonophores, south of lighthouse, Isla del Caño, Puntarenas, Pacific coast, 8°43’ N, 83°52’ W, 19 April 1984 (UCR-892).

Description: colonies erect, up to 12 mm high, arising from a creeping hydrorhiza having internal septa. Perisarc thick. Hydrocaulus monosiphonic, straight, usually with a long stolon distally, unbranched but occasionally with one to four branches; each branch arising from renovation of an old hydrotheca, divided into internodes. Basal part of hydrocaulus with one to six short athecatae internodes separated by oblique hinge-joints. Upper thecate internodes 286-432 µm long, separated by distinct or indistinct oblique to occasionally almost transverse nodes; each internode with a distal pair of frontally placed hydrothecae. Sometimes two athecate internodes separated by a short athecate internode marked by an oblique node at proximal end and oblique hinge-joint at distal end. Hydrothecal pairs opposite; pairs virtually contiguous; rarely separated frontally. Hydrothecae horn-shaped, facing outwards to obliquely upwards, each with an axis oblique to that of internode; abcauline wall 180-232 mm long, concave medially, nearly straight distally, often with a basal bulge proximally; contiguous part of adcauline wall straight, 5- 160 µm long, becoming nearly straight distally; diameter at base 55-118 µm. Perisarc of hydrothecae thickened near margin; adcauline and abcauline walls each either lacking intrathecal teeth or having one or two such teeth just below margin; hydrothecal base near adcauline wall with a small triangular perisarcal projection extending proximally into internode, and usually with a perisarcal projection protruding from base near hydropore into hydrothecal cavity. Hydrothecal orifice irregularly oval, 60-90 µm wide from adcauline to abcauline wall; margin with two lateral teeth and a smaller median adcauline tooth. Operculum consisting of a large abcauline valve. Hydranth with an abcauline diverticulum. Gonophores fixed sporosacs. Gonothecae arising from short pedicels on hydrocaulus at bases of hydrothecae, normally numbering two per hydrocaulus, elongated and oval in lateral view, irregularly oval to nearly round in cross section, with about four faint, rounded ridges. Gonothecae vary from 923 to 1024 µm long from base to orifice; diameter vary from 398 to 414 µm. Orifice nearly round, about 220 µm in diameter. Operculum and submarginal teeth present. Cnidome comprising microbasic mastigophores of two distinct sizes: (i) large, 14.0–15.2 x 3.6–4.5 µm and (ii) small, 5.2–5.9 x 2.0–2.7 µm.

Plumularia floridana Nutting, 1900
(Fig. 9)

Plumularia floridana Nutting, 1900; Plumularia alicia Torrey, 1902; Plumularia florida Wallace, 1909; Plumularia alicia var. minuta Billard, 1927; Plumularia sinuosa Fraser, 1938; Plumularia floridina Pennycuik, 1959; Plumularia sp. Pennycuik, 1959; ?Plumularia indica Mammen, 1965; Plumularia pennycuikae Millard and Bouillon, 1973; Plumularia pennycuikai Rho and Park, 1986.

Description: colony 12 mm high, with a creeping hydrorhiza. Hydrocaulus monosiphonic, branched, more or less straight basally, geniculate elsewhere, divided at regular intervals beyond basal region internodes by distinct transverse nodes; internodes 290-600 µm long, 40-90 µm wide at nodes, with conspicuous septa adjacent to nodes. Each internode with a distal apophysis and typically with two or more nematothecae, one or two axillary and one along internode on side opposite apophysis; cauline nemathothecae bithalamic, movable, cone shaped to scoop shaped. Aphophysis short, alternate, each bearing a mamelon on dorsal side near node and supporting a hydrocladium. One or two branches, irregularly arranged, inserted at an angle of about 45º to the hydrocaulus, each replacing a hydrocladium and arising from an apophysis. Hydrocladia unbranched, up to 3.4 mm long, directed outward at an angle of about 60º from axis of hydrocaulus, divided into alternating athecate and thecate internodes each with two variably developed internal septa, one at each end; most proximal internode 90-330 µm long, with a straight node proximally and an oblique node distally, lacking hydrothecae and nematotheca; thecate internodes 238-409 µm long, each with an oblique node proximally, a straight node distally, a medium inferior nematotheca; athecate internodes beyond the most proximal one 192-590 µm long, with a straight node proximally, an oblique node distally, and one intermediate nematotheca. Nematotheca of hydrocladia bithalamic, movable, cone-shaped to scoop-shaped; median nematotheca 43-55 µm long, typically ending proximal to or just reaching the base of hydrotheca; lateral nematotheca 55-65 µm long, not reaching to margin of hydrotheca. Hydrotheca 148- 178 µm deep, cup-shaped, distal third or more of adcauline wall free from internode, axis oblique to that of internode; abcauline wall slightly convex basally, concave distally; margin entire, perpendicular or nearly so to axis of hydrothecae, aperture oval, in outline, diameter 115-144 µm; without intratecal septum. Cnidome comprising microbasic mastigophores 4.4-5.1 x 1.9-2.0 µm, pseudostenoteles 8.4-10,2 x 3.9-4.4 µm; and isorhizas 7.8-8.9 x 1.9-2.5 µm.

Known range: Bermuda (Calder 1983, 1993, 1997a); western Atlantic (Calder 1983); eastern Atlantic (Billard 1927); Indian Ocean (Millard 1975); western Pacific (Hirohito 1974); eastern Pacific, Point Vicente, California (Fraser 1948).

Family Halopterididae Millard, 1962

Halopteris polymorpha (Billard, 1913)
(Fig. 10)

Plumularia polymorpha Billard, 1913; Halopteris polymorpha Vervoort, 1966; Millard and Bouillon, 1973; Millard, 1975, Hirohito, 1983; Migotto, 1993; Schuchert, 1997.

Description: colonies up to 32 mm, arising from a creeping hydrorhiza. Hydrocaulus monosiphonic, unbranched, base not segmented, lacking hydrocladia, with a variable number of nematothecae, separated by an oblique articulation; hydrocaulus divided at regular intervals by distinct transverse nodes; each caulinar internode with one hydrothecae, one apophysis laterally disposed and five to seven nematothecae (one mesial, two lateral, and two to four superior). Hydrocladia alternate, composed of one or two basal joints, followed by two athecate internodes with one nematotheca and alternately athecate. Athecate internode usually continued with the thecate one, with an inconspicuous node, or lacking a node. Hydrothecal internode with one hydrotheca and three or five nematothecae (one mesial and two laterally disposed), and with one or two reduced sessile nematothecae between hydrothecae and hydrocaulus. Hydrothecal margin 180-200 µm, and 60-128 µm deep; lateral nematothecae 76-128 µm long, placed on the apophysis. Cnidome comprising pseudostenoteles 18-22 x 7.6-9.2 mµm and microbasic mastigophores 6.0-7.1 x 1.8-2.1 µm.

Known range: Indian Ocean, Pacific (Billard 1913, Vervoort 1966, Millard and Bouillon 1973, Millard 1975, Hirohito 1983); Brazil (Migotto 1993).

Aglaophaenia dubia Nutting, 1900
(Fig. 11)

Aglaophenia gracilis Allman, 1877 [Not Aglaophenia gracilis Lamouroux, 1816]; Aglaophenia dubia Nutting, 1900; Aglaophenia flowersi Nutting, 1900; Aglaophenia lophocarpa Bennit, 1922 [Not Aglaophenia lophocarpa Allman, 1877]; Aglaophenia allmani Leloup, 1935 [Not Macrorhynchia allmani (Nutting, 1900)]; Aglaophenia elongata Leloup, 1937; Deevey, 1954; van Gemerden-Hoogeven, 1965; Vervoort, 1968; Defenbaugh, 1974 [Not A.elongata Meneghini, 1845]; ?Aglaophenia dubia Fraser, 1938; Aglaophenia acacia Svoboda, 1979; Svoboda and Cornelius, 1991 [Not A.acacia Allman, 1883]; Aglaophenia apocarpa Calder, 1993 [Not Aglaophenia apocarpa Allman, 1877]; Aglaophenia sp. Calder, 1993; Aglaophenia dubia Calder, 1997a.

Material examined: eight colonies, Punta Pitahaya, Guanacaste, Pacific coast, 10º03’55’’ N, 85º45’57’’ W, 15 June 1991 (UCR-577).

Known range: Bermuda (Bennitt 1922, Calder, 1993, 1997a); western Atlantic (Rees and Thursfield 1965); eastern Pacific (Fraser, 1938).

Aglaophenia latecarinata Allman, 1877
(Fig.12)

Sertularia pluma Bosc, 1802 [Not Aglaophenia pluma (Linnaeus, 1758)]. ?Aglaophenia pelagica Lamouroux, 1816 [Nomem dubium]. ?Plumularia simplex d’Orbigny, 1846 [Nomem dubium].

?Aglaophenia simplex Kirchenpauer, 1872 [Not Aglaophenia perforata Kirchenpauer, 1876 (Nomem nudum)] [=Macrohynchia phillipina Kirchenpauer, 1872]. Aglaophenia perpusila Allman, 1877; Aglaophenia latecarinata Allman, 1877 [Incorrect original spelling]; Aglaophenia latecarinata Tizard et al. 1885 [Justified emendation]; Migotto, 1993; Calder, 1997a. Aglaophenia minuta  Fewkes, 1881; Agalophenia perforata Allman, 1885; Aglaophenia mammilata Nutting, 1900; Aglaophenia minima Nutting, 1900; Anisocalyx (Aglaophenia) pelagica Bedot, 1905; Aglaophenia latecarinata madagascariensis Billard, 1907; glaophenoides mamillata Morris and Mogelberg, 1973; Not Aglaophenia latecarinata García et al, 1978; Aglaophenoides mammillatus Cairns et al, 1991 [Justified emendation].

Examined material: six colonies, with corbulae, Ocotal, Guanacaste, Pacific coast, 10°32’54’’ N, 85°43’30’’ W, 23 March 1984 (UCR-896).

Description: colonies up to 28 mm high, with a creeping, wrinkled hydrorhiza. Hydrocaulus monosiphonic, slender, unbranched, 0.1-0.2 mm wide basally, slightly convex frontally; perisarc thick basally, thinning progressively towards distal end. Basis of hydrocaulus with several irregular annulations just above the juncture with hydrorhiza. One or two large cauline nematothecae present; separated from hydrocladium-bearing part above by one short internode having a single nematothecae and oblique hinge-joints proximally and distally. Distal part of hydrocaulus divided into internodes by distinct to indistinct transverse nodes; each internode of hydrocaulus typically with a large, cone-shaped, inferior nematothecae, a fronto-laterally situated hydrocladial apophysis, and a pair of small, sac-shaped, axillary nematothecae; apophysis short, given off alternately from each side of hydrocaulus, adjacent pair on a given side about 0.60-1.0 mm apart; each apophysis with a cone shaped mamelon basally and supporting a hydrocladium distally; mamelom prominent, with a round to oval orifice. Hydrocladia reaching about 4 mm long, unbranched, directed outward at an angle of about 60º or more from the axis of hydrocaulus; frontal sides convex; top sides nearly straight; each hydrocladium arising directly from an apophysis and divided into short internodes by nearly transverse nodes. Hydrocladial internodes 215- 420 µm long, 50-60 µm wide at nodes, each one with one frontally placed hydrotheca, one median inferior nematotheca, and one pair of lateral nematothecae; internodal septa typically two, with one at base of intrathecal septum and another at the basis of lateral nematothecae, both well developed and reaching across internode, occasionally with an additional weakly developed septum between these two.

Hydrothecae close together, 241-318 µm deep from tip of median abcauline cusps to base, cone shaped, axis oblique to that of internode, adcauline wall convex, occasionally with a slight concavity at the proximal end, notched at the intrathecal septum, adnate to hydrocladial internode except for small free part at the distalmost end; abcauline wall convex basally, nearly straight to slightly concave from juncture of intrathecal septum to margin; prominent septum, slightly oblique, extending from adcauline to abcauline wall of hydrothecae; appearing in lateral view to divide cavity into two sections; abcauline wall with a carina of varying thickness, from thin and almost imperceptible to thick and keel-shaped, extending forward to mediam abcauline cusp, distal end of carina prow-shaped. Perisarc elsewhere of moderate thickness. Hydrothecal aperture 140- 168 µm in diameter from adcauline wall to base of median abcauline cusp, irregularly oval in outline, plane of orifice nearly perpendicular to long axis of the hydrothecae; margin with nine cusps all rounded and prominent, one median and four lateral pairs, separated by U-shaped incisions; cusps usually more or less equal in size. Median inferior nematothecae

Known range: Bermuda (Congdon 1907, Smallwood 1910, Bennitt 1922, Morris and Mogelberg 1973, Butler et al. 1983, Calder 1986, 1993, 1995, 1997); western Atlantic (Vervoort 1968); eastern Atlantic (Vervoort 1968); Indian Ocean (Millard 1975); western Pacific (Hirohito 1983).

Macrorhynchia philippina Kirchenpauer, 1872; Rees and Vervoort, 1987; Calder, 1986; Migotto, 1993; Calder, 1997. Aglaophenia philippina Kirchenpauer, 1872; Aglaophenia urens Kirchenpauer, 1876 [Nomem Nudum]; Lytocarpus urens Bale, 1888; Aglaophenia (Lytocarpus) phillipina Bale, 1888 not Lytocarpus phillipinus Congdon, 1907 [=Macrorhynchia allmani Nutting, 1900, according to Calder (1997a)]; Lytocarpus crosslandi Ritchie, 1908; Lytocarpus philippinus atlanticus Billard, 1913; Aglaophenia (Macrorhynchia) philippina Stechow, 1919; Macrorhynchia crosslandi Stechow, 1923a; Macrorhynchia urens Stechow, 1923a; Lytocarpus (Macrorhynchia) philippinus Boero and Bouillon, 1987.

Examined material: twenty-two colonies with gonophores and phylactocarps, Conchal, Guanacaste, Pacific coast, 10º23’55’’ N, 85º48’45’’ W, 31 September 1976 (UCR- 614).

Known range: Bermudas (Bennitt 1922, Calder 1983, 1986, 1993, 1997); western Atlantic (Vervoort 1968); eastern Atlantic (Ritchie 1908); Indian Ocean (Millard 1975); western Pacific (Hirohito 1983); eastern Pacific (Fraser 1948); Brazil (Migotto 1993).

Macrorhynchia sp.
Fig. 14)

Macrorhynchia Kirchenpauer, 1872, Calder, 1997a. Nematophorus Clarke, 1879. Pleurocarpa Fewkes, 1881.

Examined material: nine damaged colonies, Punta Cabuja, Guanacaste, Pacific coast, 10°22’20’’ N, 85°51’25’’ W, 16 November 1991 (UCR-894).

Lytocarpia tridentata (Verluys, 1899)
(Fig. 15)

Aglaophenia tridentata Verluys, 1899, Stechow, 1923b. Aglaophenia contorta Nutting, 1900; Bedot, 1921; Vannucci-Mendes, 1946; Vannucci, 1951. Thecocarpus contorta Totton, 1926, Aglaophenia tridentata Picard, 1951a; Vervoort, 1968; Forneris, 1969. Lytocarpia tridentata Migotto, 1993.

Examined material: nine colonies without gonophores, northern part of Isla Pájaros, Limón, Caribbean coast, 10°01’10’’ N, 83°04’45’’ W, 27 September 1996 (UCR-891).

Description: colonies up to 16 cm high, with a creeping hydrorhiza. Hydrocaulus monosiphonic and unbranched. Basal part of the colony usually without hydrocladia and nematothecae. Distal part divided into internodes by distinct transverse nodes. Internodes with one hydrocladial apophysis and four nematothecae: two axillary disposed, one inferior, and one frontal pseudonematothecae. Hydrocladia with distinct internodes, separated by oblique nodes. Each internode with one hydrotheca and three nematohecae: two laterally and one mesial. Hydrothecae with short internal septa, two almost imperceptible lateral teeth, and one conspicuos median tooth. Nine or eleven tentacles. Margin of hydrotheca about 166 mm in diameter and 280-336 µm deep. Mesial nematotheca up to 288 µm long; lateral nematothecae about 125 µm long. Perisarc of moderate thickness. Gonophores not observed. Cnidome comprising microbasic mastigophores of two categories: (i) small, 4.9-5.2 x 1.8-2.5 µm, and (ii) large, 28.0-36.0 x 4.1-5.2 µm.

Known range: western Atlantic (Vervoort 1968 as Aglaophenia tridentata); Brazil (Totton 1926, Vannucci-Mendes 1946, Vannucci 1951, Migotto 1993).

Clytia gracilis (Sars, 1850)
(Fig. 16)

Laomedea gracilis M. Sars, 1850; Laomedea (Campanularia) gracilis M. Sars, 1857; Clytia (Platypyxis) cilindrica L. Agassiz, 1862; Clytia cilindrica L. Agassiz, 1862; Platypyxis cylindrica L. Agassiz, 1862; Gonothyrea gracilis Allman, 1864; Campanularia gracilis Van Beneden, 1867; Clytia laevis Weismann, 1883; Campanularia serrulata Bale, 1888; Clytia serrulata Pictet, 1893; Laomedea (Gonothyraea) gracilis Levinsen, 1893; Campanularia attenuata Calkins, 1899; Obelia serrulata Thornely, 1900; Campanularia pelagica van Breemen, 1905; Clytia coronata Fraser, 1912; Clytia elsae-oswaldae Stechow, 1914; Clytia pelagica Billard, 1917; Clytia gracilis Stechow, 1923a; Calder, 1990. Laomedea (Phialidium) pelagica Hummelinck, 1930; Laomedea (Phialidium) gracilis Hummelinck, 1930; Laomedea pelagica Hummelinck, 1930; Laomedea (Clytia) gracilis Broch, 1933; Laomedea gracilis form pelagica Leloup, 1933; Clytia (campanularia) pelagica Kunne, 1937; Laomedea cylindrica Leloup, 1937; Clytia hemisphaerica Millard, 1966; Cornelius, 1982. Clytia elsaeoswaldea Vervoort, 1968; Campanularia (Clytia) cylindrica Vervoort, 1968; Laomedea (Clytia) pelagica Vervoort, 1972; Clytia sarsi Cornelius, 1982.

Material examined: twenty three colonies with gonothecae, Isla del Coco, Pacific coast, 5°29’81’’ N, 87°03’09’’ W, 17 June 1994 (UCR-832).

Description: colonies erect, fragile, sparingly and irregularly branched, up to 10 mm high, always arising from a creeping hydrorhiza. Hydrocaulus monosiphonic; each branch arising from a slightly curved aphophysis given off below hydrothecae of pedicel from which it arises; branches resembling primary pedicels and directed abruptly upwards, 6-10 annulations proximally and 2-6 distally placed. Pedicels about 0.5-0.4 mm high, 101- 142 µm in diameter, annulated basally and distally, with annuli or wrinkles elsewhere. Perisarc firm. Hydrothecae more or less cone-shaped, about 730-930 µm long, 390 µm wide at margin, 165-244 µm wide at diaphragm, with thin perisarc. Walls convex above diaphragm, nearly straight elsewhere; margin with about 12 to 17 deeply cut, blunt, triangular tooth cylindric by U-shaped incisions; hydrothecal walls weakly scalloped in cross section at margin. Diaphragm thin, straight; the basal chamber varied in depth but usually deep and cup-shaped. Tentacles small, 18-21 in number, but usually 20. Gonothecae urn-shaped, about 968-1000 µm long, and 404-432 mm in diameter; orifice diameter about 305- 342 µm wide, constricted just below aperture to about 320 µm wide, arising from hydrorhiza on short pedicels; walls smooth, constricted below aperture. Cnidome comprising ishorhizas about 5.8-6.5 x 1.6-1.9 µm and microbasic mastigophores of three distinct sizes; (i) small, 6.5-7.8 x 1.8-2.6 µm; (ii) medium, 8.4-9.2 x 2.8-3.1 µm; and (iii) large,13- 16.3 x 3.1-3.9 µm.

Known range: Bermudas (Bennitt 1922 and Calder 1986 as Clytia Cylindrical, Calder 1990); western Atlantic (Vervoort 1968 as Laomedea (Phialidium) pelagica); eastern Atlantic (Vervoort 1946 as Laomedea pelagica); Indian Ocean (Mammen 1965); western Pacific (Yamada 1959); eastern Pacific (Fraser 1948 as Gonothyrea gracilis); Brazil (Migotto 1993).

The list presented by Cortés (1997) includes 20 species of both Anthoathecatae and Leptothecatae orders. However, the reviewed synonym reduced this number to 19 (Table 2). With the additional contribution from this study, the actual number of Anthoathecate and Leptothecate species appears to be 35, of which only 16 are stored at the Museum of Zoology, University of Costa Rica.

The species Halopteris polymorpha (Billard 1913) previously reported in the Indian and Pacific Oceans (Billard 1913, Vervoort 1966, Millard and Bouillon 1973, Millard 1975, Hirohito 1983) and Brazil (Migotto 1993), is reported for first time from Caribbean Sea.

Another two species are new additions to the eastern Pacific fauna: Acryptolaria longitheca (Allman 1877), previously known from western Atlantic and Australia (Allman 1877, 1888, Clark 1879, Vervoort 1972, Calder 1996, Calder and Vervoort 1998), and Agloaphenia latecarinata Allman, 1877, from Bermuda (Congdon 1907, Smallwood 1910, Benitt 1922, Moris and Mogelberg 1973, Butler et al. 1983, Calder 1986, 1993, 1995, 1997a), western and eastern Atlantic (Vervoort 1968), Indian Ocean (Millard 1975), western Pacific (Hirohito 1983). Finally, the southerly range of Plumularia floridana Nutting, 1900, is now extended from Vicente Point, California to Punta Pitahaya, Guanacaste.

The Anthoathecatae and Leptothecatae hydroids are a diverse and conspicuous group of marine invertebrates. Most of the species have an apparent cosmopolitan distribution, with records from almost all oceans. However, the number of species reported from Costa Rica is still comparatively low, and unfortunately, the degree of preservation of the voucher material restricted the observation of any morphological variation. Additionally, although the hydroids are able to produce phenotypic responses to environmental changes (Kelmo and Attrill, in press), the absence of quantitative ecological data impedes the formulation of any robust conclusion about the distribution or ecology of the group. Therefore, there is a requirement for further baseline research on this group for both Pacific and Caribbean coasts of Costa Rica.

This work was made possible by financial support from CNPq-Brazil and Centro de Apoio ao Desenvolvimento Científico e Tecnológico - Secretaria de Planejamento e Tecnologia do Estado da Bahia (CADCT-SEPLANTEC). The authors wish to thank to Dale Calder from Centre for Biodiversity and Conservation Biology of Royal Ontario Museum (Canada) for critical comment of manuscript and to J. Cortés (Escuela de Biología y Centro de Investigación en Ciencias del Mar y Limnología, CIMAR), Universidad de Costa Rica for the collection of the specimens. Gratitude is expanded to Simone Souza de Moraes, Rilza da Costa Tourinho Gomes and Larissa de Siqueira, for all encouragement provided. This is a contribution of Museo de Zoología, Escuela de Biología, Universidad de Costa Rica.

Este estudio proporciona el primer informe taxonómico de los hidroides de los Ordenes Anthoathecatae y Leptothecatae presentes en las colecciones del Museo de Zoología de la Escuela de Biología de la Universidad de Costa Rica. Se informan 16 especies de hidroides: Eudendrium carneum, Pennaria disticha, Acryptolaria longitheca, Plumularia floridana, Halopteris polymorpha, Aglaophenia dubia, Aglaophenia latecarinata, Lytocarpia tridentata, Macrorhynchia phillipina, Macrorhynchia sp., Clytia gracilis, Cnidoscyphus marginatus, Thyroscyphus ramosus, Dynamena disticha, Sertularella diaphana y Tridentata distans. Se presenta una exhaustiva revisión de la sinonimia y una clave taxonómica simple e ilustraciones y descripciones para cada una de las especies.

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