Ectopleura crocea L. Agassiz 1862

Ectopleura crocea (L. Agassiz, 1862) (Figures 3���5; Tables 1���8) Parypha crocea L. Agassiz, 1862: 249, pls 23 - 2 a. Tubularia mesembryanthemum Allman 1871: 418, figs. 83���84; Hargitt 1927: 494; Yamada 1959: 16; Schmidt 1971: 32, pl. 2 B; Hirohito 1988: 18, fig. 4, pl. 1, fig. B. Tubularia crocea; Torrey 1902: 42, pl. 3, figs. 22���23; Rees 1963: 1223; Brinckmann-Voss 1970: 28, text-fig. 30���34; Calder 1971: 24, pl. 1 C; Genzano, Cuartas & Excoffon 1991: 69, pl. 5 C; Blanco 1994: 182; Genzano & Zamponi 2003: 306, 307, 309, Tables 2-3; Demicheli & Scarabino 2006: 530. Tubularia ralphi Bale 1884: 42; Watson 1980: 60, fig. 25���37; Watson 1982: 85, fig. 4.6.b, Plate 7.5. Tubularia gracilis von Lendenfeld 1885: 597, fig. 51���52. Tubularia sagamina Stechow 1907: 194; Yamada 1959: 16. Tubularia australis Stechow 1925: 196. Tubularia warreni Ewer 1953: 351, text-fig. 1���4; Millard 1959: 299; Millard 1966: 435; Millard 1975: 35, frontispiece, fig. 15 A���G. Ectopleura warreni; Migotto & Silveira 1987: 101, fig. 3; Migotto 1996: 25; Grohmann et al. 1997: 230, Table 1; Rosso & Marques 1997: 417, 420, 421, Table 1, Figure 4. Ectopleura crocea; Petersen 1990: 174, fig. 27; Schuchert 1996: 107, figs. 64 a���g; Schuchert 2001: 43; Bouillon et al. 2004: 104, fig. 55 E���F; Genzano et al. 2009: 37, 40, Tables 2-3, Fig. 3; Schuchert 2010: 357 -362, fig. 6. Ectopleura ralphi; Petersen 1990: 175; Schuchert 1996: 109; Migotto, Marques & Flynn 2001: 289, 290, 293- 297, figs. 5-6, Table 1. Pinauay ralphi; Marques & Migotto 2001: 475, 478, 480, figs. 2 B, 3, Table 1; Migotto, Marques, Morandini & Silveira 2002: 10; Marques & Migotto 1994: 173, 174, Tables 15.1, 15.2; Grohmann 2006: 103, 104, Tables 1-2; Oliveira, Marques & Migotto, 2006: 4, 10, fig. 5, Table 1; Oliveira & Marques 2007: 31; Grohmann 2007: page not numbered; Silveira & Morandini 2011: 5. Pinauay crocea; Marques & Migotto 2001: 480. Examined material. Brazil: State of Rio de Janeiro, Maca��, Cavaleiros Beach, 22 �� 24 ���S 41 �� 47 ���W, 15.viii. 2008, on rock, 95 % ethanol, coll. A.C. Morandini (MZUSP 1633); State of S��o Paulo: S��o Vicente, Vacas Beach, 23 o 58 S 46 o 23 W, 05.ix. 1991, intertidal fringe, on rock, under Phragmatopoma stripe, at the same level of Eudendrium, 4 % formalin, coll. A.C. Marques (MZUSP 414); Itanha��m, Saudade Beach, 24 o 10 'S 46 o 45 'W, 26.viii. 1991, intertidal fringe, on rock, under Phragmatopoma stripe, 4 % formalin, coll. A.C. Marques (MZUSP 406); Peru��be, Centro Beach, 24 o 19 'S 46 o 58 'W, 12.viii. 1992, intertidal fringe, on rock, sheltered place, forming an abundant stripe below Phragmatopoma, 18 o C, 28 ���, 4 % formalin, coll. A.C. Marques (MZUSP 433); Peru��be, Jureia-Itatins Ecological Station, 24 �� 34 ���S 47 �� 14 ���W, 15.ix. 2008, on rock, 95 % ethanol, coll. J.M.M. Nogueira (MZUSP 1636); Canan��ia, Argol��o Rocky Shore near S��o Jo��o Hill, 25 o00'S 47 o 57 'W, 25.viii. 1992, intertidal fringe, on rock, 19 o C, 28 ���, 4 % formalin, coll. A.C. Marques (MZUSP 444); State of Paran��: Mel Island, Encantadas Rock, 25 o 34 'S 48 o 18 'W, 06.viii.1988, 4% formalin, coll. M.A. Haddad (MZUSP 1750); Paranagu��, Yacht Club Paranagu��, 25 �� 30 ���S 48 �� 29 ���W, 10.x. 2007, on artificial substrate, 95 % ethanol, coll. M.A. Haddad, (MZUSP 1637); Guaratuba, bottom trawling net, 4 km of shore, 25 o 52 'S 48 o 33 'W, 01.xii.2003, 4% formalin, coll. M.A. Haddad (MZUSP 1751); State of Santa Catarina: Itapo��, Itapema beach, 26 o05'S 48 o 36 'W, 04.vi.2004, 4% formalin, coll. M.A. Haddad (MZUSP 1752); Penha, on a culture of mussels, 26 o 45 'S 48 o 38 'W, 17.vi.2005, 4% formalin, coll. M.A. Haddad (MZUSP 1753); Bombas, Bombas Beach, 27 o07���S 48 o 30 W, 03.xii.2006, 4% formalin, coll. M.A. Imazu (MZUSP 1754) and 95 % ethanol, coll. E. Ale, (MZUSP 1638). Argentina: Mar del Plata, Punta Cantera, 38 o04'S 57 o 32 'W, 26.i. 2002, intertidal fringe, 4 % formalin, coll. G. Genzano (MZUSP 1755), 95 % ethanol, coll. G. Genzano (MZUSP 1639). Type specimens: Ectopleura ralphi, type specimen lost, a neotype was proposed based on specimens from Australia, Victoria, Port Phillip, Yarra River Entrance Beacon, 03.iv.1977, 1��� 2m, on mussel and ascidia, formaldehyde (NMV G 3227) (Watson 1980). Ectopleura crocea, we found no reference concerning the material described by L. Agassiz, from the port of Boston. It may be lost. Description. Colonies dioecious, up to 55 mm high. Hydrorhiza and hydrocaulus with well-developed perisarc. Unbranched erect hydrocauli arising from stolonial hydrorhiza. Hydrocaulus��� coenosarc split into two longitudinal chambers with basal diameter 200���420 ��m, apical 340���1000 ��m; distal region of hydrocaulus with globular expansion supporting terminal hydranth. Hydranth with one whorl of aboral and one whorl of oral tentacles; oral tentacles adnate to hypostome up to the mouth region, circular in transversal section; aboral tentacles quadrangular in transversal section. Unbranched blastostyles of gonosomes arising immediately above aboral whorl of tentacles; main axis of each blastostyle supporting gonophores. Female gonophore cryptomedusoid, oval, with eight distal laterally compressed crests surrounding terminal aperture, terminal region of spadix projecting to outside. Male gonophore cryptomedusoid, spherical to oval, without distal crests. Early released actinulae with 8��� 11 aboral capitate tentacles. Aboral tentacles with four types of nematocysts: O-basitrichous isorhizas, rare and not measured (Figure 3 A); basitrichous isorhizas, common, 6.45���12.19 X 2.84���6.04 ��m (Figure 3 B); desmonemes, abundant, spherical to oval, 3.50���6.81 X 2.41���5.13 ��m (Figure 3 C); small stenoteles, abundant, 5.02���7.82 X 3.66���8.09 ��m (Figure 3 E). Oral tentacles with three types of nematocysts: O-basitrichous isorhiza, rare and not measured (Figure 3 A); basitrichous isorhizas, rare, 7.06���14.73 X 2.87���7.1 ��m; large stenoteles, abundant, 7.36���12.5 X 6.07���11.38 ��m and small stenoteles 4.93���7.93 X 3. 2���7.13 ��m (Figure 3 D). Distribution in the South Western Atlantic Ocean. Brazil: States of Esp��rito Santo (Grohmann et al. 1997, Grohmann 2006), S��o Paulo (Migotto & da Silveira 1987, Migotto 1996, Rosso & Marques 1997, Migotto et al. 2001, 2002, Marques & Migotto 2004, Oliveira et al. 2006, Oliveira & Marques 2007, Silveira & Morandini 2011), Paran�� (Haddad, 1992), Santa Catarina (Miranda et al. 2011) and Rio Grande do Sul (Migotto & Silveira 1987) (see Migotto et al. 2002, Marques et al. 2003). Uruguay (Demicheli & Scarabino 2006). Argentina: Provinces of Buenos Aires (Blanco 1994, Genzano et al. 1991, 2009, Genzano 1994, 1998, Genzano & Rodriguez 1998), R��o Negro and Chubut (Blanco 1994, Genzano et al. 1991), Santa Cruz and Tierra del Fuego (Oliveira et al. submitted). Remarks. General morphology. The species E. crocea and E. ralphi are morphologically similar and have been considered sister-taxa (Marques & Migotto, 2001) or synonyms (Bouillon et al. 2006, Schuchert 2010). Historically, subtle differences have been cited to differentiate the species (Table 1). For instance, Petersen (1990) differentiated the two species by describing E. crocea as having more aboral and oral tentacles than E. ralphi. However, this relationship appears to vary (Table 1). Indeed, characters related to the tentacles are generally variable (Tables 1���2; see also Agassiz 1862, Hargitt 1927, Ewer 1953, Calder 1971, Schmidt 1971, Migotto & Silveira 1987, Hirohito 1988). The SWAO specimens all have hydroucauli that broaden distally (Table 3), as described for E. crocea (Hirohito 1988, Schuchert 1996), although specimens with the same diameter throughout the hydrocaulus were reported by Petersen (1990). Each polyp is gonochoristic, although settlement of actinulae on already developed hydrocauli (Rungger 1969) may promote pseudo-hermaphroditism, a strategy also reported for other hydroids (Brinkmann-Voss 1970, Sommer 1990, Marques 2001, Schuchert 2010, Nawrocki & Cartwright 2012). There are inconsistencies in the description of the blastostyles, either characterized as unbranched (Ewer 1953 and Millard 1966 for E. ralphi; Brinckmann-Voss 1970, Hirohito 1988, and Petersen 1990 for E. crocea) or branched (Bale 1884, Millard 1975, Watson 1980, Migotto & Silveira 1987, and Petersen 1990 for E. ralphi; Schuchert 1996 for E. crocea) (Table 4). Descriptions of the neotype of E. ralphi from Melbourne (Australia) have different ways to describe the blastostyles, characterized either as ���[...] only occasionally branched��� (Schuchert 1996: 109; our underline) or as ���mature blastostyles branched [...]��� (Watson 1980: 61), or (���usually unbranched, but some branching can occur���, Schuchert 2010, p. 359). Specimens from SWAO have unbranched blastostyles (Figure 4 A���B), similar to those described by Allman (1871) and contrasting with the long bunches of gonophores described for other localities (Calder 1971, Millard 1975, Watson 1982, Petersen 1990). Millard (1959) commented on the difficulty of differentiating primary and secondary pedicels, which may explain the challenges of characterizing the ramification of blastostyles. (shaded rows), expressed as minimum���maximum values. Length and diameter are given in millimeters. T = Tubularia; E = Ectopleura. Hydranth Tentacles Locality length diameter aboral oral (n) number length number length Hydrocaulus Gonophores Locality length diameter length diameter (n) proximal medial distal S��o Vicente 14���35 240���300 300���400 340���540 260���500 180���300 (10) (20.7 �� 6.17) (280 �� 23.09) (350 �� 28.67) (436 �� 65.18) (368 �� 68.77) (228 �� 39.1) Author Female Male Blastostyles Actinulae (Original reference to species) gonophore gonophore Agassiz (1862) (Paripha 6���10 crests no crests simples or branched crocea) Allman (1871) 8 apical processes 4 small tubercles no oral tentacles (T. mesembryanthemum) Bale (1884) (T. ralphi) 4 tubercles branched Allen (1900) (Paripha crocea) 6���8 crests no crests branched Torrey (1902) (T. crocea) 8 crests top spherical, smooth or with processes Stechow (1907) (T. sagamina) 8 tubercles 6 tentacless Stechow (1925) (T. australis) 4 small tubercles 8 aboral, 5 oral tentacles Hargitt (1927) (T. mesembryanthemum) The morphology of the female gonophores (Figure 4 C) is a diagnostic feature for E. crocea (Rees & Thursfield 1965, Schmidt 1971, Petersen 1990, Schuchert 2001, 2010). Nevertheless, some variation may be observed, mainly because of the development of the gonophores, or due to their contraction (Torrey 1902, Schuchert 1996). It is hard to identify tubulariids without gonophores (Watson 1982), and this may be the cause of misidentifications between E. crocea and Ectopleura larynx (Ellis & Solander, 1786), especially for the Northwestern Atlantic (e.g. Fraser 1944). Male gonophores do not present apical crests (Figure 4 B; see also Agassiz 1862, Brinckmann-Voss 1970, Calder 1971, and Schuchert 1996, 2010 for E. crocea; Ewer 1953, Watson 1980, 1982, Migotto & Silveira 1987 for E. ralphi), although in some cases they are described with small apical processes, varying in size and development between colonies, and even in the same colony (Table 4) (Allman 1871, Hirohito 1988, and Petersen 1990 for E. crocea; Stechow 1925 and Millard 1975 for E. ralphi). Larval characters, such as tentacles, were already used to separate E. ralphi and E. crocea. A vague note in Schuchert (1996: 109, appears to refer to the observations of someone else) states that ���the actinulae of E. ralphi, however, are reported to have rudiments of oral tentacles which are absent from E. crocea ���. Other data refer to variations in morphology and number of tentacles of the actinulae (Ewer 1953, Brinckmann-Voss 1970, Millard 1975, Watson 1980, 1982, Migotto & Silveira 1987, Petersen 1990, Schuchert 1996, 2010; see Table 4). Presumably ���the oral tentacles will develop anyway immediately after the release of the actinula and the presence or absence in liberated ones is thus only a matter of timing��� (Shuchert 2010, p. 361), an interpretation that attenuates the importance of the variation. The actinulae of the SWAO present 8���10 (Brazil) or 8���11 (Argentina) capitate aboral tentacles, depending on their development, and do not have rudiments of oral tentacles (Figure 4 D��� E), but the variability of this character was never strictly assessed. Detailed studies on anatomical and histological characters have corroborated previous observations. Among these, the histological preparations confirmed that the oral tentacles are circular in a transversal section (Figure 5 A���B), while the aboral tentacles are squared (Figure 5 C), as described by Petersen (1990) for E. crocea. Also, transversal sections of the hydrocaulus have shown that its coenosarc is split into two longitudinal chambers (Figure 5 D), as already noted before (Ewer 1953 for E. ralphi; Allman 1871 and Schuchert 1996 for E. crocea), although this feature was considered to be inconstant in number and size (Millard 1959, 1975 for E. ralphi; Campbell & Campbell 1968, Hirohito 1988, and Petersen 1990 for E. crocea). Cnidome. The cnidome was uniform throughout all studied populations from SWAO (Figures 3 A, B, D, E). Literature data for E. crocea present few discrepancies (Table 5, contrasting with Table 6), for instance a cnidome restricted to stenoteles and desmonemes (Brinckmann-Voss 1970) and a potential contamination by microbasic euryteles (Schuchert 1996; in Schuchert 2010, p. 360, they are referred to ���rare euryteles���). Schuchert (1996) length 8.0��� 9.5 9.0��� 9.5 6.5 ���7.0 5.0���6.0 5.0��� 5.5 9.0 (Ectopleura crocea) width 7.0��� 9.5 3.0���4.0 5.0��� 5.5 3.5���4.5 3.0��� 3.5 4.0 S��o Vicente Itanha��m Peru��be Canan��ia Ilha do Mel Basitrichous length 8.58���10.74 6.45���10.41 7.77���12.12 8.45���10.55 * 1 8.14���11.86 (9.43 �� 0.48) (8.39 �� 0.67) (9.39 �� 0.67) (9.41 �� 0.46) (9.76 �� 0.8) Stenoteles length 5.57���7.34 5.59���7.22 5.31���7.43 5.45���7.82 5.48���7.64 (6.29 �� 0.32) (6.29 �� 0.31) (6.36 �� 0.37) (6.52 �� 0.44) (6.45 �� 0.41) (small) width 4.2���5.96 4.27���5.5 4.14���6.67 4.33���6.05 4.49���6.04 (4.93 �� 0.31) (4.85 �� 0.27) (5.07 �� 0.4) (5.06 �� 0.36) (5.27 �� 0.32) Basitrichous length 7.06���11.6 7.67���10.21 7.22���10.78 8.2���9.03 * 2 7.68���11.97 (9.32 �� 0.74) (8.67 �� 0.4) (9.46 �� 0.63) (8.62 �� 0.59) (9.79 �� 0.79) width 2.87���5.24 3.19���4.81 3.04���4.94 3.75���4.09 * 2 2.94���5.67 (3.95 �� 0.46) (3.91 �� 0.31) (3.91 �� 0.37) (3.92 �� 0.24) (4.46 �� 0.49) Oral Stenoteles length 7.37���11.88 7.43���11 7.36���11.37 8.27���11.47 * 3 7.99���11.72 (9.72 �� 0.89) (9.53 �� 0.58) (9.68 �� 0.68) (10.2 �� 0.77) (10.61 �� 0.59) tentacles (large) width 6.07���10.29 6.45���9.86 6.53���10.4 6.21���10.31 * 3 7.07���10.42 (8.36 �� 0.86) (8.23 �� 0.66) (8.81 �� 0.71) (8.79 �� 0.86) (9.37 �� 0.6) Stenoteles length 5.22���7.31 5.34���6.96 5.52���7.93 5.66���7.2 * 4 5.64���7.47 (6.19 �� 0.4) (6.06 �� 0.3) (6.29 �� 0.49) (6.4 �� 0.37) (6.42 �� 0.38) (small) width 4.24���6.5 4.1���6.45 4.33���7.02 4.34���5.82 * 4 4.41���6.46 (5.03 �� 0.37) (4.84 �� 0.32) (5.14 �� 0.53) (5.06 �� 0.34) (5.3 �� 0.45) Guaratuba Itapo�� Penha Bombas Mar del Plata Basitrichous length 8.04���12.19 8.91���11.91 8.65���11.56 7.37���10.61 7.81���10.43 (9.94 �� 0.75) (10.6 �� 0.6) (10.12 �� 0.58) (9.24 �� 0.56) (9.37 �� 0.52) width 3.06���5.87 3.73���5.82 3.37���5.29 3.54���5.33 3.17���5.26 (4.41 �� 0.5) (4.69 �� 0.4) (4.4 �� 0.4) (4.28 �� 0.37) (4.35 �� 0.43) Aboral Desmoneme length 3.94���6.81 4.57���6.66 4.51���6.6 4.67���6.54 4.02���6.17 (5.14 �� 0.53) (5.46 �� 0.39) (5.67 �� 0.41) (5.68 �� 0.44) (5.2 �� 0.37) tentacles width 2.53���4.49 3.2���5.13 3.2���4.9 3.2���4.85 3.08���4.36 (3.67 �� 0.46) (4.14 �� 0.33) (4.12 �� 0.38) (4.08 �� 0.31) (3.67 �� 0.28) Stenotele length 5.02���7.3 5.77���7.31 5.37���7.66 5.13���7.69 5.26���6.96 (6.19 �� 0.47) (6.62 �� 0.3) (6.47 �� 0.42) (6.4 �� 0.42) (6.17 �� 0.32) (small) width 3.66���5.69 4.83���8.09 4.3���6.06 4.27���6.04 4.24���5.78 (4.72 �� 0.42) (5.49 �� 0.47) (5.01 �� 0.35) (5.0�� 0.37) (4.88 �� 0.3) Basitrichous length 7.9���11.16 7.72���13.05 7.94���11.38 7.8���11.34 7.65���14.73 (9.38 �� 0.7) (10.38 �� 0.7) (9.7 �� 0.77) (9.39 �� 0.65) (9.32 �� 0.94) width 2.87���5.18 3.51���5.56 3.42���5.52 3.41���5.73 3.28���7.1 (4.21 �� 0.45) (4.55 �� 0.47) (4.28 �� 0.41) (4.23 �� 0.4) (4.24 �� 0.55) Oral Stenotele length 8.64���11.87 8.19���12.15 9.25���12.5 8.98���12.22 8.68���12.09 (10.31 �� 0.58) (10.29 �� 0.84) (10.8 �� 0.56) (10.3 �� 0.63) (10.42 �� 0.71) tentacles (large) width 7.64���9.87 7.05���11.16 8.19���11.38 7.64���10.38 7.69���10.49 (8.67 �� 0.56) (9.19 �� 0.8) (9.76 �� 0.61) (9.09 �� 0.52) (8.98 �� 0.59) Stenotele length 5.03���7.08 5.09���7.37 5.28���7.22 4.93���7.48 5.23���7.26 (6.12 �� 0.41) (6.4 �� 0.46) (6.32 �� 0.38) (6.18 �� 0.47) (6.19 �� 0.37) (small) width 3.2 ��� 3.2 ���6.00 4.21���7.13 4.43���5.78 3.9���6.02 4.05���5.84 (4.6 �� 0.43) (5.34 �� 0.4) (5.07 �� 0.32) (4.95 �� 0.43) (4.96 �� 0.38) The basitrichous isorhizas found in SWAO specimens were characterized for ��� Tubularia larynx ��� Ellis & Solander, 1786
Published as part of Imazu, Maur��cio Antunes, Ale, Ezequiel, Genzano, Gabriel Nestor & Marques, Antonio Carlos, 2014, A comparative study of populations of Ectopleura crocea and Ectopleura ralphi (Hydrozoa, Tubulariidae) from the Southwestern Atlantic Ocean, pp. 421-439 in Zootaxa 3753 (5) on pages 424-435, DOI: 10.11646/zootaxa.3753.5.2, http://zenodo.org/record/227863
{"references":["Agassiz, L. (1862) Contributions to the natural history of the United States of America. Vol. IV, Part III. Little Brown & Company, Trubner & Company, Boston, London, 398 pp.","Allman, G. J. (1871) A monograph of the gymnoblastic or tubularian hydroids. Ray Society, London, 450 pp.","Hargitt, C. W. (1927) Some hydroids of South China. Bulletin of the Museum of Comparative Zoology at Harvard College, 67, 491 - 520.","Yamada, M. (1959) Hydroid fauna of Japanese and its adjacent waters. Publications from the Akkeshi Marine Biological Station, 9, 1 - 101.","Schmidt, H. E. (1971) Some new records of hydroids from the Gulf of Aqaba with zoogeographical remarks on the Red Sea area. Journal of the Marine Biological Association of India, 13, 27 - 51.","Hirohito, Emperor Showa (1988) The hydroids of Sagami Bay. Biological Laboratory, Imperial Household, Tokyo, 179 pp.","Torrey, H. B. 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