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Zootaxa 307: 1–12 (2003) www.mapress.com/zootaxa/

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Eudendrium caraiuru sp. n. (Hydrozoa; Anthoathecata; Eudendriidae) from the southeastern coast of Brazil ANTONIO C. MARQUES1 & OTTO M.P. OLIVEIRA1,2 1

Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11461, 05422970, São Paulo, SP, Brazil; e-mail: [email protected] 2 Centro de Biologia Marinha, Universidade de São Paulo, Caixa Postal 83, 11600-970, São Sebastião, SP, Brazil; e-mail: [email protected]

Abstract Eudendrium caraiuru sp. n. is described for the southeastern coast of Brazil. The species was reported previously from the area as E. glomeratum, a common species of the Mediterranean Sea. However, morphological, morphometrical and ecological data suggest they are diverging lineages, requiring a new specific name for the Brazilian population. Key words: Hydrozoa, Eudendriidae, Eudendrium, Brazil, new species, Eudendrium glomeratum, taxonomy

Introduction Nine species of Eudendrium have been recorded to date from Brazilian waters: Eudendrium carneum Clarke, 1882 (Vannucci 1954; Tommasi 1970; Rosa 1973; Masunari 1983; Souza 1987; Correia and Loyola e Silva 1990; Pires et al. 1992; Marques and Moretzsohn 1995; Grohmann et al. 1997; Nogueira et al. 1997; Rosso and Marques 1997; Calder and Maÿal, 1998; Marques 1993, 2001); Eudendrium capillare Alder, 1856 (Vannucci 1954; Mayal 1973; Alves and Mayal 1990; Marques 1993, 2001); Eudendrium rameum (Pallas, 1766) (Mayal 1973); Eudendrium pocaruquarum Marques, 1995 (Marques 1995, 2001; Rosso and Marques 1997); Eudendrium ? fragile Motz-Kossowska, 1905 (Grohmann et al. 1997); Eudendrium nambuccense Watson, 1985 (Marques 1993, 2001; Nogueira et al. 1997); Eudendrium glomeratum Picard, 1951 (Marques 1993, 2001; Rosso and Marques 1997; Oliveira et al. 2000); Eudendrium ramosum (Linnaeus, 1758) (Marques 1993, 2001; Grohmann et al. 1997; Rosso and Marques 1997); Eudendrium merulum Watson, 1985 (Marques 1993, 2001); Eudendrium sp. (Eston et al. 1986; Costa, 1992; Nogueira et al.

Accepted: 19 September 2003; published: 24 September 2003

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1997). However, only the papers by Vannucci (1954) and Marques (1995, 2001) provide detailed descriptions of the species, and only those by Marques (1993, 1995, 2001) include data on the cnidome. Two out of the nine species (viz., E. ? fragile from Vitória, Espírito Santo State, and E. rameum from Itamaracá Island, Pernambuco State) were not included in the partial review by Marques (2001) because no material was found in the collections studied by him. Eudendrium pocaruquarum is a species so far known exclusively for Brazilian waters and the remaining six species have a wide geographical distribution. Some materials from Banyuls-sur-Mer, Villefranche-sur-Mer, Majorca and Cabrera, with the characteristic presence of large nematocysts aggregated in pads around the body of hydranth, were described and assigned to Eudendrium ramosum by Motz-Kossowska (1905). Based on this diagnostic character, Picard (1951) gave the nomen novum E. glomeratum to refer to the species described by Motz-Kossowska (1905) and, as a by product of the new name, established the importance of the study of the cnidome in the systematics of the family Eudendriidae. Eudendrium glomeratum was found to be a very common species in the Mediterranean Sea (see review in Marques et al. 2000a), but it was also recorded from the Indian Ocean (Jäderholm 1916 and Thornely 1904, after Watson 1985: 213), Pacific Ocean (Boero and Cornelius 1987) and Atlantic Ocean (Boero and Cornelius 1987; Marques 1993; 2001). The Brazilian hydroid heretofore referred to as Eudendrium glomeratum, a common shallow water species, has been described by Marques (2001). A morphometrical analysis of the species revealed some inconsistencies with the populations of E. glomeratum from the Mediterranean Sea (see Oliveira et al. 2000). A more detailed analysis of populations from both the Mediterranean and from Brazil have proved they belong to different lineages. Therefore, the purpose of this study is to describe Brazilian material as Eudendrium caraiuru sp. n.

Material and methods Colonies of Eudendrium caraiuru sp. n. were collected by hand on rock and artificial substrates in the shallow waters along the coast of the states of Rio de Janeiro (one site) and São Paulo (many sites). The specimens were anaesthetized in a 1:1 solution of 7.5% magnesium chloride solution and seawater, and preserved in 4% formaldehyde solution in seawater or alcohol 70%. The specimens were examined, measured, and drawn under microscope and stereomicroscope, both with camara lucida and photo devices. The cnidome terminology follows Weill (1934) and Mariscal (1974), and measures of nematocysts were made on non-discharged capsules. The L/W ratio (Kubota 1976) and S/C ratio (Watson 1987) are also provided. Living specimens were studied for observations on coloration of the specimens, distribution and number of nematocyst capsules, and morphology of discharged nematocysts. Stems for scanning electron microscopy (SEM) were

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preserved in 2.5% glutaraldehyde, post-fixed in 1% OsO4, dehydrated in a graded series of ethanol, dried in a critical-point drier, and sputter-coated with gold. Other study methods for Eudendriidae are from Marques (1995, 2001) and Marques and Migotto (1998). The material is deposited in the Collection of Cnidaria of the Museu de Zoologia da Universidade de São Paulo (MZUSP), Royal Ontario Museum (ROMIZ), and Museu Nacional da Universidade Federal do Rio de Janeiro (MNRJ).

Taxonomic part Eudendrium caraiuru sp. n. Figures 1–19 Eudendrium glomeratum; Marques, 1993: 68–75, pl. 3; 2001: 361–369, figs. 23–30; Migotto, 1996: 122; Rosso and Marques, 1997: 417; Oliveira et al., 2000: 519–525; Migotto et al., 2001: 289, 294–296; 2002: 11. non Eudendrium glomeratum Picard, 1951.

Type material. Holotype: Brazil: São Sebastião: Baleeiro Point, female colony, 08.iii.1988, formol, on rock, 3m, col. A.E. Migotto (MZUSP 0385; former ACM-SP029). Paratypes: Brazil: São Sebastião: Cigarras Beach: male colony, 15.vii.1988, formol, intertidal, col. A.E. Migotto (MZUSP 0388; former ACM-SP034); Pitangueiras Beach (north rocky shore): male colony, 24.x.1992, formol, on rock, intertidal, col. A.C. Marques (ROMIZ B1223; former ACM-SP162); Jarobá Point, Parque: male colony, 17.ix.1990, formol, on test panel, 2m, col. A.C. Marques (MZUSP 0394; former ACM-SP056); Baleeiro Point: ; Baraqueçaba Beach: female colonies, 12.xii.2001, formol, on metallic structures, 7m, col. H.K. Boscolo (MZUSP 0372); Lage dos Moleques: female colony, 05.xii.1991, formol, on rock, 5m, col. A.C. Marques (MZUSP 0423; former ACM-SP114). Additional material. Brazil. Rio de Janeiro: Urca: colony without gonophores, ix.1990, leg. I. Zalmon (MZUSP 0375; former ACM-RJ008); Ubatuba: Lázaro Beach: colony without gonophores, 28.vii.1992, formol, on rock, intertidal, col. A.C. Marques (MZUSP 0427; former ACM-SP120); colony without gonophores, 28.vii.1992, formol, on rock, intertidal, col. A.C. Marques (MZUSP 0428; former ACM-SP121); Maranduba Beach: colony without gonophores, 30.vii.1992, formol, on rock, intertidal, col. A.C. Marques (MZUSP 0429; former ACM-SP123); São Sebastião: Pier Sul (Petrobrás): male colony, 18.vii.1990, formol, on Perna perna, 1m, col. J.C. de Freitas (ACM-SP064); Pitangueiras Beach (north rocky shore): colony without gonophores, 18.viii.1988, formol, 6m, col. A.E. Migotto (MZUSP 0458; former ACM-SP159); male colony, 03.iv.1992, formol, on Schizoporella, 3m, col. A.E. Migotto (MZUSP 0459; former ACM-SP161); Jarobá Point (23º49,654´S 45º25,366´W): colony without gonophores 17.viii.1990, formol, on test panel, 1m, col. A.C. Marques (MNRJ 2043; former ACM-SP044); colony without

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gonophores, 17.viii.1990, formol, on test panel, 1m, col. A.C. Marques (ROMIZ B1221; former ACM-SP045); colony without gonophores, 20.viii.1990, formol, on test panel, 2m, col. A.C. Marques (MNRJ 2044; former ACM-SP048); colony without gonophores, 20.viii.1990, formol, on test panel, 2m, col. A.C. Marques (MNRJ 2045; former ACMSP049); colony without gonophores, 20.viii.1990, formol, on test panel, 2m, col. A.C. Marques (ROMIZ B1222; former ACM-SP050); colony without gonophores, 15.ix.1990, formol, on rope, 2m, col. A.C. Marques (MZUSP 0392; former ACM-SP054); male colony, 15.ix.1990, formol, on rope, 2m, col. A.C. Marques (MZUSP 0393; former ACMSP055); colonies without gonophores, 22.i.2000, formol, on test panel, 1m, col. O.M.P. Oliveira (MZUSP 0368); female colonies, 22.i.2000, formol, on test panel, 1m, col. O.M.P. Oliveira (MZUSP 0369); colonies without gonophores, 25.i.2002, alcohol, on ropes, 1m, col. O.M.P. Oliveira (MZUSP 0370); Barequeçaba Point (23º49,979´S 45º25,843´W): colonies without gomophores, 12.xii.2001, formol, on metallic structures, 7m, col. H.K. Boscolo (MZUSP 0371); female colonies, 12.xii.2001, formol, on metallic structures, 7m, col. H.K. Boscolo (MZUSP 0372); Cananéia: Cardoso Island, costão do Pereirinha: colony without gonophores, 26.viii.1992, formol, on rock, intertidal, col. A.C. Marques (MZUSP 0447; former ACM-SP145). Diagnosis. Large euryteles with shaft:capsule proportion 3.0–3.6; in pads on hydranth body, spadix of female gonophores, and in a whorl of 16–25 around hypostome. Female blastostyles with reduced hypostome and tentacles. Etymology. From the Tupi native language “cáraiurú” (= powerful mouth), in reference to the presence of large euryteles on the hypostome. Description. Colonies dioecious, arborescent, up to 54 mm in height, main stems unfascicled. Hydrocauli arising from creeping hydrorhiza; branches many, more or less alternate, occurring over entire hydrocaulus, branches up to third order, in radiate planes or rarely more or less planar; pedicels arising from main stem or branches of first, second or third order. Perisarc of main stem strongly developed, dark brown, single tubes 0.40 mm in diameter, with scarce annulations, in sets of 3–8 rings. Branches with 3–7 rings at origin, 0.20–0.25 mm in diameter. Pedicels sometimes completely annulated, yellowish, ca. 0.10 mm in diameter. Hydranths 0.18–0.75 mm in height, 0.18–0.57 mm in diameter (measured in the body region just below the tentacles), orange in color, with a distinct groove in the aboral region; tentacles 23–34 in number, occurring in a whorl below hypostome. Some hydranths with reduced tentacles juxtaposed in two close whorls. Gonophores styloids, arising from body of hydranth. Immature styloids placed in a circle around body of hydranth. Male blastostyle orange, with 10–29 sporosacs, each sporosac 1–2 chambered, linked to blastostyle body by a stalk, with a very distinct spadix over its longitudinal axis, and a terminal tubercle on its apex; distal chamber 0.12–0.18 mm in diameter. Male blastostyles completely reduced over earlier stages of their development with pedicels corrugated throughout. Female gonophores orange, arising on partially

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reduced blastostyles with highly atrophied hypostome and degenerated tentacles. Immature eggs having a simple and curved spadix over a single egg. Blastostyle reducing completely during development or at maximum with 1–5 stumps (tentacles), and spadices shed. Mature oval eggs thickened by perisarc and linked directly to the wrinkled pedicel by short and shallow concave peduncles. Eggs 4–7 in number, 0.24–0.39 mm in diameter. Nematocysts of two categories, heterotrichous microbasic euryteles and heterotrichous macrobasic (or mesobasic) euryteles.

FIGURES 1–8. Eudendrium caraiuru sp. n. 1, general aspect of the colony; 2, hydranths; 3, female blastostyle; 4, male blastostyle; 5, capsule of small microbasic eurytele; 6, discharged small microbasic eurytele; 7, capsule of large microbasic eurytele; 8, discharged large microbasic eurytele. Scale bars: 2.0 mm (1); 1.0 mm (2–4); 10 µm (5–8).



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FIGURES 9–15. Eudendrium caraiuru sp. n. 9, general aspect of the colony covered by debris. 10, oral view of the hydranth. 11, general aspect of the hydranth. 12, capsule of large microbasic euryteles. 13, primary polyp, derived from larval metamorphosis. 14, young hydranth encapsulated by perisarc. 15, degenerating hydranth. Legends: a, large microbasic euryteles on the hypostome; b, large microbasic eurytele pad on the body of the hydranth; c, perisarc involving hydranth; d, reduced tentacle. Scale bars: 1.0 mm (9); 200 µm (10); 300 µm (11,13–15); 10 µm (12).

Small microbasic euryteles (seen discharged), 6.1–8.0 X 2.9–3.9 µm, L / W = 1 : 2.05– 2.1, oval, abundant; distributed over hydranth body, hypostome, coenosarc, and tentacles. Large macrobasic euryteles (seen discharged), 18.7–22.7 X 7.1–9.3 µm, L / W 1: 2.4– 2.6, bean shaped, common. Discharged shaft up to 60 µm in length , 3.0–3.6 times length of capsule; undischarged shaft in 1.5 coils inside capsule; distributed on hydranth body sometimes forming pads to a continuous ring, up to 25 capsules on hypostome, coenosarc, terminal tubercle, and immature female spadix sometimes forming pads. Distribution. Brazil: Rio de Janeiro State: Rio de Janeiro (Marques, 2001); São Paulo: Ubatuba (Marques, 2001), São Sebastião (Oliveira et al., 2000; Marques, 2001), Cananéia (Marques, 2001).

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FIGURES 16–19. Eudendrium caraiuru sp. n. 16, general aspect of a stem; 17, oral view of the hydranth; 18, general aspect of hydranth; 19, hydranth base. Legend: e, groove. Scale bars: 1.0 mm (16); 100 µm (17–18); 40 µm (19).

Discussion The species name Eudendrium glomeratum was proposed as a nomen novum by Picard (1951) for a specimen described as Eudendrium ramosum (Linnaeus, 1758) by Motz-Kossowska (1905). Picard justified the erection of a new species based on the presence of macrobasic euryteles arranged in warts on the body of hydranth, but lacking on the hypostome (cf. Motz-Kossowska 1905; Picard 1951; Rossi 1961; Boero and Cornelius 1987; Marques et al. 2000a). The diagnosis of the species was amended two times. Initially, Watson (1985) re-defined the species to include specimens from the Mediterranean Sea in which the nematocysts formed a continuous ring around the body of hydranth. Later, it was amended again to include Brazilian specimens in which the nematocysts exhibited a number of differences from European material: shaft intermediate between macrobasic and microbasic forms, and a scarcity of these nematocysts on the hypostome, where they are never organized in warts (Marques 1993, 2001).



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The plasticity of the so called E. glomeratum was remarked by Marques et al. (2000a). These authors also included comparisons of the species with Eudendrium magnificum Yamada, 1954, and considered the possibility that the two might be conspecific. They pointed to the lack of identification of the large nematocyst type of the Japanese species (cf. Yamada 1954, 1959; Hirohito 1988, although the latter gave the dimensions of these large nematocysts). The muddled state of the taxonomy of E. glomeratum was also highlighted by Marques et al. (2000b). They compared the sexual female features of the species with those of Eudendrium cyathiferum Jäderholm, 1904, and noted the eventual broader distribution of E. glomeratum due to misidentification of the species particularly compared with E. rameum and E. ramosum. The studies above clearly pointed out the existence of a complex of species under the name E. glomeratum. Eudendrium caraiuru sp. n. has been reported so far from the Brazilian coast as Eudendrium glomeratum Picard, 1951. In fact, the two species share some characters heretofore considered diagnostic of E. glomeratum, such as the presence of large euryteles arranged in warts around the body of hydranth and on the immature unbranched female spadix, and the mature eggs encapsulated by perisarc scattered along the pedicel of completely reduced blastostyles, linked to those pedicels by short concave perisarc bases (cf. Marques et al. 2000a, 2000b). However, detailed morphological comparisons between E. caraiuru sp. n. ( “E. glomeratum” of Brazilian authors) and Mediterranean populations of E. glomeratum have shown that the height and width of Mediterranean specimens were ca. ten times those of the Brazilian specimens, and that the Brazilian colonies are unfascicled (Oliveira et al. 2000). Moreover, the cnidome of both species is also slightly different. Eudendrium caraiuru sp. n. has euryteles with smaller dimensions (18.7–22.7 X 7.1–9.3 µm, see also Marques 2001) than those of E. glomeratum from the Mediterranean Sea (22.2–27.2 X 9.5–12.0 µm, Marques et al. 2000a; 24–28 X 10–11 µm, Marinopoulos 1992) and similar to those from Australia (19.0–22.0 X 9.0–10.0 µm, Watson 1985). According to Weill’s (1934) definitions of nematocyst types, E. caraiuru sp. n. has microbasic euryteles instead of macrobasic euryteles. However, under Östman’s (2000) definition, the nematocyst is a mesobasic eurytele. The study of living material of E. caraiuru sp. n. corroborated the previous hypothesis by Marques (2001) that hydranths with two whorls of reduced tentacles are, indeed, related to the life cycle of the species. Cycles of renewal of hydranths are common in the species, taking about 8–36 hours to be completed and, during which, it is possible to see the presence of the two whorls of tentacles in resorbing hydranths. The feature is also present in certain other species of Eudendrium (cf. Marques et al. 2000b). There are also some striking differences between E. caraiuru sp. n. and E. glomeratum ecologically. After seasonal studies during 1999–2000, we found Brazilian populations of E. caraiuru sp. n. with no resting stages, living in temperatures over 21.5° C (Oliveira and Marques, unpublished data) and, during the last decade, the species was always found in

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temperatures of 19.5–27.7 ºC and salinities of 30.0–35.4 ppt (Marques 1993, and personal observations). Meanwhile, populations of E. glomeratum from the Mediterranean Sea have shown resting stages in warmer months (cf. Boero et al. 1986; Arillo et al. 1989; Bavestrello and Arillo 1992); however, the temperatures during these months were never over 21,5 ºC. Moreover, reproductive patterns are also different between the two populations (cf. Marques 1993 and Boero et al. 1986; Oliveira and Marques, unpublished data). All differences between the populations, when put together, are strong evidence that E. caraiuru sp.n. represents a different lineage from E. glomeratum. Still, evidence suggests that the remaining populations of E. glomeratum, from many places, may be cryptic species, needing detailed morphological studies (including morphometry, SEM), molecular analysis and assessment of ecological data.

Acknowledgements The authors are grateful to Alvaro E. Migotto for his help with the illustrations and for reviewing the text, and to Enio Mattos for the assistance with SEM. We thank Dale R. Calder and an anonymous referee for reviewing the text. We also thank the Centro de Biologia Marinha of the Universidade de São Paulo, in which part of the study was made, for logistic support. This research was supported, in different phases, by FAPESP (Proc. 1989/ 0625-0; 1991/1599-2; 1991/1600-0; 1995/3022-5; 1996/10544-0; 1999/11328-8; 2000/ 14932-2; 2001/02626-7) and CNPq (Proc. 300271/2001-8).

References Alder, J. (1856) A notice of some new genera and species of British hydroid zoophytes. Annals and Magazine of Natural History, 2nd series, 18, 353–362. Alves, J.M. & Mayal, E.M. (1990) Eudendrium capillare (Hydrozoa, Hidroida). Resumos do XVII Congresso Brasileiro de Zoologia, Londrina. Arillo, A., Bavestrello, G. & Boero, F. (1989) Circannual cycle and oxygen consumption in Eudendrium glomeratum (Cnidaria, Anthomedusae): studies on a shallow water population. Pubblicazione della Stazione Zoologica di Napoli, I: Marine Ecology, 10(4), 289–301. Bavestrello, G. & Arillo, A. (1992) Irradiance, temperature and circannual cycle of Eudendrium glomeratum Picard (Hydrozoa, Cnidaria). Bolletino di Zoologia, 59, 45–48. Boero, F. & Cornelius, P.F.S. (1987) Records of Eudendrium glomeratum (Cnidaria: Hydroida) in British and Irish waters, and taxonomic comments. Irish Naturalist Journal, 22(6), 244–246. Boero, F., Balduzzi, A., Bavestrello, G., Caffa, B. & Cattaneo-Vietti, R. (1986) Population dynamics of Eudendrium glomeratum (Cnidaria: Anthomedusae) on the Portofino Promontory (Ligurian Sea). Marine Biology, 92, 81–85. Calder, D.R. & Maÿal, E.M. (1998) Dry season distribution of hydroids in a small tropical estuary, Pernambuco, Brazil. Zoologische Verhandelingen, Leiden, 323, 69–78. Clarke, S.F. (1882) New and interesting hydroids from Chesapeake Bay. Memoirs of the Boston Society of Natural History, 3(4), 135–142.



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Correia, M.D. & Loyola e Silva, J. (1990) Caracterização das comunidades incrustantes e a fauna associada em painéis experimentais na Baía de Paranaguá, Paraná, Brasil. Publicações ACIESP, 71(3), 89–110. Costa, F.H.A. (1992) Some new data on description of Calliostoma pulchrum. La Conchiglia/The Shell, 264, 53–57. Eston, V.R., Migotto, A.E., Oliveira-Filho, E.C., Rodrigues, S.A. & Freitas, J.C. (1986) Vertical distribution of benthic marine organisms on rocky coasts of the Fernando de Noronha Archipelago (Brazil). Boletim do Instituto Oceanográfico, 34, 37–53. Grohmann, P., Souza, M.M. & Nogueira, C.C. (1997) Hydroids from the vicinity of a large industrial area in Vitória, Espírito Santo, Brazil. Proceedings of the VI International Conference of Coelenterate Biology, 227–232. Hirohito, Emperor Showa (1988) The hydroids of Sagami Bay. Tokio, Biological Laboratory – Imperial Household, 179 pp. Jäderholm, E. (1904) Mitteilungen über einige von der schwedischen Antarctic-Expedition 19011903 eingesammelte Hydroiden. Archives de Zoologie Expérimentale et Générale, 4(3), notes et revue, 1–14. Jäderholm, E. (1916). Hydroiden. In: Results of Dr. Mjöbergs Swedish scientific expeditions to Australia 1910-1913, XII. Kungliga svenska vetenskaps-akademiens handlingar, n. ser. 52(12), 1-9. Kubota, S. (1976) Notes of nematocysts of Japanese hydroids, I. Journal of the Faculty of Science of Hokkaido University, (6)20(2), 230–243. Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Holmiae, Laurenti Salvii, 823pp. Marinopoulos, J. (1992) Contribution à l’étude du genre Eudendrium (Hydrozoa: Hydroida) de la Méditerranée: taxonomie et phylogenie. Bulletin de l’Institut océanographique, Monaco, nº special 9, 53–66. Mariscal, R.N. (1974) Chapter 3. Nematocysts. In: Muscatine, L. & Lenhoff, H.M. (eds.), Coelenterate Biology. Academic Press, New York: p.129–178. Marques, A.C. (1993) Sistemática dos Eudendriidae L. Agassiz, 1862 (Cnidaria, Hydrozoa) do litoral paulista. M.Sc. Thesis, University of São Paulo. São Paulo, Brazil, 168pp. Marques, A.C. (1995) Eudendrium pocaruquarum n. sp. (Hydrozoa, Eudendriidae) from the southeastern coast of Brazil, with remarks on taxonomic approaches to the family Eudendriidae. Contributions to Zoology, 65(1), 35–40. Marques, A.C. (2001) O gênero Eudendrium (Hydrozoa, Anthomedusae, Eudendriidae) no Brasil. Papéis Avulsos de Zoologia, 41(22), 329–405. Marques, A.C. & Migotto, A.E. (1998) A new species of Eudendrium (Hydrozoa: Anthomedusae: Eudendriidae) from the Netherlands. Zoologische Verhandelingen, 323, 149–154. Marques, A.C. & Moretzsohn, F. (1995) Substrata used by Pteria colymbus (Röding, 1798) (Bivalvia: Pteriidae) for attachment. Publicação Especial do Instituto Oceanográfico, São Paulo, 11, 197–201. Marques, A.C., Peña-Cantero, A.L. & Vervoort, W. (2000a) Mediterranean species of Eudendrium Ehrenberg, 1834 (Hydrozoa, Anthomedusae, Eudendriidae) with the description of a new species. Journal of Zoology, 252, 197–213. Marques, A.C., Mergner, H., Höinghaus, R., Santos, C.M.D. & Vervoort, W. (2000b) Morphological study and taxonomical notes on Eudendriidae (Cnidaria, Hydrozoa: Athecatae/Anthomedusae). Zoologische Mededelingen, 74(5), 75–118. Masunari, S. (1983) Organismos do fital de Amphiroa beauvoissii Lamouroux, 1816 (Rhodophyta: Corallinaceae). I. Autoecologia. Boletim de Zoologia. Universidade de São Paulo, 7, 80–85.

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Mayal, E.M. (1973) Hidróides (Hydrozoa, Hydroida) de Pernambuco. M.Sc. Thesis, University of São Paulo. São Paulo, Brazil, 75 pp. Migotto, A.E. (1996) Benthic shallow-water hydroids (Cnidaria, Hydrozoa) of the coast of São Sebastião, Brazil, including a checklist of Brazilian hydroids. Zoologische Verhandelingen, 306, 1–125. Migotto, A.E., Marques , A.C. & Flynn, M.N. (2001) Seasonal recruitment of hydroids (Cnidaria) on experimental panels in the São Sebastião channel, southeastern Brazil. Bulletin of Marine Science, 68(2), 287–298. Migotto, A.E., Marques, A.C., Morandini, A.C. & Silveira, F.L. (2002) Checklist of the Cnidaria Medusozoa of Brazil. Biota Neotropica, 2, 1–31. Motz-Kossowska, S. (1905) Contribution à la coinnaissance des hydraires de la Méditerranée occidentale. I. Hydraire Gymnoblastiques. Archives de Zoologie Expérimentale et Générale, (4)3, 39–98. Nogueira, C.C., Grohmann, P. & Silva, V.M.A.P. (1997) Hydroids from the vicinity of a nuclear power plant site (CNAAA-Unidade I) at Angra dos Reis, Rio de Janeiro, southeastern Brazil. Proceedings of the VI International Conference of Coelenterate Biology, 365–369. Oliveira, O.M.P., Marques, A.C. & Migotto, A.E. (2000) Morphometric patterns of two fouling Eudendrium spp. (Hydrozoa, Anthomedusae, Eudendriidae) from São Sebastião (SP, SE Brazil). Brazilian Archives of Biology and Technology, 43(5), 519–526. Östman, C. (2000) A guideline to nematocyst nomenclature and classification, and some notes on the systematic value of nematocysts. Scientia Marina, 64 (suppl. 1), 31–46. Pallas, P.S. (1766) Elenchus zoophytorum sistens generum adumbrationes generaliores et specierum cognitarum succinctas descriptiones cum selectis auctorum synonymis. Hagae, Franciscum Varrentrapp, 451 pp. Picard, J. (1951) Note sur les hydraires littoraux de Banyuls-sur-Mer. Vie Milieu, 2, 338–349. Pires, D., Castro, C.B., Migotto, A.E. & Marques, A.C. (1992) Cnidários bentônicos do Arquipélago de Fernando de Noronha, Brasil. Boletim do Museu Nacional (Zoologia), 354, 1– 21. Rosa, C.N. (1973) Os animais de nossas praias. EDART, São Paulo (2th edition). 187pp. Rossi, L. (1961) Res Ligusticae CXX. Idroidi vivendi sulle scogliere del Promontorio di Portofino (Golfo di Genova). Annali del Museo Civico di Storia Naturale Giacomo Doria, 72, 68–85. Rosso, S. & Marques, A.C. (1997) Is there any conspicuous geographical pattern in intertidal hydrozoan distribution along the coast of São Paulo State, Southeastern Brazil? Proceedings of the VI International Conference of Coelenterate Biology, 415–422. Souza, M.M. (1987) Levantamento preliminar dos Hydroida (Cnidaria, Hydrozoa) do litoral do estado do Espírito Santo e considerações sobre sua biologia e ecologia. B.Sc. Thesis, Federal University of Rio de Janeiro. Rio de Janeiro, Brazil. 40pp. Thornely, L.R. (1904) Report on the Hydroida collected by Prof. Herdman, at Ceylon, in 1902. In: Herdman, W.A., Report to the Government of Ceylon on the pearl oyster fisheries of the Gulf of Manaar, by, Pt 2, suppl. Rep. 8, 107–126. Tommasi, L.R. (1970) Observações sobre a fauna bêntica do complexo estuarino-lagunar de Cananéia (SP). Boletim do Instituto Oceanográfico, São Paulo, 19, 43–56. Vannucci, M. (1954) Hydrozoa e Scyphozoa existentes no Instituto Oceanográfico. Boletim do Instituto Paulista de Oceanografia, 5(1–2), 95–150. Watson, J.E. (1985) The genus Eudendrium (Hydrozoa: Hydroida) from Australia. Proceedings of the Royal Society of Victoria, 97, 179–221. Watson, J.E. (1987) Records of Eudendrium (Hydrozoa: Hydroida) from New Zealand. Proceedings of the Linnean Society of New South Wales, 109(4), 325–330. Weill, R. (1934) Contribution à l´étude des cnidaires et de leurs nématocystes. I. Recherches sur les nématocystes (morphologie, physiologie, dévelopment). II. Le valeur taxonomique du cni-



11

ZOOTAXA

307

ZOOTAXA

307

dome. Travaux de la Station Zoologique de Wimereux, 11, 1–351. Yamada, M. (1954) Species of the genus Eudendrium from Japan. Publications of Akkeshi Marine Biological Station, 2, 1–19. Yamada, M. (1959) Hydroid fauna of Japanese and adjacent waters. Publications of Akkeshi Marine Biological Station, 9, 1–101.

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