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Accepted: 19 September 2003; published: 24 September 2003 1

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2003 Magnolia Press

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

Eudendrium caraiurusp. n. (Hydrozoa; Anthoathecata; Eudendri-idae) from the southeastern coast of Brazil

ANTONIO C. MARQUES1 & OTTO M.P. OLIVEIRA1,2

1Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11461, 05422-970, São Paulo, SP, Brazil; e-mail: marques@ib.usp.br2 Centro de Biologia Marinha, Universidade de São Paulo, Caixa Postal 83, 11600-970, São Sebastião, SP,Brazil; e-mail: ottompo@usp.br

Abstract

Eudendrium caraiurusp. n. is described for the southeastern coast of Brazil. The species wasreported previously from the area asE. 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 ofEudendriumhave been recorded to date from Brazilian waters:Euden-drium carneumClarke, 1882 (Vannucci 1954; Tommasi 1970; Rosa 1973; Masunari 1983;Souza 1987; Correia and Loyola e Silva 1990; Pireset al. 1992; Marques and Moretzsohn1995; Grohmannet al. 1997; Nogueiraet al. 1997; Rosso and Marques 1997; Calder andMaÿal, 1998; Marques 1993, 2001);Eudendrium capillareAlder, 1856 (Vannucci 1954;Mayal 1973; Alves and Mayal 1990; Marques 1993, 2001);Eudendrium rameum(Pallas,1766) (Mayal 1973);Eudendrium pocaruquarumMarques, 1995 (Marques 1995, 2001;Rosso and Marques 1997);Eudendrium? fragile Motz-Kossowska, 1905 (Grohmannetal. 1997);Eudendrium nambuccenseWatson, 1985 (Marques 1993, 2001; Nogueiraet al.1997);Eudendrium glomeratumPicard, 1951 (Marques 1993, 2001; Rosso and Marques1997; Oliveiraet al.2000);Eudendrium ramosum(Linnaeus, 1758) (Marques 1993, 2001;Grohmannet al. 1997; Rosso and Marques 1997);Eudendrium merulumWatson, 1985(Marques 1993, 2001);Eudendriumsp. (Estonet al. 1986; Costa, 1992; Nogueiraet al.

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307ZOOTAXA 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) includedata on the cnidome. Two out of the nine species (viz.,E. ? fragile from Vitória, EspíritoSanto State, andE. rameumfrom Itamaracá Island, Pernambuco State) were not includedin the partial review by Marques (2001) because no material was found in the collectionsstudied by him.Eudendrium pocaruquarumis a species so far known exclusively for Bra-zilian 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 bodyof hydranth, were described and assigned toEudendrium ramosumby Motz-Kossowska(1905). Based on this diagnostic character, Picard (1951) gave thenomen novum E. glom-eratumto refer to the species described by Motz-Kossowska (1905) and, as a by product ofthe new name, established the importance of the study of the cnidome in the systematics ofthe family Eudendriidae.Eudendrium glomeratumwas found to be a very common speciesin the Mediterranean Sea (see review in Marqueset al. 2000a), but it was also recordedfrom 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 asEudendrium glomeratum, a commonshallow water species, has been described by Marques (2001). A morphometrical analysisof the species revealed some inconsistencies with the populations ofE. glomeratumfromthe Mediterranean Sea (see Oliveiraet al. 2000). A more detailed analysis of populationsfrom both the Mediterranean and from Brazil have proved they belong to different lin-eages. Therefore, the purpose of this study is to describe Brazilian material asEudendriumcaraiuru sp. n.

Material and methods

Colonies ofEudendrium caraiurusp. n. were collected by hand on rock and artificial sub-strates in the shallow waters along the coast of the states of Rio de Janeiro (one site) andSão Paulo (many sites). The specimens were anaesthetized in a 1:1 solution of 7.5% mag-nesium chloride solution and seawater, and preserved in 4% formaldehyde solution in sea-water or alcohol 70%. The specimens were examined, measured, and drawn undermicroscope and stereomicroscope, both with camara lucida and photo devices. The cni-dome terminology follows Weill (1934) and Mariscal (1974), and measures of nemato-cysts 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 col-oration of the specimens, distribution and number of nematocyst capsules, and morphol-ogy of discharged nematocysts. Stems for scanning electron microscopy (SEM) were

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307ZOOTAXApreserved 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 methodsfor 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 daUniversidade de São Paulo (MZUSP), Royal Ontario Museum (ROMIZ), and MuseuNacional da Universidade Federal do Rio de Janeiro (MNRJ).

Taxonomic part

Eudendrium caraiurusp. 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; Oliveiraet al., 2000: 519–525; Migottoet al., 2001: 289,294–296; 2002: 11.

nonEudendrium glomeratumPicard, 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 metallicstructures, 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, onrock, intertidal, col. A.C. Marques (MZUSP 0428; former ACM-SP121); MarandubaBeach: colony without gonophores, 30.vii.1992, formol, on rock, intertidal, col. A.C.Marques (MZUSP 0429; former ACM-SP123);São Sebastião: Pier Sul (Petrobrás): malecolony, 18.vii.1990, formol, onPerna 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, for-mol, on Schizoporella, 3m, col. A.E. Migotto (MZUSP 0459; former ACM-SP161); Jar-obá 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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307ZOOTAXA 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 ACM-SP049); 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 col-ony, 15.ix.1990, formol, on rope, 2m, col. A.C. Marques (MZUSP 0393; former ACM-SP055); 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, onropes, 1m, col. O.M.P. Oliveira (MZUSP 0370); Barequeçaba Point (23º49,979´S45º25,843´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 metallicstructures, 7m, col. H.K. Boscolo (MZUSP 0372);Cananéia: Cardoso Island, costão doPereirinha: 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 hydranthbody, spadix of female gonophores, and in a whorl of 16–25 around hypostome. Femaleblastostyles with reduced hypostome and tentacles.

Etymology. From the Tupi native language “cáraiurú” (= powerful mouth), in refer-ence to the presence of large euryteles on the hypostome.

Description. Colonies dioecious, arborescent, up to 54 mm in height, main stemsunfascicled. Hydrocauli arising from creeping hydrorhiza; branches many, more or lessalternate, occurring over entire hydrocaulus, branches up to third order, in radiate planes orrarely more or less planar; pedicels arising from main stem or branches of first, second orthird order. Perisarc of main stem strongly developed, dark brown, single tubes 0.40 mm indiameter, 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. 0.10mm in diameter.

Hydranths 0.18–0.75 mm in height, 0.18–0.57 mm in diameter (measured in the bodyregion just below the tentacles), orange in color, with a distinct groove in the aboralregion; tentacles 23–34 in number, occurring in a whorl below hypostome. Somehydranths with reduced tentacles juxtaposed in two close whorls.

Gonophores styloids, arising from body of hydranth. Immature styloids placed in a cir-cle around body of hydranth. Male blastostyle orange, with 10–29 sporosacs, each sporo-sac 1–2 chambered, linked to blastostyle body by a stalk, with a very distinct spadix overits longitudinal axis, and a terminal tubercle on its apex; distal chamber 0.12–0.18 mm indiameter. Male blastostyles completely reduced over earlier stages of their developmentwith pedicels corrugated throughout. Female gonophores orange, arising on partially

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307ZOOTAXAreduced blastostyles with highly atrophied hypostome and degenerated tentacles. Imma-

ture eggs having a simple and curved spadix over a single egg. Blastostyle reducing com-pletely during development or at maximum with 1–5 stumps (tentacles), and spadicesshed. Mature oval eggs thickened by perisarc and linked directly to the wrinkled pedicelby 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 heterotrichousmacrobasic (or mesobasic) euryteles.

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

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FIGURES 9–15.Eudendrium caraiurusp. 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 microbasiceuryteles. 13, primary polyp, derived from larval metamorphosis. 14, young hydranth encapsulatedby 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); 200µm (10); 300µm (11,13–15); 10µm (12).

Small microbasic euryteles (seen discharged), 6.1–8.0 X 2.9–3.9µm, 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 60µm in length , 3.0–3.6 times lengthof capsule; undischarged shaft in 1.5 coils inside capsule; distributed on hydranth bodysometimes 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 (Oliveiraet al., 2000; Marques, 2001), Cananéia(Marques, 2001).

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

Discussion

The species nameEudendrium glomeratumwas proposed as anomen novumby Picard(1951) for a specimen described asEudendrium ramosum(Linnaeus, 1758) by Motz-Kos-sowska (1905). Picard justified the erection of a new species based on the presence ofmacrobasic euryteles arranged in warts on the body of hydranth, but lacking on the hypos-tome (cf. Motz-Kossowska 1905; Picard 1951; Rossi 1961; Boero and Cornelius 1987;Marqueset al. 2000a). The diagnosis of the species was amended two times. Initially, Wat-son (1985) re-defined the species to include specimens from the Mediterranean Sea inwhich the nematocysts formed a continuous ring around the body of hydranth. Later, itwas amended again to include Brazilian specimens in which the nematocysts exhibited anumber of differences from European material: shaft intermediate between macrobasicand microbasic forms, and a scarcity of these nematocysts on the hypostome, where theyare never organized in warts (Marques 1993, 2001).

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307ZOOTAXA The plasticity of the so calledE. glomeratumwas remarked by Marqueset al. (2000a).

These authors also included comparisons of the species withEudendrium magnificumYamada, 1954, and considered the possibility that the two might be conspecific. Theypointed 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 theselarge nematocysts). The muddled state of the taxonomy ofE. glomeratumwas also high-lighted by Marqueset al. (2000b). They compared the sexual female features of the spe-cies with those ofEudendrium cyathiferumJäderholm, 1904, and noted the eventualbroader distribution ofE. glomeratumdue to misidentification of the species particularlycompared withE. rameumand E. ramosum. The studies above clearly pointed out theexistence of a complex of species under the nameE. glomeratum.

Eudendrium caraiurusp. n. has been reported so far from the Brazilian coast asEudendrium glomeratumPicard, 1951. In fact, the two species share some characters here-tofore considered diagnostic ofE. glomeratum, such as the presence of large eurytelesarranged in warts around the body of hydranth and on the immature unbranched femalespadix, and the mature eggs encapsulated by perisarc scattered along the pedicel of com-pletely reduced blastostyles, linked to those pedicels by short concave perisarc bases (cf.Marqueset al.2000a, 2000b).

However, detailed morphological comparisons betweenE. caraiurusp. n. ( “E. glom-eratum” of Brazilian authors) and Mediterranean populations ofE. glomeratumhaveshown that the height and width of Mediterranean specimens wereca. ten times those ofthe Brazilian specimens, and that the Brazilian colonies are unfascicled (Oliveiraet al.2000). Moreover, the cnidome of both species is also slightly different.Eudendrium car-aiuru sp. n. has euryteles with smaller dimensions (18.7–22.7 X 7.1–9.3 µm, see alsoMarques 2001) than those ofE. glomeratumfrom the Mediterranean Sea (22.2–27.2 X9.5–12.0 µm, Marqueset al. 2000a; 24–28 X 10–11 µm, Marinopoulos 1992) and similarto 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. caraiurusp. n. has microbasic euryteles insteadof macrobasic euryteles. However, under Östman’s (2000) definition, the nematocyst is amesobasic eurytele.

The study of living material ofE. caraiurusp. n. corroborated the previous hypothesisby 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 thespecies, taking about 8–36 hours to be completed and, during which, it is possible to seethe presence of the two whorls of tentacles in resorbing hydranths. The feature is alsopresent in certain other species ofEudendrium(cf. Marqueset al. 2000b).

There are also some striking differences betweenE. caraiurusp. n. andE. glomeratumecologically. After seasonal studies during 1999–2000, we found Brazilian populations ofE. caraiurusp. n. with no resting stages, living in temperatures over 21.5° C (Oliveira andMarques, unpublished data) and, during the last decade, the species was always found in

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307ZOOTAXAtemperatures of 19.5–27.7 ºC and salinities of 30.0–35.4 ppt (Marques 1993, and personal

observations). Meanwhile, populations ofE. glomeratumfrom the Mediterranean Seahave 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 neverover 21,5 ºC. Moreover, reproductive patterns are also different between the two popula-tions (cf. Marques 1993 and Boeroet al.1986; Oliveira and Marques, unpublished data).

All differences between the populations, when put together, are strong evidence thatE.caraiuru sp.n. represents a different lineage fromE. glomeratum. Still, evidence suggeststhat the remaining populations ofE. glomeratum, from many places, may be cryptic spe-cies, needing detailed morphological studies (including morphometry, SEM), molecularanalysis and assessment of ecological data.

Acknowledgements

The authors are grateful to Alvaro E. Migotto for his help with the illustrations and forreviewing 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 Bio-logia Marinha of the Universidade de São Paulo, in which part of the study was made, forlogistic 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).

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