Key words
Corals, Cirripathes secchini n. sp., Antipathes hirta, Antipathes fernandezi, Brazil.
The order Antipatharia (black corals) comprises approximately 200 recognized species. Several monographs ( Brook 1889 , Schultze 1896 , Cooper 1909 , Van Pesch 1914 and Pax 1918 ) dealed with the order's taxonomy and geographical distribution, many of them associated to long range expeditions (Challenger, Siboga, British Antantic "Terra Nova" and Percy Sladen Trust). Van Pesch (1 914) carried out the last revision of the group. Since then, efforts have been made lo clarify the systematic status of the group, mainly by Opresko ( 1972 , 1974 ).
Cooper (1909) , mentions that the number of mesenteries, the presence of a branched or unbranched axis, and the continuity or partial spacing of the polyps, allow the specimens to be located in recognizable genera, like Antipathes, Cirripathes and Stichopathes. However, the characters in the species level are very poorly defined, and many species descriptions are not detailed enough lo allow a positive identification. This, along with the lost of many type specimens (Opresko, pers. comm.), causes that "misidentifications are inevitable" ( Grigg and Opresko 1977 ). Most of the descriptions were based in a single specimen or in a fragment, not considering intra-specific variation, which seems to be a very common feature in the antipatharians.
Large monographs dealing with the order were based mainly on collections from the Indo-Pacific ( Brook 1889 , Schultze 1896 , Cooper 1909 , Van Pesch 1914 and Pax 1918 ). Species from the Atlantic Ocean were poorly describes in scattered papers, being discussed later in Brook (1889) and Van Pesch (1914) monographs without a conclusivo result (Opresko 1972 ).
Opresko (1972) redescribed and reevaluated twelve Atlantic antipatharians previously describes by L.E de Pourtalès. Opresko mentions approximately 32 species frorn the Atlantic Ocean. In later works, severas new species were describes ( Wamer 1981 , Opresko and Cairns 1992 , Opresko 1993 , 1996 , Caims et al. 1993 and Opresko and Sánchez 1997 ), resulting in approximately 39 recognized species from this arca.
In the present paper a new species (Cirripathes secchiini) is proposed. The description is based in a large number of measures in twenty specimens from the same locality, in order to deal with part of the intraspecifical variation of the species. Two species (Antipathes hirta Gray, 1857, first record from the Southwestern Atlantic and Antipathes fernandezi Pourtalès, 1874 , first record from the Atlantic Ocean) are compared to the more recent redescriptions (Opresko 1972 ) and new data on the size distribution of the characters are provided. The studied material, including C. secchini holotype and paratypes, were deposited in the Cnidaria collections of the Museu Nacional do Rio de Janeiro-UFRJ. Acronym for Museu Nacional do Rio de Janeiro is MNRJ.
Holotype: MNRJ 2757. specimen 18, 1 colony wet / 18 august 1993 / C.A. Echeverría and C.B. Castro / 20 m depth.
Type locality: Recifes de Popa Verde, Abrolhos, Bahia, Brazil. 18º 00.2' S, 039º 03.1 'W; 10 to 25 m depth.
Paratypes: MNRJ 2757, 19 colonies wet, same locality.
Other material: Type specimen of Stichopathes luetkeni Brook 1889, Zoologisk Museum, Denmark.
Diagnosis: Colonies unbranched, 1.55-2.92 m in height, showing a basal plate. Axis straight for 25 to 81% of the total length, then curving in regular, step, well defined spirals or in a sinuous stem. Spiral's height (when defined) 9-57 cm, diameter 5-22 cm; spiral's number 1 to 5.5. Axis diameter just above basal plate 3.7-9 mm; at tip of colony 0.7-2 mm.
Larger spines around stem; conical, laterally compressed, rounded at tip, forming a 90º angle with axis; showing small tubercles or protuberances (Fig. 1 ). Spines size 100 to 330 
m (individual colonies average 140-320 m, n = 19 colonies). Distance between consecutiva spines 310-930 m.
Small spines distributed irregularly between large spines, with no apparent pattern (Fig. 1 ). Small spines conical, sharp, forming a 90º with axis, size 30-82 m, with no tubercles or protuberances.
Polyps white, transverse diameter about 2-3 mm; longitudinal diameter 1-2 mm, artanged in multiple irregular rows, except in apical part of axis (apparently in a single row); one side of entire stem usually free of polyps. Sagittal tentacles about 1.3-2.15 mm; lateral tentacles 1.05-1.45 mm. Oral cone about 0.50.7 mm.
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Description: Colonies unbranched, 1.55-2.92 m in height, attached to substrata (reef calcareous structure) via a basal plate. Colonies almost straight for a variable distance (about 0.8 to 2.15 m, or 25 to 81% of the total length), then recurving forming regular or irregular step spirals. Colonies total length (measured along the axis) about 1.68-4.37 m. Spirals sometimes not well defined, with distal part of colony only slightly sinuous (15% of colonies showed this pattern, n = 20). Spirals height about 9-57 cm, diameter about 5-22 cm; spirals per colony about 1 to 5.5. Axis diameter gradually tapering; diameter immediately above hasal plate (basal diameter) about 3.79 mm; at end of straight part of axis about 2.85.5 mm; at middle point of curved part (spiraled) about 1.4-3.8 mm; at colony tip about 0.7-2 mm. Colonies height / basal diameter average 0.465 (± 0.115, n = 20).
Spines (Fig. 1 ) showed two size classes: small (modal class: 50 m); and large (modal class: 160 m, n = 2 498; see Fig. 2 ). Size and distribution patterns of spines did not valy around the colony. Larger spines arranged in rows. Small spines distributed between larger ones (Fig. 1 ), not arranged in any particular fashion, showing no preferences for any side of colonies.
Larger spines mainly conical ( Fig. 1 ), slightly laterally compressed, rounded at tip, forming a 90º angle with stem, covered with small tubercles from apex to middle of spines, and sometimes to its bases. Large spines colonies average variation about 140-320 m (n = 2 498 spines in 19 colonies). Distance between consecutiva spines in the same row about 310-930 m (average per colonie range about 360-590 m).
Small spines (Fig. 1 ) conical, sharp, forming a 90º angle with stem, length about 10-80 m (colonies average about 30-82 m, n=579 spines, in 19 colonies). Few small spines, usually those in the upper size class distribution, showed tubercles or protuberances in its surface.
Polyps white (Fig. 3 ), about 2-3 mm in transverse (related to stem) diameter and 1-2 mm in longitudinal diameter (parallel to stem), measured between external basis of opposite tentacles. Polyps arranged in continuous multiple irregular rows, leaving one side of stem free of polyps along entire colony. Occasionally arranged in a single row in apical part of stem. Distance between consecutive polyps varied from imperceptible, with polyps "crowded" in younger parts of colony, to a distance of 7 mm at the older parts of colony, Common interpolypar distance approximately 2.5 mm. Sagittal tentacles long, about 1.3-2.15 mm in length. Proximal tentacles shorter, about 1.05-1.45 mm in length. Oral cone ( Fig. 3 ) always present, about 0.5-0.7 mm in height.
Underwater observations of the polyps of C. secchini reveled that they are heavily contracted when fixed, resulting in less reliable measures. Warner (1981) also found a great polyp contraction in severas species of the genus Antipathes during their fixation.
Remarks: Van Pesch (1914) carried out the last revision of the genus Cirripathes and synomized severas species. This author recognized as valid only four previously describes species: C. anguina (Dana), C. spiralis (Linnaeus), C. ? paucispina Brook, and a species without name describes by Simpson and Thompson in 1905 (see Van Pesch 1914 ). He also describes six new species: C. nana, C. translucens, C. ramosa, C. contorta, C. musculosa and C. rumphii. This author also proposed to include the two unbranched genus of antipatharians (Cirripathes and Stichopathes) as subgenus of Cirripathes, with the name of Eucirripathes and Stichopathes. In later works, authors ( Pax 1918 , Pasternak 1977 , Opresko and Genin 1990 ) argued for retaining them as separate genera. This latter position is adopted here.
The main difference between Cirripathes and Stichopathes is the polyp arrangement. Several unbranched species were describes only on the basis of colony growth form, not including polyps. Therefore, Stichopathes luetkeni type was compared to the C. secchini description.
The colony growth form in C. secchini is very similar to Brook's (1889) S. luetkeni description. Larger spines (Fig. 4 ) size (not included in Brook's description; checked by the author on the type specimen) are also very similar. The shape of the spines, however, differs in this species mainly by the fact that their are curved upwards in the type of S. luetkeni. Another differences between these species are:
1. The existence of two different kind of spines in opposite sides of S. luetkeni (Fig. 4 ). This is mentioned in Brook's description, and is typical of the antipatharians that bear polyps in only one side of the stem. Although Brook's description was based in a dry specimen and he does not mention polyps, he places this species in the genus Stichopathes (polyps unisseriately arranged in one side of the stem). In C. secchini, larger spines occur all around the axis, with the small spines between them (Fig.1 ). lt also shows the polyps distributed in severas rows without a specific arrangement, which is a characteristic of the genus Cirripathes.
2. Cirripathes secchini shows larger and better defined spirals that S. luetkeni. lt also shows a comparatively greater stem diameter. Although this can be related to the colonies development status, similar length specimens compared with the type of S. luetkeni also showed this difference. The type specimen of S. luetkeni is very similar to the C. spiralis description ( Linnaeus 1758 ), and Van Pesch (1914) suggested that they were synonymous. C. spiralis, however, is redescribed by Van Pesch as showing a great number of close spirals, about 1-2 cm in diameter. The type specimen of S. luetkeni shows spirals usually with larger diameter, but greatly varying in shape and size.
Another species bearing small secondary spines, although they are not from the Caribbean area, are compared here to C. secchini. Not all these species were considered valid by Van Pesch (1914). The genus is in a great need of revision.
]]> Cirripathes diversa Brook 1889 : Brook mentions that the axis is "spiraled as in Cirripathes spiralis" (Van Pesch synomized both in 1914), with large spines with a blunt apex and small triangular spines between them. The poor description makes difficult any discussion, except by the fact that C. spiralis shows severas small (diameter aprox. 2 cm) spirals, and C. secchini shows few spirals, but with a larger diameter.Cirripathes flagellum Brook 1889 : Brook mentions secondary spines distributed between larger spines. The axis of this species is not spiraled. Summers (1910) suggested to place this species in the genus Stichopathes.
Cirripathes gardineri Cooper 1903 : A very poor and incomplete description. Van Pesch (1914) synomized this species with Cirripathes anguina Dana, that shows only primary spines. Opresko (1974) mentions that Cooper's (1903) descriptions are very abbreviated and, although they have figures, are insufficient to determine the validity of the species.
Synonims: Antipathes hirta Gray, 1857: 293.- Opresko, 1972: 979 - 984. ]]>
Antipathes picea Pourtalès, 1880: 115, PI. 3. Figs. 9, 29.Parantipathes? hirta -- Brook, 1889: 144, PI. 2, Fig. 11, PI. 11, Fig. 1.
Antipathes picea -- Brook. 1889: 161.
Paratitipatlies hirta -- van Pesch, 1914: 20.
Material examined: Recifes de Popa Verde, Abrolhos, Bahia, Brasil: MNRJ 3071, 1 colony; MNRJ 2577, 1 colony; MNRJ 2548, 8 colonies; MNRJ2467, 2 colonies; MNRJ2579, 1 colony; Recife de Timbebas, Abrolhos, Bahia, Brasil; MNRJ 2847, 1 colony.
Description: Colony ( Fig. 5 ) sparsely branched to third or fourth order, branches arising laterally and at right angles to lower ramifications; stem and branches with four to six longitudinal rows of primary pinnules arranged bisserially and in alternating groups along the length of axis; posterior primarias 2.2-4.6 cm long (colonies average variation about 1.5-3.25 cm); secondary pinnules about 0.68-1.53 cm (colonies average variation), in a single series on basal half of posterior (abpolypar) side of primaries; tertiary pinnules few in number (one to tree) and restricted to posterior side of those secondaries nearest the base of each primary; quatemary pinnules rarely present.
Spines smooth, conical, with an acute apex; maximum polypar spine's length per colony 0.12-0.3 mm; colonies average variation 0.107-0.265 mm; maximum abpolypar spine's length per colony 0.08-0.16 mm length, colonies average variation 0.066-0.118 mm; strongly distally inclined (30º-45º) and hooked upward, arranged in eight to ten longitudinal rows.
Polyps 0.22-0.51 mm long (10-16 per centimeter) in a single series orientated perpendicular to primary dorsoventral axis of colony; tentacles 0.12-0.360 mm long; oral cone raised about 0.1-0.2 mm (Table 2 ); mouth usually sagittally elongated.
]]> Geographic distribution: West Indies, Brazil.Remarks: Warner ( 1981 , p.151) proposed a new species (Antipathes thamnea) very similar to A. hirta, and mentions that "differences arise in details of pinnation and in the size of spines and polyps". This author mentions 0.52-0.64 mm for A. thamnea polyp's size. Opresko (1972) mentions 0.70.8 mm length to A. hirta polyps. The A. hirta specimens studied herein showed even smaller polyps (0.22-0.51 mm) than A. thamiiea description (Table 3 ). This suggests that the polyp's size in A. hirta would be about 0.22-0.8 mm in length. Considering also the contraction of the polyps during fixation, polyp's size does not seem to be a very reliable character in order to separate these two species. Warner (1981) , however, mentions that the measurements where carried out in fresh specimens, and that a great contraction (1/3 to half of the original size) occurs in this process. Taking this contraction into account, there seems to exist an overlying in polyp's size gradient between A. thamnea and A. hirta. Opresko (1972) also mentions a polypar spine's length of 0.07-0.13 mm to A. hirta. Warner mentions, 0.09-0.21 mm to A. thamnea polypar spines length and 0.05-0.19 mm to the abpolypar spines. The knew polypar spine's size distribution in A. hirta would be about 0.07 mm (Opresko 1972) to 0.3 mm (this work, see Table 1 ). Again, there appears to exist an overlying on the spine's length gradient between these two species.
I found a great variability in the characters size of the herein studied specimens (Fig. 6 , Table 2 ) even in the same colony (Fig. 7 ). lt is likely that the characteristics used to define A. thamnea could he included in the intraespecific variability observed in A. hirta. The great variability in spines size and pinnules arrangement seems to be a common feature in the order Antipatharia; therefore, a large number of measures in severas specimens from diverse localities, are necessary to verify the variability of a species character. Unfortunately, most of species descriptions were based in fragments from dredgings, with soft tissues very damaged or lost.
Synonims: Antipathes femandezii Pourtalès, 1874: 47.
Antipathes fernandezii -- Pourtalés, 1880: pl. 3, fig. 20.
Parantipathes? fernandezi -- Brook, 1889: 144.
Parantipathes? fernandezi -- Looser, 1926: 272, fig. 38.
Material examines: Antipathes fernandezi: one complete colony and severas small fragments (MNRJ 3188) - off Rio Grande do Sul, between Rio Grande do Sul and Chuí. N/Oc Saldanha, Esta. 14, Sul 11, March, 1972:
Description: Colony 4.86 cm height, mainly in one plane; axis diameter above basal plate 0.75 mm; stem and branches pinnate; pinnules simple and bilateral, arranged alternately along the axis; interpinnular distance 24 mm; pinnules average size 1.39 cm; pinnules diameter 0.20-0.25 mm.
Spines long, conical and acute, forming regular groups of longer spines alternating with relatively shorter spines groups along pinnules; slightly longer in one side of axis; maximum length on branches 0.24 mm, average 0.17 mm (n = 41; S: 0.04); pinnules spines maximum 0.31 mm, average 0.21 mm (n = 42; S: 0.06); arranged in eight longitudinal rows; 55-60 per centimeter in each row. These colonies bear no polyps.
Geographic distribution: Juan Fernandez Island (Chile), Rio Grande do Sul (Brazil).
]]> Remarks: Opresko's ( 1972 ) diagnosis did not mention the spines size. In his redescription of the type specimen, however, this author mentions 0.13-0.17 mm in length. In the herein studied material, the maximum spine length is about 0.31 mm. This spine length is relatively uncommon, occurring mainly on some spines of the pinnules apex. Average length seems to be a most reliable character than maximun spines length.The split of the Antarctic Convergence waters by the South-American continent in two branches (the Falklands current, that reaches southern coast of Brazil; and Humboldt current, that reaches western coast of Chile and Juan Fernandez Island), may justify the occurrence of A. fernandezi in both sides of South America. Since antipatharians are common in the continental shelf and in deeper waters, it is unlikely the existence of a barrier to its distribution in both sides of southern south-American continent.
Acnowledgments
I wish to thank Dennis M. Opresko, for suggestions and encouragement; Ole S. Tendal (Zoologisk Museum, Denmark), for his gently loan of the type specimen of Stichopathes luetkeni. Clovis B. Castro (Museu Nacional do Rio de Janeiro/UFRJ) for field and laboratory facilities. Thanks to three anonymous reviewers for comments and suggestions that greatly contributed to improve this manuscript.
Resumen
Se describe una nueva especie, Cirripathes secchini (Cnidaria: Antipatharia) proveniente de la región de Abrolhos (Bahia, Brasil), que constituye además el primer registro de este género en Brasil. Otras dos especies (Antipathes hirta, primer registro en el Océano Atlántico sur-occidental; Antipathes fernandezi, primer registro en el Océano Atlántico) se comparan a sus redescripciones más recientes y se agregan nuevos datos sobre la distribución de tamaño de sus caracteres taxonómicos. Hasta el momento existen apenas dos registros del orden Antipatharia en Brasil.
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* Dto. de Zoología, Instituto de Biologia, Universidade Federal do Rio de Janeiro. Ccs, Bloco A, Ilha do Fundao. 21940-590, Rio de Janeiro, RJ, Brazil. Tel./Fax: 055-021-560-5993, ramal 31, e-mail: caecheve@acd.ufrj.br