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Based on live specimens of Nemalecium lighti collected in the coasts of La Réunion (Indian Ocean) and São Sebastião (Southeastern coast of Brazil) and kept in the laboratory, we observed the release of short-lived medusoids. The gonangia pass through six developmental phases: growing, ripening, migrating, stripping, liberating and spawning. The medusoids are tall, lack tentacles, bulbs, circular and radial canals, and the sexual products are packed around the eccentric manubrium; they are provided with a velum and with a subumbrellar ectoderm rich in transverse striated muscle fibers. There are refringent and isotropic corpuscles within vacuolated and ciliated large cells located around the aperture of the medusoid, which possibly function as statoliths. The corpuscles are similar to those already described for the families Plumulariidae and Aglaopheniidae. The gametes are liberated shortly after the release of the medusoid from the gonotheca. The female medusoid spawned 40-62 ova; spermatozoa exhibited a semicircular nucleus, and planulae were formed c. twelve hours after fertilization. Colonies with medusoids of only one sex or with both male and female medusoids., No disponible

Beroe forskalii Milne Edwards, 1841 (Figures 1���2; Table 1) Beroe forskalii Milne Edwards, 1841 ��� Chun 1880; Tamm & Tamm 1991, 1993; Haddock & Case 1999; Mianzan 1999; Wrobel & Mills 2003; Lindsay & Hunt 2005; Lindsay 2006; Shiganova & Malej 2009. Material examined. MZUSP 00012, a single specimen 2.5 cm long from Canal de S��o Sebasti��o, 23.8�� S, 45.4�� W, coll. A. Migotto 8 July 2010, reared to 10 July 2010 then fixed in 4% formalin. Description. Body sac-shaped, pinkish, with broad mouth; body width tapering from middle to aboral extremity of body (Fig. 1A); compressed along tentacular plane (Fig. 1C). Eight meridional canals extending from aboral region towards mouth and connected with a circular canal around mouth by a mesh of anastomosing canals; these with lateral diverticulae, mostly with blind ends in aboral half of body, but forming anastomoses along oral half (Fig. 1F). Diverticulae from subtentacular meridional canals connected with paragastric canal (Fig. 1E). Ctene rows located above meridional canals arising at aboral region. Substomodeal ctene rows extending for about 3/4 the length of the meridional canals to oral region. Subtentacular ctene rows limited to aboral 2/3 of body. Mouth wide, semicircular, occupying whole oral region, opening into large pharynx that occupies most of central inner part of animal. Two opposite paragastric canals, extending from aboral pole of pharynx to circular canal around mouth, crossing at tentacular plane. Aboral extremity of body mitriform (Fig. 1D). Apical organ including a statolith at center of polar fields. Aboral papillae prominent, branched (Fig. 1G), projecting from margins of polar fields, forming an ���hourglass��� figure, oriented in stomodeal plane when in aboral view (Fig. 1B). Each papilla with first- and sometimes second-order branches in a pinnate arrangement; branches curving toward exterior of polar field. The animal is a simultaneous hermaphrodite, with gonads formed at lateral walls of meridional canals and nearby diverticulae (Fig. 1E). Ovaries restricted to portion of meridional canals under ctene rows. Testes lying along all meridional canals and adjacent diverticula. Gonads of adjacent meridional canals facing those of same gender on the adjacent canal, i.e. female gonads face in stomodeal-stomodeal and tentacle-tentacle patterns; male gonads face in a stomodeal-tentacular arrangement. Macrocilia are robust (length 5���7 times width), with about 20 small teeth along edge (Fig. 2), covering extensive area of oral portion of stomodeal cavity, arranged in long tapering stripes that run from lip toward center of stomodeum. Remarks. The young individual on which the above description is based, was very active for the three days it was kept in the aquarium, swimming quickly, sometimes with the mouth open. It resembled Beroe mitrata Moser, 1907, except for the presence of anastomoses and the pointed end of the aboral portion (Wrobel & Mills, 2003). Based on the shape of their macrocilia, these two species form a distinct group within the genus Beroe (Tamm & Tamm 1993). In these two species, the macrocilia are very similar, differing only by their pattern of distribution along the stomodeal cavity, comprising a carpet covering all of the oral portion of the stomodeum in B. mitrata and long tapering stripes that run from the lip toward the center of the stomodeum in B. forskalii (Tamm & Tamm 1993). On this basis, inter alia, the specimen from S��o Sebasti��o Channel was certainly B. forskalii. The first description of what was probably this species was made by Forsk��l (1775), as Beroe rufescens. This name was not used by later researchers after Milne Edwards (1841) questioned its validity. The combination Beroe forskalii has been used in at least 20 publications in the last 50 years and, following the nomenclatural Principle of Stability (ICZN 1999), its use may continue. The author of the combination Beroe forskalli is taken as Milne Edwards (1841), not Chun (1880), following the Principle of Priority (ICZN 1999). Distribution. Beroe forskalii is distributed worldwide (e.g. Moser 1910; Mianzan 1999; Lindsay & Hunt 2005; Shiganova & Malej 2009; Oliveira et al. submitted)., Published as part of Oliveira, Otto M. P. & Migotto, Alvaro E., 2014, First occurrence of Beroe forskalii (Ctenophora) in South American Atlantic coastal waters, with notes on the use of macrociliary patterns for beroid identification, pp. 470-476 in Zootaxa 3779 (4) on pages 471-473, DOI: 10.11646/zootaxa.3779.4.5, http://zenodo.org/record/4910453, {"references":["Milne Edwards, M. H. (1841) Observations sur la structure et les fonctions de quelques Zoophytes, Mollusques et Crustaces des cotes de la France. Annales des Sciences Naturelles, ser. 2, 16, 193 - 232.","Chun, C. (1880) Die Ctenophoren des Golfes von Neapel und der angrenzenden Meeres-abschnitte. Eine Monographie. Fauna and flora des Golfes von Neapel, 1, Monographie, 18, 1 - 313, 18 pls.","Tamm, S. L. & Tamm, S. (1991) Reversible epithelial adhesion closes the mouth of Beroe, a carnivorous marine jelly. Biological Bulletin, 181, 463 - 474.","Tamm, S. L. & Tamm, S. (1993) Diversity of macrociliary size, tooth patterns, and distribution in Beroe (Ctenophora). Zoomorphology, 113, 79 - 89. http: // dx. doi. org / 10.1007 / bf 00403086","Haddock, S. H. D. & Case, J. F. (1999) Bioluminescence spectra of shallow and deep-sea gelatinous zooplankton: ctenophores, medusae and siphonophores. Marine Biology, 133, 571 - 582. http: // dx. doi. org / 10.1007 / s 002270050497","Mianzan, H. W. (1999) Ctenophora. In: Boltovskoy, D. (Ed.), South Atlantic Zooplankton. Backhuys Publishers, Leiden, pp. 561 - 573.","Wrobel, D. & Mills, C. (2003) Pacific Coast Pelagic Invertebrates - A Guide to the Common Gelatinous Animals. Second Printing with corrections. Sea Challengers and the Monterey Bay Aquarium, Monterey, California, iv + 108 pp.","Lindsay, D. J. & Hunt, J. C. (2005) Biodiversity in midwater cnidarians and ctenophores: submersible based results from deep- water bays in the Japan Sea and north-western Pacific. Journal of the Marine Biological Association of the United Kingdom, 85, 503 - 517. http: // dx. doi. org / 10.1017 / s 0025315405011434","Lindsay, D. J. (2006) A checklist of midwater cnidarians and ctenophores from Sagami Bay - species sampled during submersible surveys from 1993 - 2004. Bulletin of the Plankton Society of Japan, 53, 104 - 110.","Shiganova, T. & Malej, A. (2009) Native and non-native ctenophores in the Gulf of Trieste, Northern Adriatic Sea. Journal of Plankton Research, 31, 61 - 71. http: // dx. doi. org / 10.1093 / plankt / fbn 102","Forskal, P. (1775) Descriptiones Animalium, Avium, Amphibiorum, Piscium, Insectorum, Vermium; quae in itinere orientali observavit Petrus Forskal. Hauniae, ex officina Molleri, 20 + xxxiv + 164 pp., 1 map.","ICZN (1999) International Code of Zoological Nomenclature, 4 th edn. ITZN, London, 306 pp.","Moser, F. (1910) Die Ctenophoren der Deutschen Sudpolar-Expedition 1901 - 1903. Deutsche Sudpolar-Expedition, 11, Zoologie, 117 - 192."]}

Filellum contortum (Nutting, 1905) (Figure 6 A���D) Lafoea contorta Nutting, 1905: 945, pl. 3, fig. 6, pl. 9, figs. 8���9. ? Filellum contortum: Pe��a Cantero et al. 1998: 301. Non Filellum contortum: Pe��a Cantero and Gili 2006: 766 [= Filellum bouvetensis sp. nov.]. Type series. Lectotype��� Lafoea contorta Nutting, 1905; alcohol specimen on Aglaopheniidae (USNM 70715; specimen represented by Nutting 1905, pl. 3, fig. 6, pl. 9, figs. 8���9). Paralectotype��� Lafoea contorta Nutting 1905; alcohol specimen (USNM 68984). Type locality. From the lectotype, ���Albatross R/V��� Expedition, Hawaiian Explorations, Stn. 3949, north Pacific Ocean, Hawaii, north of Laysan Island, 27 o 47 ��� 45 ������N; 171 o 52 ��� 35 ������W, 108���238 m, 21 May 1902, leg. Albatross R/V. Paralectotype: ���Albatross R/V��� Expedition, Hawaiian Explorations, Stn. 4102, north Pacific Ocean, Hawaii, Pailolo Channel, between Maui and Molokai Islands, 21 o03��� 10 ������N; 156 o 45 ��� 20 ������W, 223���241 m, 23 September 1902, leg. Albatross R/V. Material examined. Lectotype USNM 70715 and paralectotype USNM 68984. Description of lectotype. Colony stolonal, with a filiform and creeping hydrorhiza growing over some 7 mm at base of an aglaopheniid hydroid; hydrorhizal tubes 0.10���0.14 mm wide, with few anastomosed perisarc. Hydrothecae sessile, arising with no definite pattern from hydrorhiza, less than half of their total length partially adnate to substratum parallel to hydrorhizal stolon, free part emerging from substrate plane at angles wide than 60 o, usually 90 o. Adnate portion of hydrothecae tubular or maximally slightly flattened, 0.41���0.73 mm (0.58 �� 0.11, n= 10) long, perisarc with many striae on abaxial hydrothecal surface of adnate part, striae extend up to curvature of hydrothecae; free part cylindrical, smooth, 0.84���1.40 mm (1.22 �� 0.15, n= 10) long; perisarc moderately thin; hydrothecae slightly widening distally, margin even and smooth, with up to two renovations, tenuous flaring; hydrothecal orifice circular, 0.21���0.27 mm (0.24 �� 0.02, n= 10) wide, perpendicular to axis of free part of hydrotheca. Hydranths badly preserved, with ca. 14 tentacles, hypostome conical. Hydranths, when retracted, lying at the adnate space of the hydrothecae. Gonothecae absent. Nematocysts of one category, heterotrichous microbasic?mastigophores (not seen discharged), 5.0���6.0 X 2.0��� 2.5 ��m (5.60 �� 0.39 X 2.35 �� 0.24, n = 10), rice-shaped, common. Distribution. Filellum contortum was recorded for the coast of Japan (Stechow 1913 b). Remarks. Filellum contortum was originally described as Lafoea contorta Nutting, 1905, and diagnosed as a ���colony parasitic, growing from a twisted rootstock��� with hydrothecae ���sessile, tubular, very long, and often bent and twisted in various ways���. The specimens described by Nutting (1905: 945) were sterile. This species is different from most of the species of Filellum in its morphometry (e.g., larger dimension of the free part of hydrotheca). The cnidome of F. contortum is also different, comprising a single type of nematocyst (heterotrichous microbasic mastigophore), comparatively smaller than those found in other species. Although these differences could characterize the species, they may be variable or dependent on the quality of specimen preservation. Besides, Nutting���s (1905) material lacks coppinia, so the crucial information concerning the reproductive structure, necessary to properly assign new material to this species, is unknown. The only coppiniae assigned to F. contortum was described by Pe��a Cantero and Gili (2006) from Bouvet Island, herein redescribed as Filellum bouvetensis sp. nov. Therefore, a reliable diagnosis of the species is difficult, and its validity remains dubious (Pe��a Cantero et al. 1998) until its reproductive parts are described. We consider F. contortum as a species inquirenda., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 22-24, DOI: 10.5281/zenodo.206783, {"references":["Nutting, C. C. (1905) Hydroids of the Hawaiian Islands collected by the steamer Albatross in 1902. Bulletin of the United States Fish Commission, 23, 931 - 959.","Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315.","Pena Cantero, A. L. & Gili, J. M. (2006) Benthic hydroids (Cnidaria, Hydrozoa) from off Bouvet Island (Antarctic Ocean). Polar Biology, 29, 764 - 771.","Stechow, E. (1913 b) Hydroidpolypen der japanische Ostkuste. II. Teil: Campanularidae, Halecidae, Campanulinidae und Sertularidae, nebst Erganzungen zu den Athecata und Plumularidae. Abhandlungen der Mathematisch-Physikalischen Klasse der Koniglich Bayerischen Akademie der Wissenschaften, 3, 1 - 162."]}

Key for the identification of the fertile specimens of Filellum 1 Abcauline wall of the adnate part of the hydrotheca with transverse striae........................................ 2 - Abcauline wall of the adnate part of the hydrotheca without transverse striae....................................... 7 2 Defensive tubes of the coppinia forming a protective structure over the gonothecae................................. 3 - Defensive tubes of the coppinia not forming a protective structure over the gonothecae........... F. nitidum Watson, 2005 3 Defensive tubes arising amongst the gonothecae. Gonotheca with short distal neck with everted rim.................... 4 - Defensive tubes exclusively on the periphery of the gonothecal mass and arching over the gonothecae................. 5 4 Defensive tubes forming a shield-shaped structure (canopy)............................... F. serratum (Clarke, 1879) - Defensive tubes simply arching over gonothecae............................. F. magnificum Pe��a Cantero et al., 2004 b 5 Gonothecae without distal neck................................................. F. antarcticum (Hartlaub, 1904) - Gonothecae with distal neck............................................................................. 6 6 Coppinia bud-shaped. Gonothecae flask-shaped. Hydrothecal aperture 140���168 ��m. Hydrothecal free part 320���420 ��m.................................................................................... F. conopeum Watson, 2003 - Coppinia corbula-shaped. Gonothecae irregular. Adnate hydrothecal part short or absent. Hydrothecal aperture 180���250 ��m. Hydrothecal free part 650���1875 ��m..................................................... F. bouvetensis sp. nov. 7 Gonothecae without juxtaposed walls. Defensive tubes absent................ F. disaggregatum Pe��a Cantero et al., 1998 - Gonothecae with juxtaposed walls. Defensive tubes present.................................................... 8 8 Hydrothecae and hydrorhiza forming a continuous layer on the substrate.......................................... 9 - Hydrothecae and hydrorhiza spaced and distinct from each other............................ F. serpens (Hassall, 1848) 9 Gonothecae fusiform............................................................. F. adhaerens (Nutting, 1901) - Gonothecae polygonal....................................................... F. parasiticum (Antsulevich, 1987), Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on page 24, DOI: 10.5281/zenodo.206783, {"references":["Watson, J. E. (2005) Hydroids of the Archipelago of the Recherche and Esperance, Western Australia: Annotated list, redescription of species and description of new species. In: Weills, F. E., Walker, D. I. & Kendrick, G. A. (Eds), The marine flora and fauna of Esperance, Western Australia. Western Australian Museum, Perth, pp. 495 - 612.","Clarke, S. F. (1879) Report on the Hydroida collected during the Exploration of the Gulf Stream and Gulf of Mexico by Alexan- der Agassiz, 1877 - 78. Bulletin of the Museum of Comparative Zoology at Harvard College, in Cambridge, 5, 239 - 252.","Pena Cantero, A. L., Svoboda, A. & Vervoort, W. (2004 b) Antarctic hydroids (Cnidaria, Hydrozoa) of the families Campanulinidae, Lafoeidae and Campanulariidae from recent Antarctic expeditions with R. V. Polarstern, with the description of a new species. Journal of Natural History, 38, 2269 - 2303.","Hartlaub, C. (1904) Hydroiden. Resultats du Voyage du S. Y. Belgica. Zoologie, 7, 1 - 19.","Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315.","Hassall, A. H. (1848) Definitions of three new British zoophytes. Zoologist, 6, 2223.","Nutting, C. C. (1901) Papers from the Harriman Alaska Expedition. XXI. The hydroids. Proceedings of the Washington Academy of Sciences, 3, 157 - 216.","Antsulevich, A. E. (1987) Gidroidy shel'fa Kuril'skykh ostrovov (Hydroids from the shelf waters of Kurile Islands). Zoologicheskii Institut, Akademiya Nauk SSSR, Leningrad, 165 pp."]}

Genus Filellum Hincks, 1868 Capsularia: Gray 1848: 151 (in part); Naumov 1960: 280; Naumov 1969: 303 [non Capsularia Cuvier, 1798: 665 = Coryne Gaertner, in Pallas, 1774; after Lamouroux et al. 1824: 224; Bedot 1901: 434, 437; Cornelius 1975: 378]. ? Conchella Gray, 1848: 88 [discussion in Cornelius & Calder 1986: 338; Cornelius 1995: 253; Calder 1991: 35]. Coppinia Hassall, 1848: 2223 [discussion in Cornelius & Calder 1986: 335���337; Calder 1991: 35]. Reticularia Thomson, 1853: 443; Ralph 1958: 311; Rees and Thursfield 1965: 85 ���88 (in part) [preoccupied by a name for a brachiopod described by McCoy, in Griffith 1844; after Vervoort 1972: 50; Cornelius 1975: 378; Calder 1991: 35 ��� Stechow 1923: 144 considered Reticularia as a mollusk]. Halia Hincks, 1855: 128 [preoccupied by a name for a gastropod described by Risso 1826; after Stechow 1923: 144 (quoting Scudder 1882, Nomenclator Zoologicus: 152); Calder 1991: 35; Cornelius 1995: 253]. Thalia L. Agassiz, 1862: 355 [incorrect subsequent spelling of Halia Hincks, 1855, after Bedot 1910: 308]. Filellum Hincks, 1868: 214; Nutting 1901: 179; 1905: 946; Stechow 1913 b: 44; Stechow 1923: 137 (comparative table of lafoeid genera), 144���145; Naumov 1960: 280; Naumov 1969: 303; Vervoort 1972: 50; Cornelius 1975: 378; Millard 1975: 175; Cornelius and Calder 1986: 335 ���339; Calder 1991: 35 ���36; Cornelius 1995: 253; Hirohito 1995: 110; Pe��a Cantero et al. 1998: 298 ���299; Vervoort and Watson 2003: 57. Fillelum Billard, 1906: 166 [incorrect subsequent spelling]. Grammaria: Vervoort 1946: 194 (in part) cf. Hirohito 1995: 110 [not Grammaria Stimpson, 1854]. Corystolona Watson, 2002: 333 ���334. Type species. Campanularia serpens Hassall, 1848, by monotypy in Hincks (1868: 214). Description. Lafoeidae with young and adult colonies stolonal; hydrorhiza filiform, branched, creeping on substratum. Hydrorhizal stolons giving rise directly to hydrothecae. Hydrothecae tubular, sessile; basal portion adherent to substratum, usually parallel to hydrorhizal stolon; distal free portion directed upwards at varying degrees and extension. Hydranth long, one distal whorl of filiform tentacles; hypostome short, dome-shaped. Hydrothecal diaphragm, hydrothecal operculum, nematothecae and nematophores absent. Gonophores as fixed sporosacs, in gonothecae forming coppiniae. Gonothecae either closely set, with lateral walls juxtaposed each other, or weakly aggregated, gonothecae being isolated. Coppinia with or without defensive tubes (modified hydrothecae). Cnidome consisting basically of heterotrichous microbasic mastigophores and/or microbasic euryteles. Distribution. Based on literature, some species of Filellum are distributed worldwide, usually epibiotic on other hydroids (Cornelius & Calder 1986: 335; Cornelius 1995: 253). Remarks. Several generic names were proposed prior to Filellum Hincks, 1868, but they were dismissed either by ICZN decision or because they were preoccupied (ICZN 1999, art. 60). The nomenclatural history of the group may be found in Stechow (1923: 144���145), Vervoort (1972: 50), Cornelius (1975: 378), Cornelius and Calder (1986: 335���341), Calder (1991: 35���36), and Pe��a Cantero et al. (1998: 299). A summary of the nomenclatural issues is given below. Conchella Gray, 1848 and Coppinia Hassall, 1848 are two generic names used in specific combinations of taxa presently referred to the generic name Filellum. The identification of the type species of Conchella, Campanularia intertexta Couch, 1844, whose type specimen is not existent (see Couch, 1844: 41���42), is doubtful. The species may refer to Filellum serpens (Hassall, 1848), Lafoea dumosa (Fleming, 1820), Orthopyxis integra (Macgillivray, 1842), or even to all of them (Cornelius & Calder 1986: 338). The last of these three was excluded as type of C. intertexa by designation as lectotype of an illustration by Couch of a specimen having coppiniae (Cornelius 1982: 122). The generic name Conchella was never used subsequently (see Cornelius & Calder 1986). Conversely, the generic name Coppinia Hassall, 1848 was based on a lafoeid reproductive structure that had been confused as a distinct taxon, presently known in the literature as ���coppinia���. The type species of the genus, Coppinia mirabilis Hassall, 1848, was eventually found to be the widely known species Filellum serpens (Hassall, 1848), as it is considered today. The generic name Coppinia, although rarely used during the 20 th century, remained valid and threatened the more widely used name Filellum (Cornelius & Calder 1986: 335���336). A case was submitted to the ICZN requesting suppression, for the purposes of the Principle of Priority, of the obscure and infrequently used names Conchella Gray, 1848 and Coppinia Hassall, 1848. In Opinion 1485 (ICZN 1988), both names were placed on the Official Index of Rejected and Invalid Generic Names in Zoology. Campanularia serpens Hassall, 1848 was transferred to the genus Reticularia Thomson, 1853 by Totton (1930: 160���161), based on its hypothesized conspecificity with the type species of the genus, Reticularia immersa Thomson, 1853. The generic name Reticularia has been used by a series of authors in combination with many specific names presently referred to Filellum, such as Reticularia annulata Watson, 1973 (Watson 1973: 164); Reticularia antarctica (Hartlaub, 1904) (Totton 1930: 160 ���161; Briggs 1938: 26; Rees & Thursfield 1965: 85 ���86; Watson 1973: 163 ���164); Reticularia serpens (Hassall, 1848) (Hastings 1943: 394; Millard 1957: 203; Rees & Thursfield 1965: 87 ���88); and Reticularia serrata (Clarke, 1879) (Ralph 1958: 312; Rees & Thursfield 1965: 86 ���87). Hincks (1868) believed that the generic name Reticularia Thomson, 1853 was preoccupied by Reticularia Bulliard, 1791, a name applied to a fungus, and founded Filellum as a replacement for it. Bulliard���s usage of the name is excluded from, and does not become a homonym in, zoological nomenclature (Vervoort 1972: 49; Calder 1991: 35; see ICZN 1999, art. 1.4). However, the name is preoccupied by Reticularia McCoy, in Griffith 1844, a fossil brachiopod. As the oldest available and valid synonym of Reticularia Thomson, 1853, Filellum becomes the name of the genus. The genus name Halia Hincks, 1855, and the binomen Halia praetenuis Hincks, 1855, were applied to what was originally thought to have been a bryozoan. The species was transferred to Hydrozoa by Hincks (1856), who held Halia to be a congener of Reticularia Thomson, 1853. In fact, the name Halia Hincks, 1855 is an invalid junior homonym of the gastropod genus Halia Risso, 1826 (see Calder 1991: 35). The generic name Thalia, as used by L. Agassiz (1862: 355), is a misspelling of Halia; as such it does not constitute a threat to Filellum. Within the family Lafoeidae, Filellum was previously considered to be related to Grammaria, either as: (1) the subgenus of Grammaria (Filellum) (e.g., Broch 1913, 1928a, 1928 b, 1933; Leloup 1933; Berezina 1948); (2) the genus Filellum (Grammaria) (e.g., Leloup 1935); or (3) directly as a junior synonym (e.g., Kramp 1913; Broch 1918; Mathisen 1928; Remane 1933; Leloup 1938, 1947; Vervoort 1942, 1946, 1959; da Cunha 1950; Arndt 1964; Fey 1970; F��ller 1970; Castric-Fey 1973). Evidence against this relationship was given by Vervoort (1972: 50), and re-affirmed by Calder (1991: 35). Similarly, the genus Filellum has also incorrectly been considered as including the subgenus Lafoea, such as in the use of Filellum (Lafoea) adhaerens (Stechow 1913 b: 30). Filellum was considered a junior synonym of Capsularia Modeer, 1793 (Cornelius 1975: 378). The generic name was used by Naumov (1960: 280; 1969: 303), probably following Gray (1848: 151, as in the binomen Capsularia serpens). Capsularia Cuvier, 1798 was considered a synonym of Coryne Gaertner, in Pallas 1774 by Lamouroux et al. (1824: 220, 224), Bedot (1901: 437), Cornelius (1975: 378, who provided the historical account on the case), and Calder (1991: 35���36). Although the genus Filellum is herein described as lacking nematothecae and nematophores, Calder (1991: 35) described the genus with ���nematothecae usually absent���. As far as we know, based on all specimens of Filellum we have studied, there is no trace of nematothecae in the members of the genus. As stated above, most authors have based their identifications and descriptions exclusively on characters of the trophosome. Nevertheless, the paramount taxonomical importance of the reproductive structures (coppiniae) is evident, as illustrated by F. disaggregatum and F. serpens (see Pe��a Cantero et al. 1998), species with indistinguishable trophosome, but easily recognized by their coppinia. As already stated, this fact makes unreliable all previous identifications of species of Filellum exclusively based on non-reproductive characters (Pe��a Cantero et al. 1998: 297). The following species were described as belonging to, or subsequently referred to, the genus Filellum. Nevertheless, to our knowledge, only the species listed in bold belong to Filellum. The remaining species belong to other genera of Lafoeidae (Grammaria, Lafoea) or to other phyla. Filellum adnatum (Fraser, 1925) species inquirenda (from Lafoea adnata Fraser, 1925) Filellum adhaerens (Nutting, 1901) (from Lafoea adhaerens Nutting, 1901) Filellum annulatum (Watson, 1973) Filellum antarcticum (Hartlaub, 1904) (from Lafoea antarctica Hartlaub, 1904) Filellum bouvieri Jullien, 1880 species inquirenda (from Filellum Bouvieri Jullien, 1880) Filellum conopeum Watson, 2003 Filellum contortum (Nutting, 1905) species inquirenda (from Lafoea contorta Nutting, 1905) Filellum disaggregatum Pe��a Cantero et al., 1998 Filellum expansum Levinsen, 1893 = Folliculina gigantea (Dons, 1910) (Ciliophora), after Dons (1910) and Vannucci (1955) Filellum gabriellae Vannucci, 1949 = Folliculina gigantea (Dons, 1910) (Ciliophora), after Vannucci (1955) Filellum magnificum Pe��a Cantero et al., 2004 b Filellum minimum Garc��a Corrales et al., 1979 = Folliculina gigantea (Dons, 1910) (Ciliophora), after Pe��a Cantero et al. 1998 Filellum nitidum Watson, 2005 Filellum parasiticum (Antsulevich, 1987) (from Lineolaria parasitica Antsulevich, 1987) Filellum plicatum (Hartlaub, 1904) species inquirenda (from Lafoea plicata Hartlaub, 1904) Filellum serpens (Hassall, 1848) (from Campanularia serpens Hassall, 1848) Filellum serratum (Clarke, 1879) (from Lafoea serrata Clarke, 1879) Filellum tubiforme Schydlowsky, 1902 = Filellum serpens (Hassall, 1848), Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 3-5, DOI: 10.5281/zenodo.206783, {"references":["Hincks, T. (1868) A history of the British hydroid zoophytes. John van Voorst, London, 338 pp.","Gray, J. E. (1848) List of the specimens of British animals in the collection of the British Museum. Part I. Centroniae or radiated animals. British Museum, London, 173 pp.","Naumov, D. V. (1960) Gidroidi i gidromedusy morskikh, solonovatovodnykh i presnovodnykh basseinov SSSR. Opredeleteli pro faune SSSR, 70, 1 - 626.","Naumov, D. V. (1969) Hydroids and Hydromedusae of the USSR. Israel Program for Scientific Translation, Jerusalem, 660 pp.","Cuvier, G. L. C. F. D. (1798) Tableau elementaire de l'histoire naturelle des animaux. Baudouin, Paris, 710 pp.","Lamouroux, J. V. F., Bory de Saint-Vincent, J. B. G. M. & Deslongchamps, E. (1824) Histoire naturelle des zoophytes ou animaux rayonnes, faisant suite a l'Histoire naturelle des Vers, de Brugiere. In: Didirot, D. & d'Alembert, J. L. R. (Eds), Encyclopedie methodique. Panckoucke, Paris, pp. 1 - 819.","Bedot, M. 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Verlag von Gustav Fischer, Jena, 66 pp.","Broch, H. (1933) Zur Kenntnis der adriatischen Hydroidenfauna von Split. Arten und Variationen. Skrifer utgitt av det Norske Videnskaps-Akademi i Oslo, I. Matematisk-Naturvidenskapelig Klasse, 4, 1 - 115.","Leloup, E. (1933) Contribution a la connaissance des hydropolypes de la cote des Pays-Bas. Bulletin du Musee Royal d'Histoire Naturelle de Belgique, 9, 1 - 30.","Berezina, N. A. (1948) Opredelitel' fauny I flory severnykh morei SSSR. Sovetskaya Nauka, Otryad Leptolina, Moscow, 19 pp.","Leloup, E. (1935) Hydraires calyptoblastiques des Indes Occidentales. (Zoologische Ergebnisse einer Reise nach Bonaire, Curacao und Aruba im Jahre 1930, Nº 13). Bulletin du Musee Royal d'Histoire Naturelle de Belgique, 2, 1 - 73.","Kramp, P. L. (1913) Hydroids collected by the \" Tjalfe \" expedition to the west coast of Greenland in 1908 and 1909. Videnskablige Meddelelser fra Dansk Naturhistorisk Forening i Kobenhavn, 66, 1 - 36.","Broch, H. (1918) Hydroida. The Danish Ingolf-Expedition, 5, 1 - 206.","Mathisen, O. (1928) Hydroids from northern Norway, with a list of unreported specimens in the Tromso Museum. Tromso Museums Aarshefter, 49, 1 - 40.","Remane, A. (1933) Verteilung und Organization der benthonischen Mikrofauna der Kieler Bucht. Wissenschaftliches Meeresuntersuchungen, Abteil Kiel, 21, 161 - 221.","Leloup, E. (1938) Quelques hydropolypes de la baie de Sagami, Japon. Bulletin du Musee Royal d'Histoire Naturelle de Belgique, 14, 1 - 22.","Leloup, E. (1947) Les coelenteres de la faune Belge. Leur bibliographie et leurs distribuition. Bulletin du Musee Royal d'Histoire Naturelle de Belgique, 107, 1 - 73.","Vervoort, W. (1942) Northern Hydroida in the collections of the Rijksmuseum van Natuurlijke Historie and the Zoological Museum at Amsterdam, with notes on their distribution. Zoologische Mededelingen, 23, 275 - 312.","Vervoort, W. (1959) The Hydroida of the tropical west coast of Africa. 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Kongliga Vetenskaps Academiens Nya Handlingar, 14, 243 - 260.","Fraser, C. M. (1925) Some new and previously unreported hydroids, mainly from the Californian coast. University of California Publications in Zoology, 28, 167 - 172.","Jullien, J. (1880) Description d'une espece nouvelle du genre Filellum. Bulletin de la Societe Zoologique de France, 5, 290 - 292.","Nutting, C. C. (1905) Hydroids of the Hawaiian Islands collected by the steamer Albatross in 1902. Bulletin of the United States Fish Commission, 23, 931 - 959.","Levinsen, G. M. R. (1893) Meduser, ctenophorer og hydroider fra Gronlands vestkyst, tilligemed bemaerkninger om hydroidernes systematik. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 5, 143 - 212, 215 - 220.","Dons, C. (1910) Zoologiske notiser I. Bermerkninger om forveksling av Folliculina med Filellum. Tromso Museums Aarshefter, 31 - 32, 189 - 194.","Vannucci, M. (1955) On the newly liberated medusa of Obelia hyalina Clarke 1879. Dusenia, 6 (1 - 2), 55 - 60.","Vannucci, M. (1949) Hydrozoa do Brasil. Boletim da Faculdade de Filosofia, Ciencias e Letras da Universidade de Sao Paulo, Zoologia, 99, 219 - 266.","Pena Cantero, A. L., Svoboda, A. & Vervoort, W. (2004 b) Antarctic hydroids (Cnidaria, Hydrozoa) of the families Campanulinidae, Lafoeidae and Campanulariidae from recent Antarctic expeditions with R. V. Polarstern, with the description of a new species. Journal of Natural History, 38, 2269 - 2303.","Garcia Corrales, P., Buencuerpo Arcas, V. & Peinado de Diego, M. V. (1979) Contribucion al conocimiento de los hidrozoos de las costas espanolas. Parte II: \" Lafoeidae \", \" Campanulinidae \", y \" Syntheciidae \". Boletin del Instituto Espanol de Oceanografia, 5, 5 - 39.","Watson, J. E. (2005) Hydroids of the Archipelago of the Recherche and Esperance, Western Australia: Annotated list, redescription of species and description of new species. In: Weills, F. E., Walker, D. I. & Kendrick, G. A. (Eds), The marine flora and fauna of Esperance, Western Australia. Western Australian Museum, Perth, pp. 495 - 612.","Antsulevich, A. E. (1987) Gidroidy shel'fa Kuril'skykh ostrovov (Hydroids from the shelf waters of Kurile Islands). Zoologicheskii Institut, Akademiya Nauk SSSR, Leningrad, 165 pp.","Schydlowsky, A. (1902) Materialy po faune gidroidov arkticheskikh morei. I. Gidroidy Belogo morya u beregov Solovetskikh ostrovov (Data on the hydroid fauna of the Arctic Seas I. Hydroids of the White Sea near the Coasts of the Solovetskie Islands). Trudy Obshchestva Ispytatelei Prirody pri Imperatorskom Khar'kovskom Universitete, 36, 3 - 268."]}

Filellum bouvetensis sp. nov. (Figure 2 A���D) Filellum contortum: Pe��a Cantero and Gili 2006: 766. Non Lafoea contorta Nutting, 1905: 945, pl. 3, fig. 6, pl. 9, figs. 8���9 [= Filellum contortum (Nutting, 1905)]. Type series. Holotype��� Filellum contortum (Nutting, 1905) (PS 65 /028), with coppiniae. Type locality. From the holotype, 54 o 22.49 ������ 54 o 22.54 ���S; 0 3 o 17.58 ������ 03o 17.21 ���E, 122���134m (Pe��a Cantero & Gili 2006: 765, 766). Material examined. ANTARCTIC OCEAN: Bouvet Island, PS 65 / 28 (25 November 2003, 122��� 134m), colonies epibiotic on various hydroids (Antarctoscyphus elongatus, Schizotricha multifurcata, Sertularella sanmatiasensis, Symplectoscyphus sp.); with coppinia. Description. Colonies stolonal, with filiform creeping hydrorhiza. Hydrotheca sessile, arising uprightly and almost completely free from hydrorhiza. When present, abcauline side of adnate part provided with a few striae. Hydrothecal aperture circular and slightly everted, usually with very long rim renovations. Free adcauline hydrothecal part 0.65 ��� 0.187 mm long. Hydrothecal aperture 0.18���0.25 mm wide. Coppiniae up to 4 mm wide and 2 mm high, some almost completely surrounding the stem of the host hydroid, forming a ring of ca. 4 mm wide. Coppinia composed of firmly adpressed gonothecae with long defensive tubes at periphery, arching over the centre of the gonothecal mass and forming a corbula-like structure. Defensive tubes unforked and with distal aperture. Gonothecae elongated, roughly polygonal in apical view, but with a rather inconstant arrangement in lateral view; distal most part of gonothecae usually free and with irregular shape. Gonothecal aperture circular, situated on a very short, frequently unconspicuous, non-everted distal neck, 0.55���0.6 mm high, 0.1���0.2 mm wide and aperture 0.55��� 0.6 mm wide. Cnidome composed of microbasic mastigophores (not seen discharged), 6.5���8.5 X 3 ���3.5 ��m. Distribution. Known only from Bouvet Island (Pe��a Cantero & Gili 2006). Remarks. The material from Bouvet Island, here described as Filellum bouvetensis sp. nov., was identified as Filellum contortum (Nutting, 1905) by Pe��a Cantero and Gili (2006). Filellum bouvetensis sp. nov. is similar to Nutting���s (1905) species in the shape and size of the hydrothecae having a very long free hydrothecal part. However, the new species has larger nematocysts than those found in the lectotype of F. c o nt o r tu m. Furthermore, as indicated above, Nutting���s species was deprived of coppinia, rendering it an insufficiently characterized species to properly assign new material to. Therefore we have decided to consider the abundant, fertile, well-preserved and characterized material from Bouvet suitable for the creation of a new species. Filellum bouvetensis sp. nov. is similar to F. a n t a rc t i c u m in the general shape of the coppinia, both with defensive tubes arching over the gonothecal mass, so that they form a sort of a corbula. Moreover, in dorsal view, the gonothecal aperture of F. bouvetensis sp. nov. is similar to that of F. a n t a rc t i c u m by the presence of a perisarc ring. However, in Hartlaub���s species the gonothecae lack any sort of neck and are distally homogeneous, being completely laterally adpressed. Both species also differ in the size and general proportions of the hydrothecae. Filellum bouvetensis sp. nov. also resembles F. conopeum Watson, 2003 in the structure of the coppinia with defensive tubes curving over the gonothecae. However, Watson (2003) described the coppinia as bud-shaped, ca. 1 mm wide and 1 mm high, whereas in F. bouvetensis they are corbula-shaped and up to 4 mm wide and 2 mm high. Anyway, both species are clearly distinguishable by the size and relative proportions of the hydrothecae. Here a new name is proposed for the Bouvet specimens, which can be clearly distinguished from the remaining known species of Filellum by the size of the nematocysts and hydrothecae, the structure of the coppinia and the shape and size of the gonothecae. Etymology. The name bouvetensis refers to Bouvet, the name of the remote Southern Ocean island where this species was found., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 10-11, DOI: 10.5281/zenodo.206783, {"references":["Pena Cantero, A. L. & Gili, J. M. (2006) Benthic hydroids (Cnidaria, Hydrozoa) from off Bouvet Island (Antarctic Ocean). Polar Biology, 29, 764 - 771.","Nutting, C. C. (1905) Hydroids of the Hawaiian Islands collected by the steamer Albatross in 1902. Bulletin of the United States Fish Commission, 23, 931 - 959."]}

Filellum adhaerens (Nutting, 1901) (Figures 1 A���E) Lafoea adhaerens Nutting, 1901: 178, pl. 21, figs. 3���4. Filellum adherens: Pe��a Cantero et al. 1998: 299 ��� 230. Type series. Neotype��� Lafoea adhaerens Nutting, 1901, former University of California no. 5245 (Nutting 1901 and specimen���s label) (USNM 43728). Type locality. From the neotype, Harriman Expedition 1899, Kodiak Harbor, Alaska, epibiotic on Abietinaria turgida (Clark, 1876). The data coincide with those of the original description. Material examined. Neotype USNM 43728. Description of neotype. Colony stolonal, growing on stem and branches of Abietinaria turgida (Clark, 1876), arising from a creeping filiform, though flattened, hydrorhiza, 0.07���0.08 mm wide; hydrorhizal tubes and hydrothecae covered by a thin sheath (possibly produced by the host), with anastomosed perisarc, forming a tight mesh with no free space left in some parts of the sertulariid substrate. Hydrothecae sessile, arising with no definite pattern from hydrorhiza, varying in form, from almost completely adnate to substratum (either parallel to or over the hydrorhizal stolon), to provided with a short free part (especially both where substrate ends and where hydrothecae ascend one another). Hydrothecae irregularly curved in diverse patterns, in varied degrees to horizontal plane, rarely emerging from substrate plane. Perisarc tube connecting hydrothecae and hydrorhiza thinner than hydrothecae themselves. Adnate portion of hydrothecae tubular or maximally slightly flattened, 0.32���0.53 mm (0.41 �� 0.08, n= 10) long, perisarc smooth; free part cylindrical, smooth, 0.00��� 0.19 mm (0.08 �� 0.06, n= 10) long; perisarc moderately thin; hydrothecae not widening distally, margin even and smooth, with up to 4 renovations, rarely slightly flaring; hydrothecal aperture circular, 0.12���0.14 mm (0.13 �� 0.01, n= 10) wide, perpendicular to axis of free part of hydrotheca. Hydranths not seen. Gonothecae packed in coppinia up to 2 mm wide. Gonothecae closely set, contiguous, perimeters distinct in dorsal view, general form elongated flask-shaped. Gonothecae 0.47���0.70 mm (0.54 �� 0.06, n= 10) high (from substrate to distal plateau), with narrowest part adnate to substratum, distal part widening until forming a distal plateau varying from flattened to roof-shaped, plateau 0.14���0.24 mm (0.18 �� 0.03, n= 10) wide; apical part with distal short neck, 0.01���0.03 mm (0.02 �� 0.01, n= 10) high (from plateau to aperture), bearing circular aperture, 0.03���0.05 mm (0.04 �� 0.01, n= 10) wide, rim even, not flared, without renovations. Protective tubes present, curved, hollow, with distal aperture, arising amongst gonothecae; some protective tubes merge with each other. Planulae diameter larger than gonothecal aperture. Nematocysts not seen. Distribution. Filellum adhaerens was recorded only at the type locality, Kodiak Harbor, Alaska by Nutting (1901). Remarks. There is a single specimen of the paratype of Filellum adhaerens (Nutting, 1901), originally Lafoea adhaerens Nutting, 1901, in the collection of the Smithsonian National Museum of Natural History (USNM 43728). This specimen has the same information as the original description on its label, which states: ��� paratype undoubtedly type lost���has Nutting���s label���. Indeed, as far as we know, there is no other specimen of the type series and, therefore, we designate the material USNM 43728 as the neotype of Lafoea adhaerens Nutting, 1901. Nutting (1901: 178), in the original description of Lafoea adhaerens, remarked that ���this interesting species is so different in appearance from the others of the genus that I was at first inclined to make it the type of a new genus���. The author described the species as ���forming an encrusting mass of adherent rootstocks disposed both longitudinally and transversely over colonies of other hydroids, the tubes of the rootstock interwoven much like the threads of a fabric. Hydrothecae sessile, tubular, often more or less curved, aperture facing upwards, entire; margin slightly expanded. The hydrothecae are very irregularly disposed, being much more crowded in some places than in others��� (Nutting 1901: 174). Inconsistently with his assignment, he described Filellum as ���stem a creeping, slender rootstock, parasitic on other hydroids, often forming a reticulate structure. Hydrothecae curved, decumbent, and partly adherent; margin entire, without operculum��� (Nutting 1901: 179). Therefore, it was only a matter of time before the species was correctly assigned to Filellum (Stechow 1913 b: 30; see also Pe��a Cantero et al. 1998: 299). Filellum adhaerens is a species characterized by the structure of its coppinia, with the juxtaposed bottle-shaped gonothecae with a short distal neck and protective tubes amongst the gonothecae. Nutting (1901) considered the coppinia similar to that of Lafoea dumosa (Fleming, 1820) and described it as ���being composed of closely packed gonangia interspersed with long, tubular, variously curved modified hydrothecae. The gonangia are flask-shaped, with a tubular neck and small aperture��� (Nutting 1901: 178)., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 5-7, DOI: 10.5281/zenodo.206783, {"references":["Nutting, C. C. (1901) Papers from the Harriman Alaska Expedition. XXI. The hydroids. Proceedings of the Washington Academy of Sciences, 3, 157 - 216.","Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315.","Clark, S. F. (1876) Report on the hydroids collected on the coast of Alaska and the Aleutian islands by W. H. Dall, U. S. Coast Survey and party from 1871 to 1874 inclusive. Proceedings of the Academy of Natural Sciences of Philadelphia, 1876, 209 - 235.","Stechow, E. (1913 b) Hydroidpolypen der japanische Ostkuste. II. Teil: Campanularidae, Halecidae, Campanulinidae und Sertularidae, nebst Erganzungen zu den Athecata und Plumularidae. Abhandlungen der Mathematisch-Physikalischen Klasse der Koniglich Bayerischen Akademie der Wissenschaften, 3, 1 - 162.","Fleming, J. (1820) Observations on the natural history of the Sertularia gelatinosa of Pallas. Edinburgh Philosophical Journal, 2, 82 - 89."]}

Filellum bouvieri Jullien, 1880 Filellum Bouvieri Jullien, 1880: 291 ���292, figure not numbered.? Filellum bouvieri: Pe��a Cantero et al. 1998: 301. Type series. We found no information on the type specimens of Filellum bouvieri Jullien, 1880, and none may exist. Distribution. Filellum bouvieri was only recorded from the Cape Verde Islands in its original description by Jullien (1880). Remarks. Filellum bouvieri was described by Jullien (1880: 291) from material collected in Cape Verde Islands, diagnosed by the hydrorhizae ���gr��les, corn��es, non entrelac��es���, with branching points ���naissant ordinairement �� angle droit les uns des autres��� and ���tortueux, alternes���. There is a single hydrotheca at the ramification diagnosed as ���p��doncul��e��� and composed of two parts, ���l'une rampante et post��rieure est renfl��e en son milieu��� and ���l'autre dress��e quelquefois jusqu'�� angle droit sur la premi��re���. The hydrotheca is ���cylindrique��� and with ���une ouverture un peu ��vas��e���. According to Jullien (1880: 292), F. bouvieri is different from F. s e r p e n s because it has hydrorhizae ���beaucoup plus gr��les��� and hydrothecae ���plus coud��es���. However, as remarked by Pe��a Cantero et al. (1998: 301), the material described by Jullien (1880) was not fertile and trophosomal characters are not enough to distinguish it from the other species of Filellum. Specimens of F. bouvieri were never collected since the original description and, therefore, the validity of this species is dubious (see also Pe��a Cantero et al. 1998: 301). We consider it as species inquirenda., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on page 22, DOI: 10.5281/zenodo.206783, {"references":["Jullien, J. (1880) Description d'une espece nouvelle du genre Filellum. Bulletin de la Societe Zoologique de France, 5, 290 - 292.","Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315."]}

Filellum adnatum (Fraser, 1925) Figures 5 A���C Lafoea adnata Fraser, 1925: 170 ���171, figs. 5 a���b. Type series. Lectotype��� Lafoea adnata Fraser, 1925 (BCPM 976 - 324 - 1). Type locality. From the lectotype, United States Bureau of Fisheries Steamer ���Albatross���, ���near Farallon Islands [California], in 33���35 fathoms [59���63 m], on algae, bryozoa, and other hydroids��� (Fraser 1925: 171). Material examined. Lectotype BCPM 976 - 324 - 1. Description of lectotype. Colony stolonal growing on algae, with filiform creeping hydrorhiza, 0.08���0.1 mm wide; hydrorhizal tubes and hydrothecae reticulated, with anastomosed perisarc, forming a more or less tight mesh. Hydrothecae sessile, arising with no definite pattern from hydrorhiza, varying in form, from almost completely adnate to substratum (either parallel to or over hydrorhizal stolon) to provided with a short free part. Perisarc tubes between hydrothecae and hydrorhiza thinner than hydrothecae themselves. Adnate portion of hydrothecae tubular or maximally slightly flattened, 0.24���0.46 mm (0.34 �� 0.07, n= 10) long, slightly constricted at base, perisarc with numerous striae on upper part up to hydrothecal curvature; free part cylindrical, arising at angles larger than 60 o, mostly 90 o, smooth, 0.2���0.32 mm (0.25 �� 0.05, n= 10) long; perisarc moderately thin; hydrothecae widening distally, margin even and smooth, with up to three renovations; hydrothecal orifice circular, 0.16���0.2 mm (0.19 �� 0.01, n= 10) wide, perpendicular to long axis of free part of hydrotheca. Hydranths badly preserved, with ca. 8���10 tentacles, hypostome conical. Hydranths, when retracted, lying at adnate space of hydrothecae. Coppinia absent. Nematocysts of one category (heterotrichous microbasic?mastigophores), but of two size classes (not seen discharged): 6.0���7.0 X 2.5 ���3.0 ��m (6.45 �� 0.50 X 2.76 �� 0.5, n= 10), rice-shaped and common, and 22.0���25.0 X 8.0���10.0 ��m (23.50 �� 1.12 X 8.80 �� 0.76, n= 5), ovoid and rare. Distribution. Filellum adnatum was recorded only for the Farallon Islands (California) by Fraser (1925). Remarks. Fraser (1925: 170���171) described Lafoea adnata as a new species with hydrotheca ���attached to a reticulate stolon, with no definite pedicels, although there is a definite constriction at the base of each; in nearly all cases the wall of the hydrotheca is attached to the stolon for a part of its length but the proportion varies very materially���. However, the specimen described by Fraser (1925) does not refer to Lafoea, a genus with ���colonies occasionally stolonial but usually erect, branched, with a polysiphonic hydrocaulus. Hydrothecae pedicellate, usually free from stem or stolon��� (Calder 1991: 36). The specimen designated as lectotype of Lafoea adnata Fraser 1925 is stolonal with sessile hydrothecae and, therefore, should be assigned to the genus Filellum. Although not remarked on by Fraser (1925), the material described by him is striated on the adnate part of the hydrotheca, a characteristic also observed in F. antarcticum, F. magnificum, F. nitidum and F. serratum. Inasmuch as the material lacks coppiniae, its identification is uncertain and, therefore, the taxon is here considered as species inquirenda., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 20-22, DOI: 10.5281/zenodo.206783, {"references":["Fraser, C. M. (1925) Some new and previously unreported hydroids, mainly from the Californian coast. University of California Publications in Zoology, 28, 167 - 172.","Calder, D. R. (1991) Shallow-water hydroids of Bermuda: The Thecatae, exclusive of Plumularioidea. Royal Ontario Museum, Life Sciences Contributions, 154, 1 - 140."]}

Filellum parasiticum (Antsulevich, 1987) (Figure 4 A���B) Lineolaria parasitica Antsulevich, 1987: 57 ���58, fig. 14. Filellum parasiticum: Antsulevich and Vervoort 1993: 431 ���433, figs. 1 a���b, 2 a���b; Pe��a Cantero et al. 1998: 302. Type series. Holotype��� Lineolaria parasitica Antsulevich, 1987, infertile colony on Abietinaria abietina (Linnaeus, 1758) (ZIRAS no. 10051 / 1). Paratypes��� Lineolaria parasitica Antsulevich, 1987, infertile colony on Abietinaria abietina (Linnaeus, 1758) (ZIRAS no. 2 / 10052); infertile colony on Abietinaria abietina (Linnaeus, 1758) (ZIRAS no. 3 / 10052.1); infertile colony on Abietinaria abietina (Linnaeus, 1758) (RMNH Coel. No. 26770 sample and 3 slides no. 1820). Type locality. From the holotype and paratypes, central part of Sea of Okhotsk, SE of Iona Island, depth 160 m, 4 August 1986, leg. AV Smirnoff. Material examined. Holotype ZIRAS 10051 / 1; Paratypes ZIRAS 2 / 10052; ZIRAS 3 / 10052.1; RMNH Coel. 26770 (sample and 3 slides no. 1820). Description of holotype. (Complemented with data from Antsulevich 1987: 57���58; after translation in Antsulevich & Vervoort 1993: 432���433): Colony stolonal, with filiform, although flattened, creeping hydrorhiza, 0.055���0.074 mm wide, growing over 20 mm long fragment of stem and proximal parts of branches of Abietinaria abietina (Linnaeus, 1758); hydrorhizal tubes and hydrothecae with anastomosed perisarc, forming a more or less tight mesh with no free space in some parts of the sertulariid substrate. Hydrothecae sessile, arising with no definite pattern from hydrorhiza, usually completely adnate to substratum, parallel to, or over hydrorhizal stolon or, exceptionally, with short free part at end of substrate or in case of overlapping hydrothecae. Hydrothecae irregularly curved to varied degree in diverse patterns with horizontal plane, but not emerging from substrate plane. Perisarc tube of hydrorhiza thinner than hydrothecae. Adnate portion of hydrothecae tubular or maximally slightly flattened, 0.42��� 0.60 mm (0.52 �� 0.05, n= 10) long, perisarc smooth; free part cylindrical, smooth, 0.00��� 0.08 mm (0.03 �� 0.03, n= 10) long; perisarc moderately thin; hydrothecae not widening distally, margin even and smooth; many hydrothecae laterally depressed, with no renovations, rarely slightly flaring; hydrothecal aperture near circular, 0.12���0.16 mm (0.14 �� 0.01, n= 10) wide, perpendicular to long axis of hydrotheca. Hydranths 0.12���0.20 mm high (0.16 �� 0.02, n= 10), 0.10���0.14 mm wide (0.12 �� 0.01, n= 10) at base of tentacles, with ca. 10���16 tentacles, hypostome conical. Hydranths, when retracted, lying at base of the hydrothecae, with very truncated base. Sometimes coenosarc preceding hydranth expanded laterally. Gonothecae not seen. Nematocysts of one category, heterotrichous microbasic euryteles (not seen discharged), 5.5���7.5 X 2.5���3.5 ��m (6.15 �� 0.41 X 2.90 �� 0.39, n= 10), bean-shaped, common. Description of paratypes. Colonies stolonal, with filiform although flattened and creeping hydrorhiza, growing over 5 mm (ZIRAS 2 / 10052) and 40 mm (ZIRAS 3 / 10052.1) long portions of stem and branches of Abietinaria abietina; hydrorhizal tubes and hydrothecae with anastomosed perisarc, forming a cover in some parts of the sertulariid substrate. Hydrothecae sessile, arising with no definite pattern from hydrorhiza, usually completely adnate to substratum, parallel to, or over hydrorhizal stolon or, maximally, with short free part either at end of substrate, or in case of overlapping hydrothecae, or renovations of hydrothecae resulting in an ascending free part. Hydrothecae irregularly curved to varied degree in diverse patterns at horizontal plan, occasionally from substrate plane. Perisarc tube of hydrorhiza thinner than hydrothecae. Adnate portion of hydrothecae tubular or maximally slightly flattened, 0.33��� 0.38 mm (0.35 �� 0.02, n= 7) long, perisarc smooth; free part cylindrical, smooth, 0.00��� 0.08 mm (0.05 �� 0.03, n= 11) long; perisarc moderately thin; hydrothecae not widening distally, margin even and smooth, many laterally depressed with projections of abcaulinar and adcaulinar walls, with up to 6 renovations emerging from substrate plane, rarely slightly flaring; hydrothecal aperture near circular, 0.11���0.15 mm (0.13 �� 0.02, n= 9) wide, perpendicular to long axis of hydrotheca. Hydranths, when retracted, lying at base of the hydrothecae, with very truncated base. Sometimes coenosarc preceding hydranth expanded laterally. Gonothecae not seen. Nematocysts of two categories: heterotrichous microbasic euryteles (not seen discharged), approximately same dimensions as holotype, bean-shaped, common; heterotrichous microbasic mastigophores (not seen discharged), rice-shaped, only one nematocyst seen, 15.5 X 4 ��m. Additional data. Antsulevich and Vervoort (1993: 432���433) described the gonosome of the species: ���the gonosome is a typical coppinia, surrounding an internode of A. abietina for length of 3 mm. Gonothecae closely packed, contiguous laterally, usually 5���6 sided as viewed from above but many varied in shape; slender, invertedpyramidal, apically contracted and there with remarkable short neck leading to circular rim. Protecting tubes numerous, arising between gonothecae, usually curving���. Distribution. Filellum parasiticum is only known from the Okhotsk Sea and Kurile Islands, NE Pacific. Remarks. This species was originally included in the genus Lineolaria for the Kuril Islands, northern hemisphere (Antsulevich 1987). Watson (1992: 81) considered the species ���to have closer affinities with the Lafoeidae than with the Lineolariidae ���, and Antsulevich and Vervoort (1993: 431���433) transferred the species to the genus Filellum after the study of fertile material with coppinia. Examination of the paratypes revealed some interesting differences and a complementary knowledge in relation to the holotype of F. parasiticum. The first important difference is the presence of renovations in this material. Unlike the holotype, these renovations are present in several hydrothecae, becoming free part not only because of the end of the substrate (as seen in the holotype). Another distinct and important difference is the presence of the rice-shaped microbasic mastigophore nematocysts. This type of nematocyst is apparently the basic type of other Lafoeidae (except for Cryptolarella; see Marques et al. 2006 a) and its occurrence would be expected in F. parasiticum as well. However, after 5 preparations, only one of these nematocysts was seen. The rarity of the nematocyst is, however, indicative of the difficulties in assigning its presence in the species of Filellum. Antsulevich and Vervoort (1993: 431���433) examined 18 colonies, one of them fertile, equally gathered from the Sea of Okhotsk (northern part, at 120 m and 150 m; central part, SE from St Jonas Is., at 147 m and 160 m, and SE of Cape Terpeniya, Sakhalin Is., at 396 m, 750 m and 1960���2000 m), growing on Abietinaria abietina, although one was growing on Sertularia sp. The fertile colony had a coppinia and, therefore, the authors transferred the species to the genus Filellum, under the name F. parasiticum (Antsulevich, 1987). The material with coppinia does not constitute part of the type series. Although it is supposedly present in the collections of the Zoological Institute of the Academy of Sciences of Russia (ZIRAS), St. Petersburg (see Antsulevich & Vervoort 1993: 431), it could not be located (S. Stepan���yants, pers. comm.). Reanalysis of the material described by Antsulevich and Vervoort (1993), with special emphasis on the cnidome composition, is most wanted because their specimen may possibly belong to a different species. The hydrothecae and hydrorhiza of F. adhaerens form a mesh laying on the substrate, somewhat similar to the pattern observed in F. parasiticum. However, F. parasiticum was considered distinguishable ���from other species of the genus by the long adnate part of hydrotheca and rather shorter free portion, by anastomosing hydrorhiza and closely packed hydrothecae forming a discontinuous carpet...the gonotheca of F. parasiticum has a short neck lacking in F. s e r p e n s (Hassall, 1848) (cf. Naumov 1960; 1969) and F. s e r r a tu m (Clarke, 1879) (cf. Ritchie 1911; Millard 1975)��� (Antsulevich and Vervoort 1993: 443, although we observed a short neck in F. serratum), similar characteristics to those listed by Pe��a Cantero et al. (1998: 302: ���characterized by the prostrate hydrothecae, typically fully adnate to the substratum, though towards the periphery of the colony the distal one-fourth to one-fifth of the hydrotheca curves upwards, being free from the substratum... frequent anastomoses of the hydrothecae and hydrorhizal stolons forming a continuous layer completely covering the substratum���)., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 15-17, DOI: 10.5281/zenodo.206783, {"references":["Antsulevich, A. E. (1987) Gidroidy shel'fa Kuril'skykh ostrovov (Hydroids from the shelf waters of Kurile Islands). Zoologicheskii Institut, Akademiya Nauk SSSR, Leningrad, 165 pp.","Antsulevich, A. E. & Vervoort, W. (1993) Some little-known species of hydroids (Cnidaria: Hydrozoa: Lafoeidae) and description of Papilionella pterophora gen. nov., spec. nov. (Sertulariidae). Zoologische Mededelingen, 67, 431 - 443.","Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315.","Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I, editio decima, reformata, Holmiae, Laurentii Salvii, 823 pp.","Watson, J. E. (1992) Revision of the family Lineolariidae Allman, 1864 (Hydrozoa: Hydroida). Proceedings of the Royal Society of Victoria, 104, 81 - 87.","Marques, A. C., Pena Cantero, A. L. & Migotto, A. E. (2006 a) An overview of the phylogeny of the families Lafoeidae and Hebellidae (Hydrozoa: Leptothecata): their composition and classification. Invertebrate Systematics, 20, 43 - 58.","Hassall, A. H. (1848) Definitions of three new British zoophytes. Zoologist, 6, 2223.","Naumov, D. V. (1960) Gidroidi i gidromedusy morskikh, solonovatovodnykh i presnovodnykh basseinov SSSR. Opredeleteli pro faune SSSR, 70, 1 - 626.","Naumov, D. V. (1969) Hydroids and Hydromedusae of the USSR. Israel Program for Scientific Translation, Jerusalem, 660 pp.","Clarke, S. F. (1879) Report on the Hydroida collected during the Exploration of the Gulf Stream and Gulf of Mexico by Alexan- der Agassiz, 1877 - 78. Bulletin of the Museum of Comparative Zoology at Harvard College, in Cambridge, 5, 239 - 252.","Ritchie, J. (1911) Hydrozoa (hydroid zoophytes and Stylasterina) of the ' Thetis' expedition. Memoirs of the Australian Museum, 4, 807 - 869.","Millard, N. A. H. (1975) Monograph on the Hydroida of Southern Africa. Annals of the South African Museum, 68, 1 - 513."]}

Filellum magnificum Pe��a Cantero, Svoboda and Vervoort, 2004 b Filellum magnificum Pe��a Cantero et al., 2004 b: 2287 ���2289, figs. 2g ���h, 5 a���c; Pe��a Cantero 2010: 765, figs. 3 d, e. Type series. Holotype��� Filellum magnificum Pe��a Cantero et al., 2004 b, several hydrothecae on Billardia subrufa, with coppinia (RMNH-Coel. 30789) (Pe��a Cantero et al. 2004 b: 2271, 2287). Type locality. From the holotype, Weddell Sea, Antarctic, ANT II- 4 Stn 502, 21 February 1984, several hydrothecae on Billardia subrufa, with coppinia (Pe��a Cantero et al. 2004 b: 2271). Material examined. German Polarstern Antarctic Cruises: ANT I- 2 Stn 135, off Riiser-Larsen Ice Shelf, Princess Martha Coast, Weddell Sea (73 �� 41.6 ���S; 20 �� 55.3 ���W, 205m, 8 February 1983), on Stegella lobata (RMNH- Coel. 30779); ANT II- 4 Stn 474, off Ronne Ice Shelf, Weddell Sea (76 �� 56.7 ���S; 49 �� 44.0���W, 220m, 14 February 1984), on Billardia subrufa (RMNH-Coel. 30788); ANT II- 4 Stn 502, Weddell Sea (21 February 1984), on B. subrufa, with coppinia (holotype, RMNH-Coel. 30789); ANT VIII- 5 Stn 16-481 (74.74 �� S; 61.13 �� W, off Ronne Ice Shelf, Weddell Sea, 620���640m, 15 February 1990), on spicules of a sponge (RMNH-Coel. 30802); ANT IX- 3 Stn 158 (72 �� 21.8 ���S; 16 �� 42.1 ���W, off Cape Norvegia, Princess Martha Coast, Weddell Sea, 623m, 13 February 1991), on B. subrufa (RMNH-Coel. 30806). Spanish Bentart 2003 Expedition: Stn 5 A, south of Peter I Island (68 �� 56 ��� 37 ��������� 68 �� 56 ��� 43 ������S; 90 �� 35 ��� 50 ��������� 90 �� 35 ��� 19 ������W, 124 m, 4 February 2003), on the ascidian Cnemidocarpa verrucosa, with coppiniae; Stn 8 A, east of Peter I Island (68 �� 50 ���05��������� 68 �� 50 ��� 16 ������S; 90 �� 21 ��� 15 ��������� 90 �� 21 ��� 16 ������W, 85���86 m, 6 February 2003), on Schizotricha vervoorti, Symplectoscyphus glacialis and? Tubularia stem, with coppinia; Stn 8 T, east of Peter I Island (68 �� 50 ���08������S; 90 �� 21 ��� 17 ������W, 87 m, 6 February 2003), on Schizotricha vervoorti, with coppiniae. Description. (After Pe��a Cantero et al. 2004 b: 2287): Colony stolonal, composed of creeping stolons giving rise to partially sessile hydrothecae. Hydrotheca tubular, sock-shaped, and adnate to substratum for just a tiny portion. Hydrothecal aperture circular, 169���208 ��m wide; rim even and smooth, though indistinctly everted. Free part of hydrotheca, 436���670 ��m long, directed upwards at varied degrees, frequently with few, long renovations. Adnate part of hydrothecae, 208���300 ��m long, provided with a few striae on dorsal side. Cnidome consisting of two size-classes of nematocysts (e.g. ANT II- 4 Stn 502): small (6.5���7.2 X 2.9 ��m) and occasionally large (18.2 X 5.2 ��m) capsules. In material from ANT VIII- 5 Stn 16-481 only the small nematocysts were observed (5.9���7.2 X 2.6���3.3 ��m). Coppinia consisting of firmly adpressed gonothecae with defensive tubes. Gonothecae 423���455 ��m long, 98��� 111 ��m wide, aperture circular, 52���58 ��m wide, on a short and distinctly everted distal neck. Defensive tubes situated on periphery of the group of gonothecae, but also arising amongst them, and provided with large, distal, circular aperture. Distribution. Filellum magnificum has been recorded only for the Antarctic, ���at depths from 205 to 640 m [���] south and east coasts of the Weddell Sea (off Ronne Ice Shelf, off Riiser-Larsen Ice Shelf and off Cape Norvegia, Princess Martha Coast)��� (Pe��a Cantero et al. 2004 b: 2289) and the south and east coast of Peter I Island, in the Bellingshausen Sea (Pe��a Cantero 2010). Remarks. See Pe��a Cantero et al. (2004 b: 2288���2289)., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on page 14, DOI: 10.5281/zenodo.206783, {"references":["Pena Cantero, A. L., Svoboda, A. & Vervoort, W. (2004 b) Antarctic hydroids (Cnidaria, Hydrozoa) of the families Campanulinidae, Lafoeidae and Campanulariidae from recent Antarctic expeditions with R. V. Polarstern, with the description of a new species. Journal of Natural History, 38, 2269 - 2303.","Pena Cantero, A. L. (2010) Benthic hydroids (Cnidaria: Hydrozoa) from Peter I Island (Southern Ocean, Antarctica). Polar Biology, 33, 761 - 773."]}

Filellum annulatum (Watson, 1973) Reticularia annulata Watson, 1973: 164, figs. 5���6. Filellum annulatum: El Beshbeeshy 1991: 71 ���73, figs. 15 a���b; Pe��a Cantero et al. 1998: 299. Non Corystolona annulata: Watson 2002: 334 ���335, figs. 1 a���e. Type series. Holotype������ NMV G 1922, microslide; G 2091, preserved material, remainder of holotype colony���S [denoting the sheltered side of Pearson Island], 17 m, on a small calcareous bryozoa��� (Watson 1973: 164). Type material was not examined. Type locality. From the holotype, sheltered side of Pearson Island, Great Australian Bight (Watson 1973). Remarks. Watson (1973) described and figured Reticularia annulata as a new species assigned to the family Lafoeidae. When Reticularia was recognized as a preoccupied name for a brachiopod, the species referred to this genus were assigned to the genus Filellum by Pe��a Cantero et al. (1998) (see discussion above). Watson (1973: 164) characterized her species ���by the close thecal rings [���] of uniform size, and the distance between them varies little along the entire length of the hydrotheca. They have developed by continuous apical reduplication during growth of the hydrotheca, the flange of each rib being a relict margin���. Pe��a Cantero et al. (1998: 299) considered Reticularia annulata Watson, 1973 actually to belong to the pterobranch hemichordate genus Rhabdopleura, but the authors gave no arguments supporting their conclusion. Filellum annulatum was subsequently recorded from Patagonia and redescribed in the dissertation of El Beshbeeshy (1991), including details and figure of the hydranth. El Beshbeeshy���s data undoubtedly refer to a hydroid and, most likely, to a species of Filellum. Whether the specimens assigned to F. annulatum by El Beshbeeshy are conspecific to the geographically distant material originally described by Watson is open to question. Watson (2002) proposed the new hydroid genus Corystolona, assigned it to the family Clavidae, and mistakenly placed it in ���Leptothecatae���. The holotype of the type species of Corystolona annulata (Watson, 1973) is NMV G 1922, as described by her in 1973. However, Watson (2002: 333) also collected ���Abundant colonies of a hydroid previously described as Reticularia annulata Watson, 1973 ��� and, based on this new material [(see Watson 2002: 334: ���Description (of Tasmanian material)���], redescribed and provided illustrations (her Figure 1) of a specimen that is undoubtedly a member of ��� Clavidae ��� (today considered part of family Oceaniidae, see Schuchert 2009). The rationale of the redescription and new assignment was that ���the small sample of Reticularia annulata from Pearson Island in the eastern Great Australian Bight had deeply withdrawn, partly decomposed, sterile hydranths and disposition of the specimen on the bryozoan host was such that the bases of the hydrocauli were obscured; the specimen was thus mistakenly referred to Reticularia Thomson, 1853 (junior synonym, Filellum Hincks, 1868). The smaller dimension of the type compared with those of the present specimens is almost certainly due [to] its being a young, infertile colony while the Tasmanian material ranges from young to aged��� (Watson 2002: 334). The dimensions of perisarc of the two specimens are obviously different, as well as their trophosomal morphology. Watson (2002) neither remarked about the reassignment of her species made by Pe��a Cantero et al. (1998) as a hemichordate, nor its subsequent redescription by El Beshbeeshy (1991). We believe that the two papers by Watson describe different species; the 1973 one being a Filellum, and the 2002 one being a species of Anthoathecata and indeed a new species of Oceaniidae. However, once Watson (2002) explicitly considered Corystolona annulata as linked to the holotype NMV G 1922, both the species and generic name shall not be considered for nomenclatural purposes, because the holotype was intended to be a Filellum, not an oceaniid. Therefore, we consider F. annulatum a valid name based on the holotype material NMV G 1922 (a microslide, and the remainder preserved material of the holotype colony NMV G 2091). The Tasmanian material NMV F 91280 (malinol mounted microslide, female colony, Port of Launceston, Tasmania, colonies from bryozoans on wharf pilings 1���4 m, Aquenal Pty Ltd, May, 2001, see Watson 2002: 334) undoubtedly represents a species of Corydendriinae (sensu Schuchert, 2004), and perhaps even the genus Corydendrium van Beneden, 1844, whose description was emended as ���erect, branching or stolonal hydroid colonies [���]��� (Schuchert, 2004: 335), therefore encompassing the diagnostic character ���strictly stolonal colonies��� (Watson, 2002: 334) presented by Corystolona., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 7-8, DOI: 10.5281/zenodo.206783, {"references":["Watson, J. E. (1973) Pearson Island Expedition 1969. 9. Hydroids. Transactions of the Royal Society of South Australia, 97 (3), 153 - 200.","El Beshbeeshy, M. (1991) Systematische, morphologische und zoogeographische unter-suchungen an den Thekaten Hydroiden des Patagonoschen schelfs. PhD Dissertation. Hamburg University, Hamburg, 390 pp.","Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315.","Watson, J. E. (2002) Corystolona, a new hydroid genus (Leptolida: Leptothecatae) from southern Australia. Memoirs of the Museum Victoria, 59, 333 - 336.","Schuchert, P. (2009) World Hydrozoa database. Available from http: // www. marinespecies. org / hydrozoa (accessed 24 October 2011).","Thomson, W. (1853) Notes on some British zoophytes. Annals and Magazine of Natural History, 2 nd series, 11, 442 - 444.","Hincks, T. (1868) A history of the British hydroid zoophytes. John van Voorst, London, 338 pp.","Schuchert, P. (2004) Revision of the European athecate hydroids and their medusa (Hydrozoa, Cnidaria): families Oceanidae and Pachycordylidae. Revue Suisse de Zoologie, 111, 315 - 369."]}

Filellum disaggregatum Pe��a Cantero, Garc��a Carrascosa and Vervoort, 1998 (Figure 3 A���C) Filellum disaggregatum Pe��a Cantero et al., 1998: 308 ���311, figs. 3 a���d, 4 a���e; Pe��a Cantero and Garc��a Carrascosa 2002: 53. Type series. Holotype��� Filellum disaggregatum Pe��a Cantero et al., 1998, alcohol specimen with coppiniae (University of Valencia Coel. No. 26 /MED CONG 57). Paratype��� Filellum disaggregatum Pe��a Cantero et al., 1998, (RMNH Coel. no. 27771, two slides no. 3558) (Pe��a Cantero et al. 1998: 298). Type locality. From the holotype and paratype, Chafarinas Islands, Mediterranean Sea, Stn. CONG 57, sunk boat, on algae, 10.5 m, August 1991, leg. University of Valencia (Pe��a Cantero et al. 1998: 298, 308). Material examined. CHAFARINAS ISLANDS: CONG 57 (��� 10.5 m depth���, 4 August 1991, ���Barco hundido���), several colonies on the algae Cystoseira spinosa Sauvageau, Rhodymenia sp. and Acrosorium uncinatum (Turner) Kylin; with coppiniae��� (Pe��a Cantero et al. 1998: 298, 308), holotype and paratype of Filellum disaggregatum Pe��a Cantero et al., 1998 (University of Valencia Coel. No. 26 / MED CONG 57, RMNH Coel. no. 27771, two slides no. 3558, respectively). Description of holotype. Colonies stolonal, arising from a filiform and creeping hydrorhiza growing on diverse substrata; hydrorhizal tubes 0.04���0.06 mm wide, with anastomosed perisarc, forming a more or less tight mesh, depending on substratum. Hydrothecae sessile, arising with no definite pattern from hydrorhiza, more than half of their total length adnate to substratum, parallel to hydrorhizal stolon, free part emerging from substrate plane at angles wide than 45 o, usually 90 o. Adnate portion of hydrothecae tubular or maximally slightly flattened, 0.25���0.33 mm (0.30 �� 0.03, n= 11) long, perisarc smooth; free part cylindrical, smooth, 0.12���0.25 mm (0.18 �� 0.04, n= 11) long; perisarc moderately thin; hydrothecae widening distally, margin even and smooth, with up to four renovations, tenuous flaring; hydrothecal aperture circular, 0.06���0.12 mm (0.09 �� 0.02, n= 11) wide, perpendicular to long axis of free part of hydrotheca. Hydranths 0.06���0.13 mm high (0.09 �� 0.02, n= 10), 0.08���0.11 mm wide (0.10 �� 0.01, n= 10) at the base of tentacles, with ca. 12���16 filiform tentacles, hypostome conical. Hydranths, when retracted, lying on the adnate space of the hydrothecae. Gonothecae arranged in coppinia, up to 3 mm wide, varying from closely to more loosely packed, their limits distinct in dorsal view. Gonothecae 0.18���0.24 mm (0.21 �� 0.02, n= 10) high (from substrate to aperture), with widest part adnate to substratum, distal part bent away from substratum; general form variable, from flask-shaped to pyriform, with variable diameter up to distal part, apical part with distal neck bearing circular (sometimes oval) aperture, 0.05��� 0.07 mm (0.06 �� 0.01, n= 10) wide, rim even and flared, occasionally with up to three renovations. Protective tubes absent, defensive function made by hydrothecae around or among gonothecae. Planulae complete their development outside gonothecae, connected to them through a mucous thread; planulae diameter larger than gonothecal aperture. Nematocysts of one category, heterotrichous microbasic mastigophores (not seen discharged), 21���23 X 4.5��� 5.5 ��m (22.10 �� 0.88 X 5.00 �� 0.33, n= 10), rice-shaped, common. Distribution. Filellum disaggregatum was recorded only for the Chafarinas Islands, Alboran Sea (Pe��a Cantero et al. 1998). Remarks. The trophosomal morphology of Filellum disaggregatum is indistinguishable from that of F. s e rpens, and both species may be differentiated only by their coppiniae. The coppinia of F. disaggregatum is unique among species of Filellum, with pyriform gonothecae, with long necks, irregularly distributed through the coppinia and lacking protective tubes (Pe��a Cantero et al. 1998)., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 12-14, DOI: 10.5281/zenodo.206783, {"references":["Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315.","Pena Cantero, A. L. & Garcia Carrascosa, AM. (2002) The benthic hydroid fauna of the Chafarinas Islands (Alboran Sea, western Mediterranean). Zoologische Verhandelingen, 337, 3 - 180."]}

Filellum conopeum Watson, 2003 Filellum conopeum Watson, 2003: 159 ���160, figs. 9 a���c. Type series. Holotype��� Filellum conopeum Watson, 2003, malinol-mounted microslide, sparse fertile colony on stem of Acryptolaria patagonica (NMV F 91342) (Watson 2003: 159). Type locality. From the holotype, off Macquarie Island (53 ˚ 55.8��S��� 53 ˚ 55.7��S; 159 ˚ 5.5��E��� 159 ˚ 4.7��E), 453 m, (Watson 2003). Description. See Watson (2003: 159���160). Distribution. Filellum conopeum is known only from its original description, off Macquarie Island (53 ˚ 55.8����� 53 ˚ 55.7��S; 159 ˚ 5.5����� 159 ˚ 4.7��E) (Watson 2003). Remarks. Filellum conopeum was described by Watson (2003: 159) as bearing ���Hydrothecae stolonal [���]. Proximal quarter to one third of hydrotheca adnate to stolon, dorsal abcauline wall furrowed by many close, sharpedged ridges with minute ragged frill of perisarc; ridges fading on adnate wall. Adnate wall becoming free at a sharp upward bend, free part cylindrical or weakly expanding from bend to margin, free part straight to broadly curved, walls smooth, occasionally with several regenerations. Margin circular, transverse, with smooth, distinctly everted rim. Perisarc of walls fairly thick, thinning distally. Hydranth with c. 12 tentacles and clavate hypostome���. Trophosome features of F. conopeum are found in other species of the genus, such as F. serratum, F. a n t a rc t i c u m or F. magnificum, so they alone do not allow a proper identification of the species, although F. mangnificum differs from Watson���s species by the distinctly larger hydrothecae, particularly in relation to the diameter of the hydrothecal aperture. As with other species of the genus, truly diagnostic characters of F. conopeum are based on features of the coppiniae: ���Coppinia bud-shaped, c. 1 mm wide and 1 mm high, comprising many tightly packed gonothecae enclosed within a cone of protective nematophorous tubules. Gonotheca flask-shaped (lateral view), base rounded, body expanding a little from base to shoulder then narrowing into a short straight or slightly curved neck tapering to a circular aperture; in transverse view gonothecae polygonal. Nematophorous tubules similar in length, not forked, conjoined just above gonothecae then becoming free, most narrowing distally and inwardly curved to meet above gonotheca; terminal orifice circular. Perisarc of gonothecae and tubes moderately thick; perisarc of tubes somewhat roughened. Planulae enclosed in gonothecae small, spherical.��� (Watson 2003: 160). The general structure of the coppinia of this species resembles that of F. a n t a rc t i c u m and F. magnificum, in which the defensive tubes are situated on the periphery of the mass of gonothecae, like a fence, arching over the gonothecae (in F. magnificum the defensive tubes also arise among the gonothecae). They are different, however, because the gonothecae lack a distal neck in F. antarcticum, and have a short, clearly differentiated distal neck with an everted rim in F. magnificum. In Watson���s species, however, the ���body expanding a little from base to shoulder then narrowing into a short straight or slightly curved neck tapering to a circular aperture���. Although Watson did not give measurements of the gonothecal neck, this seems distinctly longer in general and variable in shape. The cnidome of F. conopeum is unknown and, therefore, no comparison concerning this character can be made., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 11-12, DOI: 10.5281/zenodo.206783

Filellum bouvetensis sp. nov. (Figure 2 A–D) Filellum contortum: Peña Cantero and Gili 2006: 766. Non Lafoea contorta Nutting, 1905: 945, pl. 3, fig. 6, pl. 9, figs. 8–9 [= Filellum contortum (Nutting, 1905)]. Type series. Holotype— Filellum contortum (Nutting, 1905) (PS 65 /028), with coppiniae. Type locality. From the holotype, 54 o 22.49 ’– 54 o 22.54 ’S; 0 3 o 17.58 ’– 03o 17.21 ’E, 122–134m (Peña Cantero & Gili 2006: 765, 766). Material examined. ANTARCTIC OCEAN: Bouvet Island, PS 65 / 28 (25 November 2003, 122– 134m), colonies epibiotic on various hydroids (Antarctoscyphus elongatus, Schizotricha multifurcata, Sertularella sanmatiasensis, Symplectoscyphus sp.); with coppinia. Description. Colonies stolonal, with filiform creeping hydrorhiza. Hydrotheca sessile, arising uprightly and almost completely free from hydrorhiza. When present, abcauline side of adnate part provided with a few striae. Hydrothecal aperture circular and slightly everted, usually with very long rim renovations. Free adcauline hydrothecal part 0.65 – 0.187 mm long. Hydrothecal aperture 0.18–0.25 mm wide. Coppiniae up to 4 mm wide and 2 mm high, some almost completely surrounding the stem of the host hydroid, forming a ring of ca. 4 mm wide. Coppinia composed of firmly adpressed gonothecae with long defensive tubes at periphery, arching over the centre of the gonothecal mass and forming a corbula-like structure. Defensive tubes unforked and with distal aperture. Gonothecae elongated, roughly polygonal in apical view, but with a rather inconstant arrangement in lateral view; distal most part of gonothecae usually free and with irregular shape. Gonothecal aperture circular, situated on a very short, frequently unconspicuous, non-everted distal neck, 0.55–0.6 mm high, 0.1–0.2 mm wide and aperture 0.55– 0.6 mm wide. Cnidome composed of microbasic mastigophores (not seen discharged), 6.5–8.5 X 3 –3.5 µm. Distribution. Known only from Bouvet Island (Peña Cantero & Gili 2006). Remarks. The material from Bouvet Island, here described as Filellum bouvetensis sp. nov., was identified as Filellum contortum (Nutting, 1905) by Peña Cantero and Gili (2006). Filellum bouvetensis sp. nov. is similar to Nutting’s (1905) species in the shape and size of the hydrothecae having a very long free hydrothecal part. However, the new species has larger nematocysts than those found in the lectotype of F. c o nt o r tu m. Furthermore, as indicated above, Nutting’s species was deprived of coppinia, rendering it an insufficiently characterized species to properly assign new material to. Therefore we have decided to consider the abundant, fertile, well-preserved and characterized material from Bouvet suitable for the creation of a new species. Filellum bouvetensis sp. nov. is similar to F. a n t a rc t i c u m in the general shape of the coppinia, both with defensive tubes arching over the gonothecal mass, so that they form a sort of a corbula. Moreover, in dorsal view, the gonothecal aperture of F. bouvetensis sp. nov. is similar to that of F. a n t a rc t i c u m by the presence of a perisarc ring. However, in Hartlaub’s species the gonothecae lack any sort of neck and are distally homogeneous, being completely laterally adpressed. Both species also differ in the size and general proportions of the hydrothecae. Filellum bouvetensis sp. nov. also resembles F. conopeum Watson, 2003 in the structure of the coppinia with defensive tubes curving over the gonothecae. However, Watson (2003) described the coppinia as bud-shaped, ca. 1 mm wide and 1 mm high, whereas in F. bouvetensis they are corbula-shaped and up to 4 mm wide and 2 mm high. Anyway, both species are clearly distinguishable by the size and relative proportions of the hydrothecae. Here a new name is proposed for the Bouvet specimens, which can be clearly distinguished from the remaining known species of Filellum by the size of the nematocysts and hydrothecae, the structure of the coppinia and the shape and size of the gonothecae. Etymology. The name bouvetensis refers to Bouvet, the name of the remote Southern Ocean island where this species was found.

Filellum nitidum Watson, 2005 Filellum nitidum Watson, 2005: 504 ���505, figs. 3 c���e. Type series. Holotype��� Filellum nitidum Watson, 2005, fertile colony on stem of Gymnangium sp. (NMV F 101650). Paratype��� Filellum nitidum Watson, 2005, fertile colony on basal stems of additional colonies of Gymnangium sp. (WAM Z 29955) (Watson 2005: 504). Type locality. From the holotype, Australia, Middle Island, 25 m (Watson 2005: 504). Description. See Watson (2005: 504). Distribution. Filellum nitidum is only known from its original description, from Australia, Middle Island (Watson 2005). Remarks. Filellum nitidum was described by Watson (2005: 504) as bearing ���Hydrothecae sessile [���] adnate part of base fairly short, free part long, perisarc of abcauline wall faintly wrinkled; perisarc of free wall rather thick, marginal rim slightly everted and thickened��� and ���hydrothecae with several regenerations along length���. Surprisingly, in spite of the original description and the figure 3 C, where wrinkled hydrothecae are shown, Watson (2005: 505) also stated that the ���absence of marked abcauline rugosity and absence of a distinctly everted rim��� would make the trophosome of the species similar to that of F. antarcticum. This seems to be an inapt description, since F. antarcticum has distinctly marked striae on the upper side of the adnate portion of the hydrothecae (Pe��a Cantero et al. 2004 b). On the other hand, as also remarked by Watson (2005), both species have clearly different gonothecae. Other morphological features listed by Watson (2005) do not distinguish this species from the others of the genus Filellum. Therefore, we believe that reliable diagnosis of F. nitidum must be based on coppiniae characters, described as ���surface of coppinia a mosaic covered by a thin pellicle of tissue perforated by small openings flush with surface; hydrothecae protruding above surface together with large inverted funnel-shaped tubules, some tubules bifurcated, apices of others closed��� and the sectional view of ���numerous rather haphazardly arranged elongate gonothecae��� (Watson 2005: 504���505). The figure illustrating the coppinia of this species (Watson 2005: 503, figs. 3 D���E) shows protective tubes among the gonothecae; this, together with the different shape of the gonothecae, make the species distinct from F. magnificum, for which the defensive tubes are mainly situated on the periphery of the mass of gonothecae, like a fence, arching over the gonothecae. There is no information on the cnidome of the species, and no further comparison concerning this character can be made., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on page 15, DOI: 10.5281/zenodo.206783, {"references":["Watson, J. E. (2005) Hydroids of the Archipelago of the Recherche and Esperance, Western Australia: Annotated list, redescription of species and description of new species. In: Weills, F. E., Walker, D. I. & Kendrick, G. A. (Eds), The marine flora and fauna of Esperance, Western Australia. Western Australian Museum, Perth, pp. 495 - 612.","Pena Cantero, A. L., Svoboda, A. & Vervoort, W. (2004 b) Antarctic hydroids (Cnidaria, Hydrozoa) of the families Campanulinidae, Lafoeidae and Campanulariidae from recent Antarctic expeditions with R. V. Polarstern, with the description of a new species. Journal of Natural History, 38, 2269 - 2303."]}

Filellum antarcticum (Hartlaub, 1904) Lafoea antarctica Hartlaub, 1904: 11, pl. 2, fig. 2; Vanh��ffen 1910: 311 ���312, figs. 31 a���c. Reticularia antarctica: Totton 1930: 160 ���161, fig. 17. Filellum antarcticum: Millard 1964: 10; 1975: 177, figs. 58g ���h; 1978: 171, 172, 192; Pe��a Cantero et al. 1998: 300; 2004 b: 2283���2287, figs. 2 c���f, 3, 4; Vervoort and Watson 2003: 57 ���58; Pe��a Cantero 2008: 453; 2010: 764���765, figs. 3 a���c. Type series. The type specimen of Filellum antarcticum is lost (Pe��a Cantero et al. 2004 b: 2285). Neotype��� Lafoea antarctica Vanh��ffen, 1910; specimens on Symplectocyphus sp. (ZMB Cni 14222) (Pe��a Cantero et al. 2004 b: 2285��� 2286). Type locality. From the original description of Lafoea antarctica Hartlaub, 1904, it would be ���S. Y. Belgica ��� Expedition, Antarctic, Schwabber VII (70 �� 23 ���S; 82 �� 47 ���W, ca. 500 m, 8 October 1898), specimens on bryozoan (Hartlaub 1904: 5). From neotype designation it is ���Deutschen Southpolar Expedition���, Gauss Station, 65 �� 21 ���S; 86 ��06���E, 385 m (Pe��a Cantero et al. 2004 b: 2285). Material studied. ANTARCTIC: German Polarstern expedition���ANT II- 4 Stn 310, Ronne Ice Shelf, Weddell Sea (76 ��52.0���S; 50 �� 40.4 ���W, 252 m, 20 January 1984), ���on Billardia subrufa, with coppinia��� (RMNH-Coel. 30782); ANT II- 4 Stn 341, Ronne Ice Shelf, Weddell Sea (76 �� 39.2 ���S; 52 ��09.0���W, 297 m, 26 January 1984), ���on B. subrufa and Stegella lobata, with coppinia��� (RMNH-Coel. 30783); ANT II- 4 Stn 438, Larsen Ice Shelf, Antarctic Peninsula, Weddell Sea (67 ��09.7���S; 54 �� 21.4 ���W, 423 m, 7 February 1984), ���on B. subrufa, with coppinia��� (RMNH- Coel. 30785); ANT II- 4 Stn 474, Ronne Ice Shelf, Weddell Sea (76 �� 56.7 ���S; 49 ��44.0���W, 220 m, 14 February 1984), ���on B. subrufa, with coppinia��� (RMNH-Coel. 30788);?ANT II- 4 Stn 524, Cape Norvegia, Princess Martha Coast, Weddell Sea (71 �� 23.9 ���S; 13 �� 58.8 ���W, 325 m, 25 February 1984), ���on Halecium sp., with coppinia��� (RMNH-Coel. 30790); ANT VIII- 5 Stn 16-399, Riiser-Larsen Ice Shelf, Princess Martha Coast, Weddell Sea (72.86 ��S; 19.30 ��W, 380���390 m, 30 December 1989), ���Bottom: large rocks���, ���on B. subrufa, with coppinia��� (RMNHCoel. 30794); ANT VIII- 5 Stn 16-405, Ronne Ice Shelf, Weddell Sea, (76.52 ��S; 52.63 ��W, 330 m, 7 January 1990), ���Bottom: mud���, ���on B. subrufa,? Opercularella belgicae, and axis of gorgonian, with coppinia��� (RMNH-Coel. 30796); ANT VIII- 5 Stn 16-456, Cape Norvegia, Princess Martha Coast, Weddell Sea (71.25 ��S; 12.01 ��W, 200 m, 26 January 1990), ���on B. subrufa, with coppinia��� (RMNH-Coel. 30816); ANT VIII- 5 Stn 16-459, Cape Norvegia, Princess Martha Coast, Weddell Sea (70.96 ��S; 11.19 ��W, 350���380 m, 28 January 1990), ���on B. subrufa, with coppinia��� (RMNH-Coel. 30799); ANT VIII- 5 Stn 16-486, Ronne Ice Shelf, Weddell Sea (76.50 ��S; 52.15 ��W, 330���340 m, 17 February 1990), ���on B. subrufa, with coppinia��� (RMNH-Coel. 30803); ANT IX- 3 Stn 135, McDonald Ice Rumples, Caird Coast, Weddell Sea (75 �� 27.9 ���S; 26 �� 45.4 ���W, 221 m, 4 February 1991), ���on B. subrufa, with coppinia��� (RMNH-Coel. 30805). Deutsche S��dpolar Expedition���Gauss Station (65 �� 21 ���S; 86 ��06���E, 385 m), Museum f��r Naturkunde der Humboldt-Universit��t zu Berlin (neotype, ZMB Cni 14222) (Pe��a Cantero et al., 2004 b: 2284��� 2285). Spanish Bentart 95 Expedition���Stn 3 A, south of Livingston Island (62 �� 37.7018 ���S; 60 �� 22.8167 'W, 92 m, 17 January 1995), on polychaete tube, with coppinia. Spanish Bentart 2003 Expedition���Stn 5 A, south of Peter I Island (68 �� 56 ��� 37 ��������� 68 �� 56 ��� 43 ������S; 90 �� 35 ��� 50 ��������� 90 �� 35 ��� 19 ������W, 124 m, 4 February 2003), on Stegella lobata and the ascidian Cnemidocarpa verrucosa, with coppinia. Description of neotype. (Complementing the description by Pe��a Cantero et al. 2004 b: 2285): Colony stolonal, growing on monosiphonic stems of Symplectoscyphus sp., stolons creeping on substratum, hydrorhiza ca. 0.72���0.80 mm wide. Hydrothecae sessile, sock-shaped, arising with no definite pattern from hydrorhiza, varying in form, from about half of hydrotheca adnate to substratum to some completely free. Hydrothecae irregularly curved in diverse patterns, emerging from substrate plane at various angles. Adnate portion of hydrothecae tubular or maximally slightly flattened, 0.24���0.35 mm long, with numerous external transverse ridges on exposed side; free part cylindrical, smooth, 0.30���0.35 mm long; total adcauline wall 0.59���0.65 mm long, abcauline wall 0.55���0.63 mm long; hydrothecae slightly widening distally, margin even and smooth, with up to four renovations, slightly flaring; hydrothecal aperture circular, 0.104���0.130 mm wide. Hydranths not seen. Gonothecae arranged in coppinia, ca. 2.4 mm wide. Gonothecae closely set, juxtaposed, their limits almost distinct in dorsal view, general shape tubular. Gonothecae 0.48 mm high, 0.11 mm wide, distally truncated with unraised circular aperture, 0.055 mm wide, rim even, not flared, without renovations. Protective tubes present, long, hollow, either forked or unforked, with distal circular aperture. Protective tubes usually curved towards center of gonothecal mass and merged to each other over gonothecae, forming a nest or corbula acting as an incubatory chamber. In one case, disk-shaped coppinia with defensive tubes forming two fused and disk-shaped structures, protecting gonothecae and eggs; in another case, coppinia as a semicircular structure. Eggs remaining inside gonothecae, immersed in a tissue mass presumed to be nourishing. Cnidome composed of two types of nematocysts, small (5.5��� 6.5 X 2.6 ��m) and large ones (10.4���12.4 X 4.2���5.2 ��m). Distribution. Filellum antarcticum seems to be ���a shelf species��� (Pe��a Cantero et al. 1998: 300), being recorded from 13.5 m (Millard 1964, 1975) to 423 m (Pe��a Cantero et al. 2004 b). The species has always been recorded epibiotic, on hydroids (Vanh��ffen 1910; Totton 1930; Pe��a Cantero et al. 2004 b; Pe��a Cantero 2010), gorgonians (Pe��a Cantero et al. 2004 b), bryozoans (Vanh��ffen 1910; Millard 1964, 1975), cidarid spines (Vanh��ffen 1910), polychaete tubes (Pe��a Cantero 2008), and ascidians (Pe��a Cantero 2010). Coppinia found in austral summer (Pe��a Cantero et al. 2004 b: 2297). It is known from South Africa (Millard 1964, 1975) and Antarctica, in the Davis Sea (Vanh��ffen 1910), Ross Sea (Totton 1930), Weddell Sea (Pe��a Cantero et al. 2004 b), the South Shetland Islands (Pe��a Cantero 2008), and off Peter I Island, in the Bellingshausen Sea (Pe��a Cantero 2010). Remarks. Filellum antarcticum was originally described as Lafoea antarctica by Hartlaub (1904) who, according to Pe��a Cantero et al. (1998: 300), diagnosed it as ���sessile hydrothecae, with a short portion adnate to the substratum, followed by a long part directed upwards (this part being longer than in Filellum serpens), and by the presence of numerous renovations of the hydrothecal rim���. However, these features are not sufficient to characterize a species of the genus. The first solution in adequately describing Filellum antarcticum would be to study the type specimen. However, the type material of Lafoea antarctica Hartlaub, 1904 is reportedly lost (Pe��a Cantero et al. 2004 b: 2285). Therefore, Pe��a Cantero et al. (2004 b) proposed the material described by Vanh��ffen (1910) as the neotype of the species, since this specimen was the basis for the first description of the coppinia. Vanh��ffen���s (1910) material was diagnosed by Pe��a Cantero et al. (2004 b: 2286) as having ���several coppiniae [���] of varied shape [���] all have in common a set of gonothecae surrounded by a fence of forked or unforked, defensive tubes provided with a distal circular aperture���. Pe��a Cantero et al. (2004 b: 2286) emphasize that ���this variability seems related to the nature of the substratum where the colony was growing��� and when ���there is enough surface for attachment of the stolonal tubes [���] the coppinia is like a nest with the gonothecae surrounded by a fence of defensive tubes which arch over the gonothecae, whereas when there is not so much space, [���] the coppinia has a disk or semicircular shape���. This plasticity is corroborated by several authors (Totton 1930: 160; Millard 1975: 177) and calls attention to the difficulty in identifying boundaries of species in the genus Filellum. Concerning the trophosome, as many other species congeners (viz. F. magnificum, F. nitidum, F. serratum), F. antarcticum has striations on the outer part of the adnate wall and, therefore, the morphology of the coppinia is essential for the identification of the species. Filellum antarcticum has characteristic coppiniae with tubular gonothecae without distal neck and with bent protective tubes at the periphery of the coppinia, forming a structure that calls to mind a corbula. Differently, the gonothecae of F. magnificum are provided with a distal neck and the protective tubes may be either at the periphery or arise amongst gonothecae in the central area of the coppinia. Also, in F. nitidum, the coppinia has the surface covered by a thin, perforated, tissue layer; in addition, the hydrothecae of F. nitidum are spread on the coppinia together with the thin inverted, occasionally bifid, protective tubes. Finally, Pe��a Cantero et al. (2004 b: 2286���2287) remarked that the differences between F. antarcticum and F. serratum are as follows: ���the distally closed defensive tubes arise from amongst the gonothecae and curve over them forming a protective structure like a canopy��� in F. serratum, whilst ���the gonothecae are surrounded by a fence of distally open defensive tubes usually bent towards the centre of the gonothecal mass, constituting a nest or corbula��� in F. antarcticum. Additionally, the gonothecae of F. serratum are ���bottle-shaped, provided with a short and everted distal neck bearing the aperture, and are slightly shorter and wider���. The similarities in the trophosome of F. antarcticum and F. serratum led Pe��a Cantero et al. (2004 b) to disregard the previous records of Filellum antarcticum not based on fertile specimens., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 8-9, DOI: 10.5281/zenodo.206783, {"references":["Hartlaub, C. (1904) Hydroiden. Resultats du Voyage du S. Y. Belgica. Zoologie, 7, 1 - 19.","Vanhoffen, E. (1910) Die Hydroiden der deutschen Sudpolar-Expedition 1901 - 1903. Deutsche Sudpolar-Expedition 1901 - 1903, 3, 266 - 340.","Totton, A. K. (1930) Coelenterata. Part V. Hydroida. British Antarctic (\" Terra Nova \") Expedition, 1910. Natural History Report, Zoology, 5, 131 - 252.","Millard, N. A. H. (1964) The Hydrozoa of the south and west coasts of South Africa. Part II. The Lafoeidae, Syntheciidae and Sertulariidae. Annals of the South African Museum, 48, 1 - 56.","Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315.","Vervoort, W. & Watson, J. E. (2003) The marine fauna of New Zealand: Leptothecata (Cnidaria: Hydrozoa) (Thecate Hydroids). National Institute of Water and Atmospheric Research (NIWA), 119, 1 - 538.","Pena Cantero, A. L. (2008) Benthic hydroids (Cnidaria: Hydrozoa) from the Spanish Antarctic expedition Bentart 95. Polar Biology, 31, 451 - 464.","Pena Cantero, A. L., Svoboda, A. & Vervoort, W. (2004 b) Antarctic hydroids (Cnidaria, Hydrozoa) of the families Campanulinidae, Lafoeidae and Campanulariidae from recent Antarctic expeditions with R. V. Polarstern, with the description of a new species. Journal of Natural History, 38, 2269 - 2303.","Millard, N. A. H. (1975) Monograph on the Hydroida of Southern Africa. Annals of the South African Museum, 68, 1 - 513.","Pena Cantero, A. L. (2010) Benthic hydroids (Cnidaria: Hydrozoa) from Peter I Island (Southern Ocean, Antarctica). Polar Biology, 33, 761 - 773."]}

Filellum serpens (Hassall, 1848) Sertularia arcta Dalyell, 1847: 224 ���226, pl. 42, figs. 1���15. Campanularia serpens Hassall, 1848: 2223; Hassall and Coppin, 1852: 163, pl. 21, fig. 4. Capsularia serpens: Gray 1848: 151. Coppinia mirabilis Hassall, 1848: 2223. Reticularia immersa Thomson, 1853: 443, pl. 16 a, figs. 2���3. Filellum serpens: Hincks 1868: 214 ���215, pl. 41, figs. 4 a���b (synonym of Reticularia immersa Thomson, 1853); Fraser 1944: 215 ��� 216, pl. 44, figs. 198 a���d; Naumov 1960: 281, figs. 47, 170, 171; Naumov 1969: 303, figs. 47, 170, 171; Vervoort 1972: 49 ��� 50; Cornelius 1975: 378 ���381, fig. 2; Pe��a Cantero et al. 1998: 302 ���304. ? Filellum tubiforme Schydlowsky, 1902: 168, pl. 3, fig. 29. Lafoea serpens: Vanh��ffen 1910: 311. Grammaria serpens: Vervoort 1946: 194 ���196, fig. 82. Filellum serpens serpens: Naumov 1969: 303, figs. 170���171. ? Filellum serpens tubiforme: Naumov 1969: 303, fig. 172. Reticularia serpens: Rees and Thursfield 1965: 87 ���88. [More complete synonymy in Pe��a Cantero et al. 1998: 302���303] Type series. Holotype������Dublin, 1842, infertile colony on herbarium specimen of Abietinaria abietina (Linnaeus, 1758), coll. A.H. Hassall, 1973.10.8.4; mentioned Gray, 1848: 151 ��� (Cornelius 1975: 379). Type locality. From the holotype, Dublin, Ireland (Cornelius 1975: 379). Description after literature data. (Trophosome after Pe��a Cantero & Garc��a Carrascosa 1995: 20, translated; coppinia after Hassall 1848: 2223): Colony with barely visible stolonal hydrorhiza. Hydrothecae arising from hydrorhiza; about half of hydrotheca adnate to substratum. Adnate portion of hydrothecae 0.26���0.28 mm long; free part 0.10���0.22 mm long. Hydrotheca bearing circular aperture, 0,08��� 0.10 mm wide, rim even, slightly flared, without renovations. Coppinia������Polypidom parasitic, massive, hirsute; polype cells elongated, tubular, often curved, arising at irregular distances (and generally at the angles of junction) out of a cellular basis, the apertures of the cells or spaces of which are often themselves covered in by a lid perforated by a small tubular orifice���. Distribution. Eurybathic species, living as an epibiont mainly other species of hydroids (Pe��a Cantero et al. 1998). It has been recorded around the world, but records are mainly based on infertile material. According to records based on fertile material, the species seems to have a temperate-cold boreal distribution (cf. Pe��a Cantero et al. 1998). Remarks. Following the general decision concerning the generic name Filellum after application of case by Cornelius and Calder (1986), the ICZN (Opinion 1485, ICZN 1988) also ruled in favor of the conservation of the specific name serpens Hassall, 1848, as published in the binomen Campanularia serpens Hassall, 1848, the type species of the genus Filellum. The identity of the species Campanularia intertexta Couch, 1844 was questioned by Cornelius and Calder (1986), because the specimens analysed by Couch (1844) were growing on Lafoea dumosa and Sertularella polyzonias, the latter being a typical substrate for Filellum serpens. The authors consider that it is possible that the type material of Campanularia intertexta Couch, 1844, includes F. serpens, besides L. dumosa and Orthopyxis integra, as proposed by Cornelius (1982). Hassall (1848) described the trophosome and gonosome of F. s e r p e n s as two different species, Campanularia serpens and Coppinia mirabilis, respectively. The first author who united them was Hincks (1868), and the name ���coppiniae��� since then refers to the peculiar reproductive structures found in some genera of the family Lafoeidae. Coppinia mirabilis Hassall, 1848 is considered a junior synonym of Sertularia arcta Dalyell, 1847, as evidenced by Hincks (1868) and reviewed by Cornelius and Calder (1986). Reticularia immersa Thomson, 1853 is considered a synonym of Filellum serpens (Hassall, 1848) and its conspecificity was corroborated by Cornelius (1975) through the analyses of its type material. Although data on the coppinia of the species is sketchy (e.g., ���Coppinia of typical structure���, Naumov 1969: 303; ������coppiniae typical of the sub-family���, Cornelius 1975: 380) the trophosome of F. s e r p e n s may be differentiated from that of most species of the genus (viz. F. antarcticum, F. magnificum, F. nitidum, and F. serratum), but it is similar to those of F. adhaerens and F. disaggregatum, with more than half of the hydrothecae adnate to the substrate. However, the hydrothecae and hydrorhiza of F. adhaerens form a mesh laying on the substrate, which is also found in F. parasiticum. Concerning the sexual structures, F. disaggregatum has a completely different coppinia, whereas F. serpens, F. parasiticum and F. adhaerens have similar coppiniae, with gonothecae having juxtaposed walls and a distal neck, and curved protective tubes arising among the gonothecae. However, there are some differences in the gonothecal morphology, polygonal in F. adhaerens (Nutting 1901: 178, pl. 21, fig. 4) and F. parasiticum, and fusiform in F. serpens (Dalyell 1847, pl. 42, fig. 6, also in Pe��a Cantero et al. 1998: 304). The taxonomic status of F. s e r p e n s is dubious, because of the imprecise morphological descriptions associated with the species. It is likely that a group of different species may be confused and incorrectly assigned to the species under discussion. However, since F. serpens is the type species of the genus, it is by definition the senior synonym, although other congeners appear to be much more clearly described. Therefore, dismissing the name serpens would demand major nomenclatural acts at the genus level that we would prefer to avoid at this time., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on pages 18-19, DOI: 10.5281/zenodo.206783, {"references":["Hassall, A. H. (1848) Definitions of three new British zoophytes. Zoologist, 6, 2223.","Dalyell, J. G. (1847) Rare and remarkable animals of Scotland, represented from living subjects: with practical observations on their nature. John Van Voorst, London, 270 pp.","Hassall, A. H. & Coppin, J. (1852) Descriptions of three species of marine zoophytes. Transactions of the Microscopical Society of London, 3, 160 - 163.","Gray, J. E. (1848) List of the specimens of British animals in the collection of the British Museum. Part I. Centroniae or radiated animals. British Museum, London, 173 pp.","Thomson, W. (1853) Notes on some British zoophytes. Annals and Magazine of Natural History, 2 nd series, 11, 442 - 444.","Hincks, T. (1868) A history of the British hydroid zoophytes. John van Voorst, London, 338 pp.","Fraser, C. M. (1944) Hydroids of the Atlantic coast of North America. The University of Toronto Press, Toronto, 447 pp.","Naumov, D. V. (1960) Gidroidi i gidromedusy morskikh, solonovatovodnykh i presnovodnykh basseinov SSSR. Opredeleteli pro faune SSSR, 70, 1 - 626.","Naumov, D. V. (1969) Hydroids and Hydromedusae of the USSR. Israel Program for Scientific Translation, Jerusalem, 660 pp.","Vervoort, W. (1972) Hydroids from Theta, Vema and Yelcho cruises of the Lamont-Doherty geological observatory. Zoologische Verhandelingen, 120, 1 - 247.","Cornelius, P. F. S. (1975) A revision of the species of Lafoeidae and Haleciidae (Coelenterata: Hydroida) recorded from Britain and nearby seas. The Bulletin of the British Museum (Natural History), Zoology, 28, 373 - 426.","Pena Cantero, A. L., Garcia Carrascosa, A. M. & Vervoort, W. (1998) On the species of Filellum Hincks, 1868 (Cnidaria: Hydrozoa) with the description of a new species. Journal of Natural History, 32, 297 - 315.","Schydlowsky, A. (1902) Materialy po faune gidroidov arkticheskikh morei. I. Gidroidy Belogo morya u beregov Solovetskikh ostrovov (Data on the hydroid fauna of the Arctic Seas I. Hydroids of the White Sea near the Coasts of the Solovetskie Islands). Trudy Obshchestva Ispytatelei Prirody pri Imperatorskom Khar'kovskom Universitete, 36, 3 - 268.","Vanhoffen, E. (1910) Die Hydroiden der deutschen Sudpolar-Expedition 1901 - 1903. Deutsche Sudpolar-Expedition 1901 - 1903, 3, 266 - 340.","Vervoort, W. (1946) Fauna van Nederland. Hydrozoa (CI). A. Hydropolypen. A. W. Sijthoff's Uitgeversmaatschappij N. V., Leiden, 336 pp.","Rees, W. J. & Thursfield, S. (1965) The hydroid collections of James Ritchie. Proceedings of the Royal Society of Edinburgh, 69, 34 - 220.","Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I, editio decima, reformata, Holmiae, Laurentii Salvii, 823 pp.","Pena Cantero, A. L. & Garcia Carrascosa, A. M. (1995) Hidrozoos bentonicos de la Campana Antartida 8611. Publicaciones Especiales del Instituto Espanol de Oceanografia, 19, 1 - 148.","Cornelius, P. F. S. & Calder, D. (1986) Filellum serpens (Hassall, 1848) (Cnidaria, Hydrozoa): proposed conservation of both generic and specific names. A. N. (S.) 2508. Bulletin of Zoological Nomenclature, 43, 335 - 341.","Couch, R. Q. (1844) A Cornish fauna; being a compendium of the natural history of the county. Part III containing the zoophytes and calcareous corallines. Royal Institution of Cornwall, Truro, 162 pp.","Cornelius, P. F. S. (1982) Hydroids and medusae of the family Campanulariidae recorded from the eastern North Atlantic, with a world synopsis of genera. The Bulletin of the British Museum (Natural History), Zoology, 42, 37 - 148.","Nutting, C. C. (1901) Papers from the Harriman Alaska Expedition. XXI. The hydroids. Proceedings of the Washington Academy of Sciences, 3, 157 - 216."]}

Filellum plicatum (Hartlaub, 1904) Lafoea plicata Hartlaub, 1904: 12, pl. 2, fig. 3. Type series. Holotype������the material ��� is lost��� (Pe��a Cantero et al. 2004 b: 2287). Type locality. From the original description, ���Schwabber VIII���, ���N�� 923 ���, Antarctic, 70 ��00���S; 80 �� 48 ���W, 55 m, 18 October 1898 (Hartlaub 1904: 12). Remarks. Lafoea plicata was originally described by Hartlaub (1904), who remarked on the resemblance of his species with F. serratum based on the adnate part of the hydrotheca being surrounded by a thin membrane provided with fine pleats resembling indentations in lateral view (see also Pe��a Cantero et al. 2004 b: 2287). The type material was infertile and is reportedly lost. According to Pe��a Cantero et al. (2004 b: 2287) ���the size of the hydrothecae in L. plicata is closer to that of the material here considered as F. antarcticum (e.g. 104���111 mm diameter at the aperture)���. However, based on the lack of diagnostic characters of the trophosome, we decided to consider this taxon as species inquirenda., Published as part of Marques, Antonio C., Pe��a, ��lvaro L., Miranda, Tha��s P. & Migotto, Alvaro E., 2011, Revision of the genus Filellum Hincks, 1868 (Lafoeidae, Leptothecata, Hydrozoa), pp. 1-28 in Zootaxa 3129 on page 24, DOI: 10.5281/zenodo.206783, {"references":["Hartlaub, C. (1904) Hydroiden. Resultats du Voyage du S. Y. Belgica. Zoologie, 7, 1 - 19.","Pena Cantero, A. L., Svoboda, A. & Vervoort, W. (2004 b) Antarctic hydroids (Cnidaria, Hydrozoa) of the families Campanulinidae, Lafoeidae and Campanulariidae from recent Antarctic expeditions with R. V. Polarstern, with the description of a new species. Journal of Natural History, 38, 2269 - 2303."]}

Key for the identification of the fertile specimens of Filellum 1 Abcauline wall of the adnate part of the hydrotheca with transverse striae........................................ 2 - Abcauline wall of the adnate part of the hydrotheca without transverse striae....................................... 7 2 Defensive tubes of the coppinia forming a protective structure over the gonothecae................................. 3 - Defensive tubes of the coppinia not forming a protective structure over the gonothecae........... F. nitidum Watson, 2005 3 Defensive tubes arising amongst the gonothecae. Gonotheca with short distal neck with everted rim.................... 4 - Defensive tubes exclusively on the periphery of the gonothecal mass and arching over the gonothecae................. 5 4 Defensive tubes forming a shield-shaped structure (canopy)............................... F. serratum (Clarke, 1879) - Defensive tubes simply arching over gonothecae............................. F. magnificum Peña Cantero et al., 2004 b 5 Gonothecae without distal neck................................................. F. antarcticum (Hartlaub, 1904) - Gonothecae with distal neck............................................................................. 6 6 Coppinia bud-shaped. Gonothecae flask-shaped. Hydrothecal aperture 140–168 µm. Hydrothecal free part 320–420 µm.................................................................................... F. conopeum Watson, 2003 - Coppinia corbula-shaped. Gonothecae irregular. Adnate hydrothecal part short or absent. Hydrothecal aperture 180–250 µm. Hydrothecal free part 650–1875 µm..................................................... F. bouvetensis sp. nov. 7 Gonothecae without juxtaposed walls. Defensive tubes absent................ F. disaggregatum Peña Cantero et al., 1998 - Gonothecae with juxtaposed walls. Defensive tubes present.................................................... 8 8 Hydrothecae and hydrorhiza forming a continuous layer on the substrate.......................................... 9 - Hydrothecae and hydrorhiza spaced and distinct from each other............................ F. serpens (Hassall, 1848) 9 Gonothecae fusiform............................................................. F. adhaerens (Nutting, 1901) - Gonothecae polygonal....................................................... F. parasiticum (Antsulevich, 1987)

Clytia elsaeoswaldae Stechow, 1914 (Figures 2 & 3; Table 2) Clytia elseae-oswaldae Stechow, 1914: 125 Clytia hemisphaerica: Migotto, 1996: 82 ���84 Clytia cf. gracilis sp. 2: Lindner, 2000: 46 Material examined. Syntypes: colony with gonangia, on alga. (Charlotte Amalie, St. Thomas, U.S. Virgin Islands) [Zoologische Staatssamlung, M��nchen, microslides 20041517 (part of colony with six hydranths and one gonangium), 20041518 (part of colony with nine hydranths and six gonangia) and 20041519 (part of colony with 39 hydranths and ~ 25 gonangia)]. Collected by E. Stechow and E. Oswald, 18 March 1912. Additional material. (1) Colony with gonangia on posts of a pier, growing on sponges and barnacles (TEBAR, S��o Sebasti��o, SP, Brazil; coordinates: 23 �� 47.97 ���S; 45 �� 22.98 ���W; water depth: 5���20 m) [MZUSP 962, 963]; collected by A. Lindner and A.E. Migotto, 22 October 1999. (2) Colony with gonangia on iron structure, growing on sponge, Hypnea sp. and Eudendrium carneum Clarke, 1882 (Hydrozoa: Eudendridae) (Saco do Sombrio, Ilhabela, SP, Brazil; coordinates: 23 �� 53.77 ���S; 45 �� 14.74 ���W; water depth: ~ 5 m) [MZUSP 959, 960, 961]; collected by A. Lindner, 26 September 1999. (3) Colony with gonangia growing on Sargassum sp. and 39 one to 21 days-old medusae liberated from colony and reared in the laboratory (Saco Grande, S��o Sebasti��o, SP, Brazil; coordinates: 23 �� 49.72 ���S; 045�� 25.52 ���W; water depth: 1���3 m) [colony: MZUSP 937, 938, 940, 941, USNM 1078726; medusae: MZUSP 939, 942, 943, 945, 946, 947, 948, 949, 950, 951, 952, 953, 954, 955, 956, 957, 958]; colony collected by A. Lindner, 19 February 1999. (4) Colony with gonangia on Sargassum sp., Hypnea sp. and bryozoan (Praia de Bareque��aba, west shore, S��o Sebasti��o, SP, Brazil; water depth: ~ 0.5 m) [MZUSP 967, USNM 1078725]; collected by L.P. de Andrade, 0 5 April 2000. (5) Colony with gonangia on Sargassum sp. and ~ 100 newly liberated medusae (Praia de Bareque��aba, east shore, S��o Sebasti��o, SP, Brazil; water depth: ~1.0 m) [colony: MZUSP 965, USNM 1078727; medusae: MZUSP 966]; colony collected by L.P. de Andrade, 0 4 April 2000. (6) Colony with gonangia on Sargassum sp. and on dead hydrocaulus of Aglaophenia latecarinata (Hydrozoa: Aglaopheniidae) (Praia do Guaec��, west shore, S��o Sebasti��o, SP, Brazil; water depth: ~ 1 m) [MZUSP 964]; collected by L.P. de Andrade, 0 1 February 2000. (7) Colony on Sargassum sp. (S��o Sebasti��o, SP, Brazil) [USNM 1078728]; collected by O.M.P. de Oliveira. Diagnosis. Clytia elsaeoswaldae is distinguished from its congeners and other campanulariids by the combination of the following characters: polyp: stems monosiphonic or, less common, polysiphonic, branching dichotomously. Hydrothecal cusps usually inclined to one side. Gonothecae only on hydrorhiza, with smooth, slightly undulated walls. B-type microbasic mastigophores 15���18 ��m long in vivo and 13���16 ��m long in formalin. adult medusa: bell 3.6���5.5 mm in marginal diameter, up to 16 tentacles and at least 21 statocysts. A��� type microbasic mastigophores, ~ 11 ��m long, forming a row at the level of the circular canal (no character or combination of characters so far observed is diagnostic for young medusae). Description. Hydroid . Colonies stolonal or with erect stems branching dichotomously up to 4 times (Figure 2). Erect stems up to 5.6 mm high. Hydrorhiza frequently growing on pedicels, sometimes forming incipient polysiphonic stems (Figure 2 F) or more developed polysiphonic stems up to 23 mm high and bearing up to 60 hydranths. Pedicels usually smooth, slightly curved, with 5���15 proximal and 2���16 distal annuli; median region of pedicels sometimes annulated. Distal end of pedicels bearing one hydrotheca. Internodes with a upward-curved apophysis supporting the next internode or the distal pedicel (Figure 2 A,B). Hydrothecae cylindrical, 368���753 ��m long and 255���408 ��m wide at margin, with thin perisarc and walls almost parallel in lateral view, except a basal rounded portion (Figure 2). Hydrothecal diaphragm thin, transverse, near base of hydrotheca; basal chamber 16���78 ��m long and 78���165 ��m wide at diaphragm. Hydrothecal margin with 9���14 cusps (39���86 ��m high); cusps usually inclined to right side when hydrotheca is seen in lateral view under microscope (Figure 2). Hydrotheca width:length ratio: 0.48���0.84 (Table 2). Gonothecae smooth, with undulated walls, 630���1019 ��m long and 255���463 ��m in maximum diameter, with a constriction (216���294 ��m in diameter) under the truncated distal margin (205���345 ��m in diameter). Gonothecae growing only from hydrorhiza (Figure 2); on Sargassum, small ���aggregations��� of gonothecae common (Figure 2 H). Gonothecal pedicels absent or having up to 4 annulli. Up to six medusae in each gonangium. Gonotheca width:length ratio: 0.38���0.50. Hydranth column, on average, 5 times as long as broad when extended [493 �� 83 ��m (252���600, N= 47) long and 103 �� 14 ��m (72���144, N= 47)] wide; hypostome pedunculated, 120���264 ��m wide, spherical in oral and lateral views (Figure 2 G); 20���34 filiform, amphicoronate tentacles, 420���1080 ��m long; rings of A-type nematocysts every 15���20 ��m along the tentacles. Coenosarc whitish. A- and B-type microbasic mastigophores on hydranths and along cenosarc, only A-type on tentacles. A-type microbasic mastigophores 8.1 �� 0.6 ��m [7.0��� 9.5, N= 46] long and 2.0 �� 0.1 ��m [2.0��� 2.5, N= 46] wide in vivo and 7.1 �� 0.2 ��m [6.5 ���8.0, N= 40] long and 2.0 �� 0.2 ��m [1.5���2.5, N= 40] wide in formalin. Discharged capsules 6.8 �� 0.4 ��m [6.0��� 7.5, N= 36] long and 2.0 �� 0.2 ��m [1.5���2.5, N= 20] wide in vivo, with a proximal armature 6.8 �� 0.4 ��m [6.0���8.0, N= 36] long. B-type microbasic mastigophores 16.4 �� 0.6 ��m [15.0���18.0, N= 40] long and 3.8 �� 0.3 ��m [3.0���4.0, N= 40] wide in vivo and 14.6 �� 0.5 ��m [13.0���16.0, N= 180] long and 3.2 �� 0.3 ��m [3.0���4.0, N= 150] wide in formalin. Discharged capsules 14.4 �� 0.5 ��m [14.0���15.0, N= 17] long and 3.2 �� 0.4 ��m [3.0��� 4.5, N= 16] wide in vivo, with a proximal armature 18.1 �� 1.4 ��m [15.0���20.0, N= 16] long. Newly released medusa. Umbrella hemispherical, with ring canal, four perradial canals; four perradial bulbs with tentacles; two to four interradial developing bulbs; D-type nematocysts on the exumbrella; 4���8 adradial statocysts, each with one statolith. Some medusae also liberated with two interradial tentacles developing in opposite quadrants, i.e., with six tentacles (four perradial and the two incipient interradial tentacles). Gonads on median region of radial canals, 31���39 ��m wide and 55���78 ��m long in newly liberated medusae. Manubrium quadrate, ca. 0.4���0.5 height of bell; lips smooth with A-type nematocysts. Velum broad. Tentacles hollow, with A- and C-type nematocysts and a terminal nematocyst cluster with only C-type nematocysts. Adult medusa. Umbrella saucer-shaped, 3.6���5.5 mm in diameter. Bell margin with up to 16 bulbs with tentacles and at least 21 statocysts (Table 1; Figure 3). Gonads oval when fully developed, on distal 1 / 3 of radial canals, approximately at the same level of velum margin in oral and aboral views (Figure 3 B). Manubrium short, cruciform, with four undulated lips in oral view; lips with A-type nematocysts. Marginal tentacles hollow, long, with Atype and atrichous isorhiza nematocysts. C-type, D-type, and terminal nematocyst cluster absent. A-type nematocysts, larger than those in the tentacles, forming a row at the level of the circular canal, perpendicular to the canal (Figure 3 C). Length of A-type nematocysts in a 21 day old medusa: on tentacles, 7.9 �� 0.3 ��m [7.0��� 8.5, N= 20]; on circular canal, 10.7 �� 0.5 ��m [10.0��� 11.5, N= 20]. Bell transparent; gonads, manubrium, bulbs and tentacles assuming the colour of food. Female medusae not seen. Distribution. Known from S��o Sebasti��o and Ilhabela, SP, Brazil (Migotto 1996) and the type-locality, St. Thomas, U.S. Virgin Islands (Stechow 1914), but reports of C. hemisphaerica and C. gracilis from other western Atlantic localities may also have been based on the species. Biological notes. Clytia elsaeoswaldae is one of the most abundant shallow-water hydroids occurring off the coast of S��o Sebasti��o and Ilhabela, southeastern Brazil (see Migotto 1996, as C. hemisphaerica). Some colonies growing on Sargassum have several hundred hydranths and gonangia, and a few hours after collecting even small fragments of such colonies were observed to release several hundred medusae in the laboratory. Only small size differences were observed among colonies of C. elsaeoswaldae from southeastern Brazil (Table 2), and all fertile colonies had gonangia growing exclusively on the hydrorhiza. Colonies from Brazil are also similar to the syntypes from off the U.S. Virgin Islands, which also have gonothecae exclusively on the hydrorhiza (as noticed by Stechow 1914: 125) but differ in having smaller gonothecae (Table 2). This difference, however, is only in size, and the shape of the gonothecae is similar, with a mean width:length ratio of 0.4 (Table 2). The specimens from Brazil and the Virgin Islands also differ slightly in that the hydrothecal cusps are less tilted and their tips more acute in the latter specimens, but otherwise all specimens are similar in both shape and size (Table 2). Because of permanent preservation in microslides, cnidome of syntypes could not be determined and DNA sequences could not be obtained, and we regard the specimens from southeastern Brazil conspecific to those from the Virgin Islands on the basis of perisarc morphology. Medusae of C. elsaeoswaldae are released with small gonads, eight statocysts, and either four or six tentacles. After 2 days of culture at 22���24 ��C, medusae attained up to 8 tentacles, and after six days additional statocysts started to develop between perradial and adradial marginal bulbs. D-type nematocysts on the exumbrella disappeared during medusa development, as well as C-types in the tentacles. Maximum diameter of the umbrella (i.e., 5.5 mm) was reached after 12 days, and no additional growth was observed until the culture ended 21 days after liberation [maximum diameter of the umbrella, on average: 4.8 mm (N= 5) on the 14 th day; 4.9 mm (N= 10) on the 16 th day; 4.6 mm (N= 10) on the 20 th]. New statocysts continue to develop after medusae reached their maximum size: up to 16 and 21 statocysts in 12 and 20 days old medusae, respectively. Maximum number of fully developed tentacles varied from 12 to 16. Phylogenetics. The combined maximum likelihood analysis indicates that Clytia elsaeoswaldae and other Clytia gracilis -like species do not form a monophyletic group (Figure 4). Similarly, none of the four genes support a monophyletic C. gracilis clade when analyzed individually or in mitochondrial (16 S and COI) and nuclear (18 S and calmodulin) partitions (results not shown). The phylogenetic results show that the here re-described C. elsaeoswaldae is most closely related to a clade including C. hemisphaerica and two C. gracilis -like species (species A and B in Figure 4)., Published as part of Lindner, Alberto, Govindarajan, Annette F. & Migotto, Alvaro E., 2011, Cryptic species, life cycles, and the phylogeny of Clytia (Cnidaria: Hydrozoa: Campanulariidae), pp. 23-36 in Zootaxa 2980 on pages 26-30, DOI: 10.5281/zenodo.208048, {"references":["Stechow, E. (1914) Zur Kenntnis neuer oder seltener Hydroidpolypen, meist Campanulariden, aus Amerika und Norwegen. Zoologischer Anzeiger, 45, 120 - 136.","Migotto, A. E. (1996) Benthic shallow-water hydroids (Cnidaria, Hydrozoa) of the coast of Sao Sebastiao, Brazil, including a checklist of Brazilian hydroids. Zoologische Verhandelingen, 306, 1 - 125.","Lindner, A. (2000) Redescricao e ciclo de vida de Clytia gracilis e Clytia linearis (Cnidaria, Hydrozoa, Campanulariidae). M. S. thesis, Universidade de Sao Paulo, Sao Paulo, Brazil, 82 pp.","Clarke, S. F. (1882) New and interesting hydroids from Chesapeake Bay. Memoirs of the Boston Society of Natural History, 3 (4), 135 - 142."]}

Beania americana n. sp. (Figs 14, 15, 24; Table 2) Beania hirtissima: Marcus 1937: 62, pl. 14, fig. 31; 1941: 19, fig. 12; Osburn 1940: 397; Lagaaij 1963: 180, pl. 8, fig. 4; Maturo 1966: 579, fig. 17; Winston 1982: 131, fig. 56; Vieira et al. 2008: 17 (checklist). Non Diachoris hirtissima Heller, 1867. Material examined. Holotype: MZUSP 415. Paratype: MZUSP 416, Additional material: Marcus collection (19), Beania hirtissima E. Marcus det. on rocks; MZUSP 417���428; VMNH 760.00, Beania hirtissima, Walton Rocks, South Hutchinson Island, St Lucie County, Florida, USA, coll. J. E. Winston, 28.xi, 1998, intertidal beach rock; VMNH 2612.00, Beania hirtissima, Juno Ledge, off Palm Beach, Florida, USA, coll. Littler and Little, 18.vii. 1999, drift dive, 20 m, on alga; VMNH 3150.01, Beania hirtissima, North Beach, Fort Pierce, St Lucie County, Florida, USA, coll. J. E. Winston 21.vi. 2003, beach drift alga, with B. mirabilis (= B. mirabilissima). Description. Colony reticulate, semierect, grayish-brown in color, often heavily coated with mud and debris. Zooids with lightly calcified curving lateral and basal walls and a membranous frontal surface; zooids joined by 6 tubular processes and attached loosely to substratum by tubular radicles. Frontal surface constricted into two parts, a smaller oval opercular area surrounded by a halo of 8���10 long, straight stout spines, and a larger oval proximal region with 9���13 pairs of spines arched over the frontal membrane plus some projecting outward from the lateral walls. Tubular radicles extend from the proximal portion of zooids. Avicularia and ovicells absent. Remarks. Beania hirtissima has been considered to have a worldwide geographical distribution in shallow, warm-temperate to tropical seas (Winston 1982; Cook 1985; Hayward & McKinney 2002). Hayward and McKinney (2002) examined the type of Beania hirtissima (Heller, 1867) and designated its lectotype, a specimen with vicarious avicularia, autozooids 0.8 ���1.0 mm long with seven stout spines surrounding the operculum like a halo (three distal and two on each side), and only five erect spines on each lateral margin of the frontal membrane, each with a shorter spine at its base, curving medially over the frontal membrane. They remarked that records of Beania hirtissima should ���be reexamined in light of the characteristics of the lectotype ���. They examined one such taxon with reticulate colonies that had often been attributed to Beania hirtissima f. or var. cylindrica, and designated it as Beania cylindrica (Hincks, 1886). Cook (1985) showed that B. hirtissima may represent a species complex and, more recently, Tilbrook (2006) noted that some reports previously attributed to this species represent undescribed species. Beania cylindrica (Hincks, 1886) has bigger autozooids (1.0 x 0.25 mm), a terminal operculum orientated perpendicular to the frontal membrane, only six distolateral spines and another 8���10 pairs of slightly incurved lateral spines. In the erect tufts of Beania cylindrica, each autozooid is linked to six neighbors by tubular connections, three proximal and closely spaced and the other three mid-basal. In contrast, colonies of B. hirtissima are unilaminar sheets with autozooids closely spaced and linked by a ring of six short connecting tubes around their proximal ends. Marcus (1937) characterized colonies of Beania hirtissima from Santos (S��o Paulo state, Brazil) as having a distal halo of 10 long spines and 12 pairs of shorter lateral spines; additionally, he described some basal spines, sometimes bi- or trifurcated. We consider that Beania hirtissima sensu Marcus (1937, 1941) is a previously undescribed species, named herein as Beania americana n. sp. Although B. americana is similar to the lectotype of B. hirtissima in the general habit of the colony and characters of the autozooid, it is clearly distinguishable, particularly by the number and size of spines and size of zooids (see Table 2). Shared features of the Brazilian and Western Atlantic colonies also led us to reassign some specimens previously recorded as Beania hirtissima (viz. Osburn 1940; Lagaaij 1963; Maturo 1966; Winston 1982) to B. americana. measurements from Hayward and McKinney (2002). MA, distance between midpoints of adjacent apertures. Mean (St Dev) 0.503 (0.054) 0.508 (0.030) 0.521 (0.036) 0.499 (0.065) 0.549 (0.092) Min���Max 0.385���0.615 0.462���0.569 0.440 ���0.600 0.394 ���0.600 0.377���0.792 Beania americana and B. cylindrica are similar in the number of spines arched over the frontal membrane, but B. americana differs in having smaller zooids (reaching 0.723 x 0.338 mm), a halo of 8���10 stout distal spines, and autozooids linked to six neighbors by long, well-spaced tubular connections. The other similar Brazilian species, described herein as Beania correiae n. sp., is readily distinguished by the number of marginal spines arched over the frontal membrane. Biological notes. Beania americana was found on rocky shores from the intertidal to 20 m deep on rocks, macroalgae including red algae and Sargassum sp., and other bryozoans [viz. Amathia spp. and Anguinella palmata]; it was also present on shallow soft bottoms (6���10 m) on rocks and shell fragments (Table 6). In Florida (USA), it was found year-round, epibiotic on hydroids and encrusting beach rocks partially buried in sand (Winston 1982). Colonies are also utilized as a substratum by other cheilostome bryozoans. Colonies coated with sediment can hardly be seen in situ. Distribution. Atlantic. Brazil: Paran�� (Marcus 1941; present study), S��o Paulo (Marcus 1937; present study) and Alagoas (present study); Porto Rico and Bermuda (Osburn 1940); USA: North Carolina, Georgia (Maturo 1966), Florida (Maturo 1966; Winston 1982; present study) and Gulf of Mexico (Lagaaij 1963)., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Shallow-water species of Beania Johnston, 1840 (Bryozoa, Cheilostomata) from the tropical and subtropical Western Atlantic, pp. 1-20 in Zootaxa 2550 on pages 9-11, DOI: 10.5281/zenodo.196806, {"references":["Marcus, E. (1937) Bryozoarios marinhos brasileiros, 1. Boletim da Faculdade de Filosofia, Ciencias e Letras da Universidade de Sao Paulo, Zoologia, 1, 5 - 224.","Osburn, R. C. (1940) Bryozoa of Porto Rico with a resume of the West Indian bryozoan fauna. Scientific Survey of Porto Rico and the Virgin Island, 16, 321 - 486.","Lagaaij, R. (1963) New additions to the bryozoan fauna of the Gulf of Mexico. Publication of the Institute of Marine Science (University of Texas), 9, 162 - 236.","Maturo, F. J. S. (1966) Bryozoa of the south-east coast of the United States: Bugulidae and Beaniidae (Cheilostomata: Anasca). Bulletin of Marine Science, 16, 556 - 583.","Winston, J. E. (1982) Marine bryozoans (Ectoprocta) of the Indian River area, Florida. Bulletin of the American Museum of Natural History, 173, 99 - 176.","Vieira, L. M., Migotto, A. E. & Winston, J. E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810, 1 - 39.","Heller, C. (1867) Die Bryozoen des adriatischen Meeres. Verhandlungen der Zoologisch-botanischen Gesellschaft in Wien, 17, 77 - 136.","Cook, P. L. (1985) Bryozoa from Ghana. Tervuren, Belgie Koninklijk Museum voor Midden-Afrika. Zoologische Weteschappen, 238, 1 - 315.","Hayward, P. J. & McKinney, F. K. (2002) Northern Adriatic Bryozoa from the vicinity of Rovinj, Croatia. Bulletin of the American Museum of Natural History, 270, 1 - 139.","Hincks, T. (1886) The Polyzoa of the Adriatic, I. Annals and Magazine of Natural History, ser. 5, 17, 254 - 271.","Tilbrook, K. J. (2006) Cheilostomatous Bryozoa from the Solomon Islands. (Studies in Biodiversity, 3). Santa Barbara Museum of Natural History Monographs, 4, 1 - 386.","Marcus, E. (1941) Briozoarios marinhos do litoral Paranaense. Arquivos do Museu Paranaense, 1, 7 - 36."]}

Beania mirabilissima n. sp. (Figs 12, 13, 22, 23; Table 1) Beania mirabilis: Osburn 1914: 189; 1940: 398; Marcus 1937: 60, pl. 12, fig. 29; Winston 1982: 133, fig. 45; 1986: 5; López-Gappa 2001: 73, figs 1, 2, 9; Vieira et al. 2008: 18 (checklist); Ramalho et al. 2010: 503, fig. 3. Non Beania mirabilis Johnston, 1840: 272, fig 1, 2; 1847: 372, figs 69–70; Busk 1852 b: 32, pl. 24, figs 4–5; Hincks 1880: 96, pl. 4, figs 8–10; Hayward & Ryland 1998: 244; Hayward & McKinney 2002: 26, figs 10 C,D. Material examined. Holotype: MZUSP 405. Paratype: MZUSP 406. Additional material: MZUSP 407–414; UFAL 0 49, Maceió, Alagoas state, 27.iv. 2006, on hydroids; VMNH 2992.00, Beania mirabilis, Walton Rocks, South Hutchinson Island, St Lucie County, Florida, USA, coll. J. E. Winston 15.vii. 2002, intertidal on hydroid stolon; VMNH 3116.00, Beania mirabilis, Fort Pierce Inlet, USA, coll. J. E. Winston, 14.vii. 2003, beach drift on aluminum can, with Aetea sp. Description. Colony uniserial, branched, lightly attached to substratum, translucent white in color. Zooids oblong, suberect, with an erect boat-shaped portion and a long and slender connective tube. Erect portion inclined, convex basally with a flat, expanded membranous frontal surface. Connective tube proximal, about 0.05 mm in diameter, joined to basal part of zooid. Orifice about 0.06 mm long and 0.12 mm wide. Two pairs of short distal spines around operculum and 5–8 (often 6) pairs of small spines overarching the frontal membrane; sometimes the proximal spines are slightly longer than the distal ones. Tubular basal radicles present in all zooids, between connective tubes. Avicularia and ovicells absent. Polypides with 18–20 tentacles. Remarks. Johnston (1840) described a new British genus and species, Beania mirabilis, found loosely attached to shell. The species has been characterized by several authors as having slender zooids with 5–11 pairs of lateral spines in addition to the four distal spines. It has been considered to be a species with a worldwide distribution in warm and temperate waters (e.g. Winston 1982; Ramalho et al. 2010). However, Cook (1985) suggested that B. mirabilis may represent a species complex in the warm waters of the Atlantic and Indo-Pacific oceans. Later, López-Gappa (2001) observed slight differences (zooid length, number and position of spines) between British and Patagonian specimens of putative B. mirabilis, but did not introduce a new species name for the southwestern Atlantic material. Recently, Ramalho et al. (2010) remarked that specimens from Rio de Janeiro (Brazil) had fewer spines, but considered that the number of spines of those specimens fell within the range for the species. Tilbrook et al. (2001) redescribed Johnston’s material of Beania mirabilis, characterized by zooids about 0.70 mm long, connective tubes 1.5 times longer than the boat-shaped portion, two pairs of oral spines and 9– 10 pairs of lateral spines overarching the frontal membrane; neanic colonies identified by Johnston from the type locality have only 2–6 spines. Tilbrook et al. (2001) also noted that several specimens previously attributed to B. mirabilis represent undescribed species. Later, Hayward and McKinney (2002) described and figured specimens of Beania mirabilis from the Adriatic; they are similar to Johnston’s specimens. These descriptions have made it possible to compare precisely the Brazilian colonies with material from North Atlantic and Adriatic. Both English and Adriatic ephebic specimens differ from those found in shallowwater localities of the West Atlantic in the presence of 10 pairs of short, short-spaced lateral spines, and deeper (about 0.2 mm), wider zooids (about 0.28 mm wide) connected by longer proximal tubes. Beania mirabilis sensu strict is probably restricted to the northeastern Atlantic (Johnston 1840; Busk 1852 b; Hincks 1880), Red Sea (Dumont 1981) and Mediterranean-Adriatic (Gautier 1962; Zabala & Malquer 1988; Hayward & McKinney 2002). The material from Vanuatu previously identified as Beania cf. mirabilis (Tilbrook et al. 2001) was recently redescribed by Tilbrook (2006) as Beania lagenula, also characterized by a large gap between marginal and oral spines, but differing from those found in the western Atlantic in the shape of the zooid and the presence of 6–8 pairs of marginal spines that overlap each other in the midline. Beania mirabilissima n. sp., which name alludes to the morphological likeness with Beania mirabilis Johnston, 1840, is characterized by oblong, shallower zooids (about 0.17 mm deep) joined by a shorter proximal connective tube (about 1.1 times longer than the boat-shaped portion), 5–8 pairs of lateral spines overarching the frontal membrane but not reaching its midpoint, and a large gap between the marginal and oral spines. Two small colonies from deep water off Brazil deposited at the MZUSP (MZUSP 0 40, 088) are readily distinguished from B. mirabilissima by their smaller, deeper zooids (about 0.21 mm deep) connected by a longer proximal connective tube (ca 1.5–2.5 times longer than the boat-shaped portion) and 8–10 shorterspaced pairs of lateral spines with slightly curved distal tips; these resemble European B. mirabilis and the eastern Atlantic colonies described by Cook (1985) but have slightly shorter zooids (about 0.4 mm). We suggest molecular genetics and further morphometric studies will resolve the identity and morphological variations of different populations of B. mirabilis with 8–11 spines in Atlantic waters. Biological notes. Colonies of B. mirabilissima were found on rocky shores from the intertidal to 20 m deep, epiphytic on algae including Halimeda sp. and Sargassum sp., and epizoitic on bryozoans (e.g. Amathia sp.) and hydroids (Table 6). It was previously recorded in the West Atlantic by several authors (Osburn 1914; Marcus 1937; Winston 1982; Ramalho et al. 2010) as Beania mirabilis, growing on rocks, shells, other bryozoans and hydroid stems. It was previously recorded from Caribbean at 9–36 m deep (Osburn 1914, 1940). According to Winston (1982), embryos probably develop inside the zooids, and as she remarked, there is still no information available on its reproduction. Specimens collected in São Paulo state, Brazil, have about 18–20 tentacles while African specimens of B. mirabilis collected on shells have 12–14 (Cook 1985) and probably represent an undescribed species. Distribution. Western Atlantic: Argentina (López-Gappa 2001); Brazil: Santa Catarina (present study), Paraná (present study), São Paulo (Marcus 1937; present study), Rio de Janeiro (Ramalho et al. 2010) and Alagoas (Vieira et al. 2007; present study); USA: Tortugas Island (Osburn 1940) and Florida (Winston 1982; present study).

Beania metrii n. sp. (Figs 26���29; Table 2) Material examined. Holotype: MZUSP 401. Description. Colony reticulate, loosely encrusting, white in color, surface often heavily coated with mud and debris. Zooids close spaced and slightly overlapping distally; lightly calcified curving lateral and basal walls with some spines; zooids joined by 6 very short tubular processes. Frontal surface oval, membranous, broader proximally than distally, with the operculum in a narrower distal semicircular area surrounded by a halo of 8 spines; 8���10 pairs of small, often straight, lateral spines usually directed over the frontal membrane; lateral walls also with several straight spines, 6���16 on each side, projecting outwards. Avicularia and ovicells absent. Remarks. Beania metrii n. sp., named after Rafael Metri (Universidade Estadual do Centro-Oeste, Brazil) who collected the specimen, is very distinct from the other reticulate species described here in its zooidal dimensions (bigger than other species), the number of frontal spines and very short connective tubes joining autozooids. The distance between the midpoints of adjacent apertures of Beania metrii are similar to those described for Adriatic specimens of both B. cylindrica and B. hirtissima, but B. metrii is clearly distinguished by the number of spines in the distal halo. Beania hirtissima also differs in having bigger zooids and only five pairs of lateral spines. Biological notes. The species occurred in a bank of calcareous algae (Halimeda sp.) at Arvoredo Island (Santa Catarina state) between algal nodules in water ca 6���20 m deep (Table 6). Distribution. Atlantic. Brazil: Santa Catarina (present study)., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Shallow-water species of Beania Johnston, 1840 (Bryozoa, Cheilostomata) from the tropical and subtropical Western Atlantic, pp. 1-20 in Zootaxa 2550 on page 14, DOI: 10.5281/zenodo.196806

Beania australis Busk, 1852 b (Fig 2 ���4, 18; Table 1) Beania australis Busk, 1852 b: 32, pl. 16, fig 1���3; Marcus 1937: 63, pl. 13, fig. 32; Marcus 1949: 2; Vieira et al. 2008: 17 (checklist). Material examined. MZUSP 0 23, Santos, S��o Paulo state, Brazil, E. Marcus det. (1938) on rock; MZUSP 377���378. Description. Colony uniserial, branched, white in color. Zooids oblong, suberect, with an erect boatshaped portion and a long and slender connective tube. Erect portion obliquely inclined, convex basally, with a flat wholly membranous frontal surface. Connective tube proximal, about 0.271���0.802 mm in length and 0.037���0.043 mm in width, joined to the basal part of zooid; each zooid with 1���3 basal tubes. Two pairs of short distal (oral) spines around operculum and 10���19 opesial spines on each side, overarching frontal membrane, sometimes overlapping. Opesial spines reaching or sometimes exceeding the midline of membrane. Tubular radicles with branched ends present proximal to boat-shaped portion of some zooids. Avicularia absent. Ovicells absent. Remarks. Beania australis was described from Cape Horn by Busk (1852 b), as characterized by autozooids with 18���20 spines on each side of the frontal membrane, reaching to midpoint of the opesia. Marcus (1937) recorded this species in Santos (S��o Paulo state, Brazil) and described zooids with 17���22 pairs of lateral spines. However, specimens identified by Ernst Marcus and deposited in the Museu de Zoologia (MZUSP 023) have only 10���15 pairs. In the Brazilian specimens, the number of opesial spines may increase with the size of the zooid. Hayward and Ryland (1995) described a new species, Beania pectinata, collected on Sargassum sp. from Heron Island (Australia). It resembles B. australis in several aspects ��� dimensions of autozooids, presence of four short distal spines, four basal connective tubes, peri-opesial spines forming a comb-like frontal shield, and lack of avicularia ��� but the 25 pairs of closely aligned lateral spines in B. pectinata are highly distinctive. The frontal membrane of specimens of B. australis from Alagoas state (Brazil) is about 0.5 mm long, whereas, according to a drawing by Marcus (1937) of specimens from Santos (Brazil), the length of the frontal membrane is about 0.85 mm. We examined specimens from Santos in the Museu de Zoologia da Universidade de S��o Paulo (MZUSP 023) identified by Ernst Marcus and found that their length to be ca 0.60 mm, slightly longer than specimens from Alagoas. The beaniid genus Stolonella, established monotypically for S. clausa Hincks, 1883 from the Australian region, resembles B. australis in autozooid appearance (with a comb-like frontal shield and lack of avicularia), but the zooids are borne directly on kenozooids (true stolons). The basal tubular processes in autozooids of Beania australis are similar to those in B. mirabilis, being formed by the slender proximal tube of the zooid rather than a kenozooid, indicating a closer relationship with Beania. Biological notes. The colonies of B. australis studied were collected on rocky shores in shallow water, where it is epiphytic on red algae and epilithic on rocks (Table 6). Some colonies were also found in shrimp bycatch in the south of Alagoas state, at ca 15 m deep on the bryozoan Celleporaria sp. Distribution. Atlantic: Cape Horn and Patagonia (Busk 1852 b); Brazil: S��o Paulo (Marcus 1937; present study), Esp��rito Santo (Marcus 1949) and Alagoas (present study). Beania cupulariensis Osburn, 1914: 190, figs 6���7; 1940: 399; Marcus 1944: 1, figs 1���4; 1955: 291, pl. 4, figs 41���43; Fransen 1986: 80, figs 26 a���d; Winston 1986: 5; Vieira et al. 2008: 17 (checklist). Material examined. MZUSP 0 0 7, 1948, Ilhabela, S��o Paulo state, Brazil, E. Marcus det. on algae; MZUSP 379���381. Description. Colony uniserial, rarely branched, white���translucent in color. Zooids oblong, boat-shaped, higher than wide, with lightly calcified walls; connective tubes short, the distal end of the zooid overlapping the proximal end of the subsequent zooid. Frontal membrane occupying the whole frontal surface. Marginal spines numbering 5���7 on each side, overarching the frontal membrane with only the most proximal pair directed outward; one pair of distal spines. One pair of pedunculate avicularia on distal corners of zooids, with rostrum hooked, mandible hard and curved. Ovicells absent. Remarks. Beania cupulariensis was described from the Tortugas, Florida (USA) by Osburn (1914). This species is characterized by zooids connected by short tubes, the distal ends of zooids overlapping the proximal ends of subsequent zooids, the frontal membrane overarched by 5���7 pairs of spines, with the most proximal pair directed to the outside, and one pair of distal spines. Harmer (1926, p. 419, pl. 28, figs 13���14) recorded two forms of B. cupulariensis. The first form, collected from the Philippines, has broader zooids with 7���9 pairs of uniformly spaced spines directed to the outside. An Indonesia form has longer and narrower zoecia, with 3���4 shorter pairs of marginal spines. These forms probably represent two distinct, undescribed species. The number of tentacles found in Beania cupulariensis is variable: the Brazilian specimens have 26 tentacles (Marcus 1955), but Cook (1985) reported polypides with 20���22 tentacles in material from Ghana; a figured specimen (ibid., fig. 37) also differs in lacking the most proximal pair of spines directed to the outside. It too may represent a different species. Biological notes. The colonies of B. cupulariensis studied by us were collected from rocks in shallow water about 5 m deep (Table 6), where the current flow is strong. Colonies from S��o Paulo state were found on mollusk shells and algae. Vieira et al. (2007) noted the association of this species with the bryozoan Vasignyella ovicellata (Catenicellidae) in northeastern Brazil. Osburn (1914, 1940) found some colonies of B. cupulariensis on dorsal surfaces of Cupuladria sp. from 18���40 m depth while the colonies reported by Marcus (1955) from the Brazilian coast were epiphytic on algae. Distribution. Atlantic. Brazil: S��o Paulo (Marcus 1944, 1955; present study) and Alagoas (Vieira et al. 2007; present study); Porto Rico (Osburn 1940); Cura��ao (Fransen 1985); USA: Tortugas Island (Osburn 1914);? Ghana (Cook 1985)., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Shallow-water species of Beania Johnston, 1840 (Bryozoa, Cheilostomata) from the tropical and subtropical Western Atlantic, pp. 1-20 in Zootaxa 2550 on pages 3-6, DOI: 10.5281/zenodo.196806, {"references":["Busk, G. (1852 b) Catalogue of Marine Polyzoa in the Collection of the British Museum, I. Cheilostomata (Part). Trustees of the British Museum (Natural History), London. 54 p.","Marcus, E. (1937) Bryozoarios marinhos brasileiros, 1. Boletim da Faculdade de Filosofia, Ciencias e Letras da Universidade de Sao Paulo, Zoologia, 1, 5 - 224.","Marcus, E. (1949) Some Bryozoa from the Brazilian coast. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 3 (53), 1 - 33.","Vieira, L. M., Migotto, A. E. & Winston, J. E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810, 1 - 39.","Hayward, P. J. & Ryland, J. S. (1995) Bryozoa from Heron Island, Great Barrier Reef. 2. Memoirs of the Queensland Museum, 38, 533 - 573.","Hincks, T. (1883) Contributions towards a general history of the marine Polyzoa. Part XI. Foreign Cheilostomata (Australia and New Zealand). Annals and Magazine of Natural Histor, ser. 5, 11, 193 - 202.","Osburn, R. C. (1914) The Bryozoa of the Tortugas Islands, Florida. Publication of the Carnegie Institution of Washington, 182, 183 - 222.","Marcus, E. (1944) Beania cupulariensis Osb. (Bryozoa Cheilost.) nova para o Brasil. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 1 (12), 1 - 3.","Fransen, C. H. J. M. (1986) Caribbean Bryozoan: Anasca and Ascophora Imperfecta of the inner bays of Curacao and Bonaire. Studies of Fauna Curacao and other Caribbean Islands, 68 (210), 1 - 19.","Harmer, S. F. (1926) The Polyzoa of the Siboga Expedition, Part 2. Cheilostomata Anasca. Siboga-Expeditie, 28 b, 183 - 501.","Marcus, E. (1955) Notas sobre briozoos marinhos brasileiros. Arquivos do Museu Nacional do Rio de Janeiro, 42, 273 - 341.","Cook, P. L. (1985) Bryozoa from Ghana. Tervuren, Belgie Koninklijk Museum voor Midden-Afrika. Zoologische Weteschappen, 238, 1 - 315.","Vieira, L. M., Gordon, D. P. & Correia, M. D. (2007) First record of a living ditaxiporine catenicellid in the Atlantic, with a description of Vasignyella ovicellata n. sp. (Bryozoa). Zootaxa, 1582, 49 - 58.","Osburn, R. C. (1940) Bryozoa of Porto Rico with a resume of the West Indian bryozoan fauna. Scientific Survey of Porto Rico and the Virgin Island, 16, 321 - 486."]}

Beania correiae n. sp. (Figs 16, 17, 25; Table 2) Material examined. Holotype: MZUSP 402. Paratype: MZUSP 403. Description. Colonies reticulate, semierect, grayish-brown in color, often heavily coated with mud and debris. Zooids with lightly calcified, curving lateral and basal walls and a membranous frontal surface; zooids joined by 6 tubular processes and attached loosely to the substratum by tubular radicles. Frontal surface constricted into two parts, a smaller oval opercular area surrounded by a halo of 8 long, symmetrical stout spines plus 5 long distally directed spines at its base, and a larger oval proximal region with 14���20 pairs of spines (often 18) arched over the frontal membrane. Long spines project outwards from the lateral walls on each side of the frontal membrane; 3 regular clusters of 1���3 spines (often 2): the first clusters placed closest to constriction below the halo, the other 2 placed in distal third and half portion of zooid. Long spines on basal wall; 2���5 spines (often 5) in distal and 4���5 spines (often 5) in proximal basal portion of zooid. Tubular radicles in proximo-lateral portion of zooids. Avicularia and ovicells absent. Remarks. Beania correiae n. sp., named in honour of Dr Monica Dorigo Correia (Universidade Federal de Alagoas, Brazil) who encouraged the first author to study bryozoans, is characterized by eight long stout spines in the distal halo plus five distally directed spines at its base, and often up to 16 pairs of long spines arched over the frontal membrane and overlapping each other across its midline. Colonies of B. correiae resemble B. cylindrica, but the last differs from the former in having bigger zooids and fewer spines around the distal halo and the frontal membrane. Biological notes. Beania correiae grew on rocks found in shrimp bycatch in the south of Alagoas state (Brazil), in water about 15 m deep. It was also found on bryozoans (e.g. Celleporaria spp.) and shells (Table 6). Some colonies were heavily coated with mud and debris, like B. americana and B. metrii n. sp., the other reticulate species described here. Distribution. Atlantic. Brazil: Alagoas (present study)., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Shallow-water species of Beania Johnston, 1840 (Bryozoa, Cheilostomata) from the tropical and subtropical Western Atlantic, pp. 1-20 in Zootaxa 2550 on pages 11-14, DOI: 10.5281/zenodo.196806

Beania metrii n. sp. (Figs 26–29; Table 2) Material examined. Holotype: MZUSP 401. Description. Colony reticulate, loosely encrusting, white in color, surface often heavily coated with mud and debris. Zooids close spaced and slightly overlapping distally; lightly calcified curving lateral and basal walls with some spines; zooids joined by 6 very short tubular processes. Frontal surface oval, membranous, broader proximally than distally, with the operculum in a narrower distal semicircular area surrounded by a halo of 8 spines; 8–10 pairs of small, often straight, lateral spines usually directed over the frontal membrane; lateral walls also with several straight spines, 6–16 on each side, projecting outwards. Avicularia and ovicells absent. Remarks. Beania metrii n. sp., named after Rafael Metri (Universidade Estadual do Centro-Oeste, Brazil) who collected the specimen, is very distinct from the other reticulate species described here in its zooidal dimensions (bigger than other species), the number of frontal spines and very short connective tubes joining autozooids. The distance between the midpoints of adjacent apertures of Beania metrii are similar to those described for Adriatic specimens of both B. cylindrica and B. hirtissima, but B. metrii is clearly distinguished by the number of spines in the distal halo. Beania hirtissima also differs in having bigger zooids and only five pairs of lateral spines. Biological notes. The species occurred in a bank of calcareous algae (Halimeda sp.) at Arvoredo Island (Santa Catarina state) between algal nodules in water ca 6–20 m deep (Table 6). Distribution. Atlantic. Brazil: Santa Catarina (present study).

Beania maxilladentata Ramalho, Muricy & Taylor, 2010 (Fig 9 ���11, 21; Table 1) Beania maxilladentata Ramalho et al., 2010: 504, fig. 4. Material examined. MZUSP 400. Remarks. Beania maxilladentata was recently described from Rio de Janeiro state on bryozoans, shells, serpulids and algae in 3���6 m of water. The species is characterized by small spines that never overarch the frontal membrane and by the presence of a strongly curved avicularian mandible with one or two tooth-shaped accessory structures. Distribution. Atlantic. Brazil: Rio de Janeiro (Ramalho et al. 2010; present study)., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Shallow-water species of Beania Johnston, 1840 (Bryozoa, Cheilostomata) from the tropical and subtropical Western Atlantic, pp. 1-20 in Zootaxa 2550 on page 8, DOI: 10.5281/zenodo.196806, {"references":["Ramalho, L. V., Muricy, G. & Taylor, P. D. (2010) Taxonomy of Beania Johnston, 1840 (Bryozoa, Flustrina) from Arraial do Cabo, Rio de Janeiro state, Brazil. Arquivos do Museu Nacional do Rio de Janeiro, 66, 499 - 508."]}

Beania klugei Cook, 1968 (Figs 7, 8, 20; Table 1) Beania intermedia: Marcus 1937: 61, pl. 13, fig. 30; 1939: 113; 1949: 2; Osburn 1940: 398; Maturo 1966: 579, fig. 18. Non Diachoris intermedia Hincks, 1881. Beania klugei Cook, 1968: 165, figs 2 A,B; Winston 1982: 131, fig. 46; Cook 1985: 119, fig. 4 A; Winston & H��kansson 1986: 18, fig. 39; Fransen 1986: 82, figs 27 A���C; Tilbrook et al. 2001: 46; Vieira et al. 2008: 18 (checklist); Ramalho et al. 2010: 501, fig. 2. Material examined. MZUSP 0 23, Beania intermedia, Santos, S��o Paulo state, Brazil, E. Marcus det. (1938) on rock; MZUSP 0 72, 210���211, 252, 265, 269, 382 ��� 390, 404; UFAL 0 34, Macei��, Alagoas state, 27.iv. 2006, on hydroids; UFAL 0 51, Macei��, Alagoas state, 24.iv. 2006 on bryozoans; VMNH 2569.00, Beania klugei, Spanish Anchor, off West Palm Beach, Florida, USA, coll. Littler and Littler, 15.vii. 1999 on algae; VMNH 3150.01, Beania klugei, North Beach, Fort Pierce, St. Lucie County, Florida, USA, coll. J. E. Winston 21.vi. 2003, on drift alga with Beania hirtissima (= B. americana). Description. Colony uniserial, sometimes branching laterally, attached loosely to the substratum, often with a portion free from the substratum. Zooids transparent yellowish-tan in color, large, boat-shaped, with a flat frontal membrane and rounded abfrontal surface; short connective tubes slender: a proximal tube with zooid budded from the one proximal or proximal-lateral to it, with occasional lateral chains of zooids. Operculum delicately chitinized and barely visible. Marginal spines absent. One pair of pedunculate avicularia on distal ends of zooids. Polypides with a large lophophore and 26 tentacles. Ovicells absent. Remarks. Marcus (1937) used the name Beania intermedia for this species, but Cook (1968) showed that B. intermedia Hincks, 1881 is a spinous species with small zooids, and she introduced the name Beania klugei for populations of a large, uniserial Beania with small paired avicularia and no marginal spines. The species was recently discussed by Ramalho et al. (2010), who noted small differences between Brazilian and Florida (USA) materials. Although there are slight differences in size between Brazilian and Florida colonies described by Winston (1982), specimens of both places are characterized by zooids without spines, similarshaped avicularia, and a lophophore with 26 tentacles. Specimens recorded from the Great Barrier Reef by Hayward and Ryland (1995, p. 542) as B. klugei were redescribed by Tilbrook et al. (2001) as Beania cookae; this species differs from B. klugei by the shape of the avicularia and by the presence of short lateral spines on some zooids. Biological notes. The specimens studied by us were found from the intertidal to 15 m deep on rock surfaces, algae including Sargassum sp., ascidians, bryozoans [viz. Anguinella palmata, Bugula spp., Crisia pseudosolena and Amathia vidovici] and hydroids (Table 6). Some zooids of B. klugei were found with other bryozoans on a shipwreck off the coast of Pernambuco state (Brazil), at 24 m deep. This species was previously recorded from the Caribbean at 10���64 m deep (Osburn 1940). In Florida (USA), it is common on hydroid stems and benthic algae, and on the drift alga Soliera tenera (Winston 1982); a two-zooid colony was also found attached to a vibraculum of a Cupuladria doma colony (Winston & H��kansson 1986). Both African (Cook 1985) and Brazilian specimens have lophophores with 24���26 tentacles. Distribution. Widely distributed in tropical and subtropical waters (Tilbrook et al. 2001). Brazil: Paran�� (present study), S��o Paulo (Marcus 1937; present study), Rio de Janeiro (Marcus 1949; Ramalho et al. 2010; present study), Alagoas (present study), Pernambuco (Marcus 1939; present study) and Cear�� (present study)., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Shallow-water species of Beania Johnston, 1840 (Bryozoa, Cheilostomata) from the tropical and subtropical Western Atlantic, pp. 1-20 in Zootaxa 2550 on pages 6-7, DOI: 10.5281/zenodo.196806, {"references":["Cook, P. L. (1968) Bryozoa (Polyzoa) from the coasts of tropical West Africa. Atlantide Report, 10, 11 - 262.","Marcus, E. (1937) Bryozoarios marinhos brasileiros, 1. Boletim da Faculdade de Filosofia, Ciencias e Letras da Universidade de Sao Paulo, Zoologia, 1, 5 - 224.","Osburn, R. C. (1940) Bryozoa of Porto Rico with a resume of the West Indian bryozoan fauna. Scientific Survey of Porto Rico and the Virgin Island, 16, 321 - 486.","Maturo, F. J. S. (1966) Bryozoa of the south-east coast of the United States: Bugulidae and Beaniidae (Cheilostomata: Anasca). Bulletin of Marine Science, 16, 556 - 583.","Hincks, T. (1881) Contributions towards a general history of the marine Polyzoa. Part VIII. Foreign Cheilostomata (miscellaneous). Annals and Magazine of Natural History, ser. 5, 8, 132 - 136.","Winston, J. E. (1982) Marine bryozoans (Ectoprocta) of the Indian River area, Florida. Bulletin of the American Museum of Natural History, 173, 99 - 176.","Cook, P. L. (1985) Bryozoa from Ghana. Tervuren, Belgie Koninklijk Museum voor Midden-Afrika. Zoologische Weteschappen, 238, 1 - 315.","Winston, J. E. & Hakansson, E. (1986) The interstitial fauna of the Capron Shoals, Florida. American Museum Novitates, 2865, 1 - 98.","Fransen, C. H. J. M. (1986) Caribbean Bryozoan: Anasca and Ascophora Imperfecta of the inner bays of Curacao and Bonaire. Studies of Fauna Curacao and other Caribbean Islands, 68 (210), 1 - 19.","Tilbrook, K. J., Hayward, P. J. & Gordon, D. P. (2001) Cheilostomatous Bryozoa from Vanuatu. Zoological Journal of the Linnean Society, 131, 35 - 109.","Vieira, L. M., Migotto, A. E. & Winston, J. E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810, 1 - 39.","Ramalho, L. V., Muricy, G. & Taylor, P. D. (2010) Taxonomy of Beania Johnston, 1840 (Bryozoa, Flustrina) from Arraial do Cabo, Rio de Janeiro state, Brazil. Arquivos do Museu Nacional do Rio de Janeiro, 66, 499 - 508.","Hayward, P. J. & Ryland, J. S. (1995) Bryozoa from Heron Island, Great Barrier Reef. 2. Memoirs of the Queensland Museum, 38, 533 - 573.","Marcus, E. (1949) Some Bryozoa from the Brazilian coast. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 3 (53), 1 - 33.","Marcus, E. (1939) Bryozoarios marinhos brasileiros, 3. Boletim da Faculdade de Filosofia, Ciencias e Letras da Universidade de Sao Paulo, Zoologia, 3, 111 - 353."]}

Marcusadorea corderoi (Marcus, 1949) n. comb. (Figures 6–11; Table 2) Coleopora corderoi Marcus 1949: 18, figs 23–25; Vieira, Migotto & Winston 2008: 30. Material examined. MZC Hassler Expedition Box 33, 36; Coleopora corderoi Marcus; off North of Rio de Janeiro, Brazil. Description. Colonies encrusting, unilaminar. Autozooids large (1.251 mm long, 0.766 mm wide for MCZ Hassler colonies), irregularly polygonal to rectangular, separated by distinctly raised lateral wall junctions. Frontal shield convex, surface granular with small scattered pores, these becoming obscured with a tuberculated frontal wall in increasing calcification; some larger marginal areolar pores remain visible. Primary orifice bell-shaped, wider proximally, anter arched, U-shaped, separated from broad and shallow poster by barely projecting condyles (mean 0.140 mm long, 0.178 mm wide for MCZ Hassler colonies). No oral spines. Peristome tubular, entire. Secondary orifice mostly rounded to oblong, with U- to V-shaped proximal edge (0.464 mm long, 0.401 mm wide for MCZ Hassler colonies). Suboral avicularium occasionally present, positioned laterally on inner side of peristome, in V-shaped portion, with curved and cuspidate mandible; none was present on the two colony fragments measured. Vicarious avicularia lacking. Ooecia not observed, but Marcus (1949) described them as prominent, globular, with porous surface, opening inside peristome. Ancestrula not observed. Remarks. When introducing the new species Coleopora corderoi, Marcus (1949) distinguished it from other Coleopora species by features of the peristome, absence of tubes on frontal wall, presence of avicularia with curved cuspidate mandible, and prominent ooecium opening inside the peristome. He described very large zooids, reaching 1.4–1.8 mm in length and 0.7 mm in width, while his drawing shows zooids about 1.2 mm long (see Marcus 1949: fig. 23). It was not possible to reexamine Marcus’ specimens as the holotype of C. corderoi is missing, along with other Brazilian specimens identified by Ernst Marcus that were deposited in the Museum of Natural History (UK) (M. Spencer Jones pers. comm., 2009) and Museu de Zoologia of Universidade de São Paulo (Brazil). Some of the colonies of M. corderoi collected north of Rio de Janeiro by the Hassler Expedition have smaller zooids (MCZ Hassler Box 33), more like those of Marcusadorea jamaicensis specimens, but colonies with large zooids (MCZ Hassler Box 36) as described by Marcus (1949) were also found. The Brazilian colonies found in MCZ collection (Figures 1–6) lack avicularia and ovicells. Other authors have already described further species with large zooids, raised peristomes, irregularly porous frontal shields, and raised lateral walls adjacent to the areolar pores, which we consider as the diagnostic features of the new genus Marcusadorea. The species M. tubulosa (Canu & Bassler, 1928) differs from M. corderoi by having smaller zooids and suboral avicularia with curved spatulate mandibles; colonies in which suboral avicularia are lacking might be difficult to identify; however, colonies of Marcusadorea without avicularia may be distinguished by features of the peristome and orifice. Marcusadorea corderoi has an almost rounded secondary orifice at the end of the peristome in most zooids. In the other three species of the genus, the peristome of most zooids is incomplete; there is a characteristically proximally incomplete Ushaped slit in Marcusadorea jamaicensis n. sp., a broader proximal V-shaped to U-shaped gap in M. tubulosa, and a quadrangular to subtriangular one in M. efatensis (Tilbrook, 2006). Distribution. Western Atlantic – Brazil: Rio de Janeiro (present study) and Espírito Santo states (Vieira et al. 2008)., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Marcusadorea, a new genus of lepralioid bryozoan from warm waters, pp. 57-68 in Zootaxa 2348 (1) on pages 61-63, DOI: 10.11646/zootaxa.2348.1.4, http://zenodo.org/record/5307125, {"references":["Marcus, E. (1949) Some Bryozoa from the Brazilian coast. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 3 (53), 1 - 33.","Vieira, L. M., Migotto, A. E. & Winston, J. E. (2008) Synopsis and annotated checklist of Recent marine Bryozoa from Brazil. Zootaxa, 1810, 1 - 39.","Canu, F. & Bassler, R. S. (1928) Fossil and Recent Bryozoa of the Gulf of Mexico region. Proceedings of the United States National Museum, 72, 1 - 199.","Tilbrook, K. J. (2006) Cheilostomatous Bryozoa from the Solomon Islands. (Studies in Biodiversity Number 3). Santa Barbara Museum of Natural History Monographs, 4, 1 - 386."]}

Marcusadorea tubulosa (Canu & Bassler, 1928) n. comb. (Figures 12–15, 18–19; Table 3) Holoporella (?) tubulosa Canu & Bassler 1928: 147, pl. 24, figs 1–6, text-fig. 33a. Coleopora tubulosa: Winston 1986: 16; Montoya-Cadavid, Flórez-Romero & Winston 2007: 169. Non Coleopora tubulosa Canu & Bassler 1925. Material examined. USNM 7532; Holoporella (?) tubulosa (Cotypes); Gulf of Mexico and Cuba. UFAL 0006; Marcusadorea tubulosa (Canu & Bassler); Coral Reefs of Pajuçara, Maceió, Alagoas State, Brazil, 9°40'49" S, 35°43'05" W, 0–3 m, coll. M.D. Correia, 20 September 2006, 1 encrusting colony. UFAL 0016; Marcusadorea tubulosa (Canu & Bassler); Coral Reefs of Amores, Maceió, Alagoas State, Brazil, 9°40'40" S, 35°42'10" W, 0–5 m, coll. M.D. Correia, 31 November 2006, 1 encrusting colony. UFAL 0021; 22 September 2006, 1 encrusting colony. MZUSP 0175; 0–3 m, coll. V. R. Cedro, 17 October 2005, 1 encrusting colony. MZUSP 0176; 0–5 m, coll. L.M. Vieira and M.D. Correia, 02 April 2007, 1 encrusting colony. MZUSP 0177; 1 encrusting colony (gold-coated specimen for SEM). MZUFBA 145; Coleopora corderoi Marcus; V.E.S. Almeida det., Costa do Dendê, Bahia State, Brazil, 13°31'25" S, 38°48'38" W, 45 m, coll. J.M.L. Dominguez and R. Silva, March 2002, 1 encrusting colony. MZUFBA 146; 13°44'37" S, 38°48'40" W, 47.7 m, coll. J.M.L. Dominguez and R. Silva, March 2002, 1 erect colony. MZUFBA 147; 13°59'48" S, 38°55'34" W, 16.6 m, coll. J.M.L. Dominguez and R. Silva, 2002, 2 encrusting colonies. Redescription. Colonies encrusting, unilaminar, or erect cylindrical with bifurcated branches, about 1.5– 3.0 mm diameter, disposed in longitudinal irregular series around the axis, reddish-brown in color. Autozooids large (0.911 mm long, 0.684 mm wide for Brazilian colonies), irregularly rectangular to oblong, separated by shallow grooves and distinctly raised lateral walls. Frontal shield convex, granular, irregularly porous, with numerous marginal areolar pores. Primary orifice widest in distal third, with a poster comprising two-thirds of a full circle, demarcated from the deep and broadly U-shaped anter by very short and blunt condyles. Primary orifice partly subcircular, bell-shaped, U-shaped anter and broader poster. No oral spines. Peristome tubular, proximally incomplete, forming a U-shaped pseudosinus. Secondary orifice rounded (0.226 mm long, 0.233 mm wide in Brazilian colonies), proximally incomplete. Large proximal suboral avicularium (0.233 mm long, 0.094 mm wide in Brazilian colonies) occasionally present, set obliquely on the side of the peristome, with the rostrum directed towards the proximal end of the pseudosimus, with curved spatulate mandible; rostrum oblong with rounded corners, slightly curving to outside, crossbar complete. Vicarious avicularia lacking. Ooecium prominent (0.383 mm long, 0.489 mm wide in Brazilian colonies), subglobular, partly incorporated into frontal shield of distal zooid, with some medium-sized pores scattered across granular-tubercular surface; ovicell opening inside peristome. Ancestrula not observed. Remarks. Marcusadorea tubulosa is characterized by large zooids, approximately 1.20 mm in length in Caribbean specimens (Canu & Bassler 1928), a thin-walled proximally incomplete peristome surrounding the orifice, and a large proximal suboral avicularium placed laterally on the rim of the peristome. Canu & Bassler (1928) noted that the budded zooids of M. tubulosa are distinct from those of other species of Holoporella Waters, 1909 (= Celleporaria Lamouroux, 1821). They noted similarities with the peristome of Coleopora species, which lack attachment marks around the operculum. They considered their material as an undetermined genus owing to the absence of ovicells and the growth mode of the colonies in which frontal budding is lacking occur. However, their plate clearly illustrates ovicells (Canu & Bassler 1928: pl. 26, fig. 6), the presence of which we confirmed by the examination of a paratype of Holoporella (?) tubulosa deposited in the United States National Museum (USNM 7532). The examination of this material also revealed that in some zooids there is a proximal suboral avicularium placed laterally on the peristome, occupying one third of it, and globular ovicells opening inside the peristome, as in the Brazilian material figured herein (Figures 13– 14). Prior to the original description of the genus Coleopora, Canu & Bassler (1925) described the Austrian fossil species Coleopora tubulosa, later regarded by David & Pouyet (1974) as a junior synonym of Coleopora insignis (Reuss, 1874). Teuchopora sp. recorded by Banta & Carson (1977) from Costa Rica resembles Marcusadorea tubulosa in general dimensions, frontal-shield formation, orifice, and the hemispherical aperture of ovicell opening into the base of the peristome, but it apparently lacks avicularia. Among the type material of M. tubulosa were found erect colonies, comprising cylindrical bifurcated branches, figured by Canu & Bassler (1928: pl. 24, figs 1, 4–5). Erect cylindrical colonies have also been collected from Bahia state, northeastern Brazil (Figure 15). The detached erect colony has irregular to oblong autozooids, somewhat erratically disposed along the axis; the avicularia are unusual (V. E.S. Almeida pers. comm. 2009), as in specimens from Albatross Station 2319 (J. Sanner pers. comm. 2009), and are infrequent, as in encrusting specimens collected in Alagoas. Both forms of colonies found in Brazil have similar-sized zooids, orifices and avicularia. Distribution. West Atlantic – Brazil: Bahia and Alagoas states (present study); Colombian Caribbean (Montoya-Cadavid et al. 2007); Gulf of Mexico and Cuba (Canu & Bassler 1928); USA: Florida (Winston 1986)., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Marcusadorea, a new genus of lepralioid bryozoan from warm waters, pp. 57-68 in Zootaxa 2348 (1) on pages 63-66, DOI: 10.11646/zootaxa.2348.1.4, http://zenodo.org/record/5307125, {"references":["Canu, F. & Bassler, R. S. (1928) Fossil and Recent Bryozoa of the Gulf of Mexico region. Proceedings of the United States National Museum, 72, 1 - 199.","Winston, J. E. (1986) An annotated check-list of coral-associated bryozoans. American Museum Novitates, 2859, 1 - 39.","Montoya-Cadavid, E., Florez-Romero, P. & Winston, J. E. (2007) Checklist of the marine Bryozoa of the Colombian Caribbean. Biota Colombiana, 8 (2), 159 - 184.","Canu, F. & Bassler, R. S. (1925) Contribution a l'etude des Bryozoaires d'Autriche et de Hongrie. Bulletin de la Societe Geologique de France, series 4, 24, 627 - 690.","Waters, A. W. (1909) Reports on the Marine Biology of the Sudanese Red Sea, from collections made by Cyril Crossland, M. A., B. Sc., F. Z. S.; together with collections made in the Red Sea by Dr. Hartmeyer. XII. The Bryozoa, Part 1, Cheilostomata. Linnean Society of London Journal of Zoology, 31, 123 - 181.","Lamouroux, J. V. F. (1821) Exposition methodique des genres de l'ordre des polypiers, avec leur description et celles des principales especes figurees dans 84 planches; les 63 premieres appartenant a l'Histoire naturelle des zoophytes d'Ellis et Solander. V. Agasse, Paris, 115 pp.","David, L. & Pouyet, S. (1974) Revision des Bryozoaires cheilostomes miocenes du Bassin de Vienne (Autriche). Documents des Laboratoires de Geologie de la Faculte des Sciences de Lyon, 60, 83 - 257.","Reuss, A. E. (1874) Die fossilen Bryozoen des osterreichisch-ungarischen Miocans. Sitzungsberichte der kaiserlichen Akademie der Wissenschaften, Wien, 68, 219 - 222.","Banta, W. C. & Carson, R. J. M. (1977) Bryozoa from Costa Rica. Pacific Science, 31 (4), 381 - 424."]}

Marcusadorea tubulosa (Canu & Bassler, 1928) n. comb. (Figures 12–15, 18–19; Table 3) Holoporella (?) tubulosa Canu & Bassler 1928: 147, pl. 24, figs 1–6, text-fig. 33a. Coleopora tubulosa: Winston 1986: 16; Montoya-Cadavid, Flórez-Romero & Winston 2007: 169. Non Coleopora tubulosa Canu & Bassler 1925. Material examined. USNM 7532; Holoporella (?) tubulosa (Cotypes); Gulf of Mexico and Cuba. UFAL 0006; Marcusadorea tubulosa (Canu & Bassler); Coral Reefs of Pajuçara, Maceió, Alagoas State, Brazil, 9°40'49" S, 35°43'05" W, 0–3 m, coll. M.D. Correia, 20 September 2006, 1 encrusting colony. UFAL 0016; Marcusadorea tubulosa (Canu & Bassler); Coral Reefs of Amores, Maceió, Alagoas State, Brazil, 9°40'40" S, 35°42'10" W, 0–5 m, coll. M.D. Correia, 31 November 2006, 1 encrusting colony. UFAL 0021; 22 September 2006, 1 encrusting colony. MZUSP 0175; 0–3 m, coll. V. R. Cedro, 17 October 2005, 1 encrusting colony. MZUSP 0176; 0–5 m, coll. L.M. Vieira and M.D. Correia, 02 April 2007, 1 encrusting colony. MZUSP 0177; 1 encrusting colony (gold-coated specimen for SEM). MZUFBA 145; Coleopora corderoi Marcus; V.E.S. Almeida det., Costa do Dendê, Bahia State, Brazil, 13°31'25" S, 38°48'38" W, 45 m, coll. J.M.L. Dominguez and R. Silva, March 2002, 1 encrusting colony. MZUFBA 146; 13°44'37" S, 38°48'40" W, 47.7 m, coll. J.M.L. Dominguez and R. Silva, March 2002, 1 erect colony. MZUFBA 147; 13°59'48" S, 38°55'34" W, 16.6 m, coll. J.M.L. Dominguez and R. Silva, 2002, 2 encrusting colonies. Redescription. Colonies encrusting, unilaminar, or erect cylindrical with bifurcated branches, about 1.5– 3.0 mm diameter, disposed in longitudinal irregular series around the axis, reddish-brown in color. Autozooids large (0.911 mm long, 0.684 mm wide for Brazilian colonies), irregularly rectangular to oblong, separated by shallow grooves and distinctly raised lateral walls. Frontal shield convex, granular, irregularly porous, with numerous marginal areolar pores. Primary orifice widest in distal third, with a poster comprising two-thirds of a full circle, demarcated from the deep and broadly U-shaped anter by very short and blunt condyles. Primary orifice partly subcircular, bell-shaped, U-shaped anter and broader poster. No oral spines. Peristome tubular, proximally incomplete, forming a U-shaped pseudosinus. Secondary orifice rounded (0.226 mm long, 0.233 mm wide in Brazilian colonies), proximally incomplete. Large proximal suboral avicularium (0.233 mm long, 0.094 mm wide in Brazilian colonies) occasionally present, set obliquely on the side of the peristome, with the rostrum directed towards the proximal end of the pseudosimus, with curved spatulate mandible; rostrum oblong with rounded corners, slightly curving to outside, crossbar complete. Vicarious avicularia lacking. Ooecium prominent (0.383 mm long, 0.489 mm wide in Brazilian colonies), subglobular, partly incorporated into frontal shield of distal zooid, with some medium-sized pores scattered across granular-tubercular surface; ovicell opening inside peristome. Ancestrula not observed. Remarks. Marcusadorea tubulosa is characterized by large zooids, approximately 1.20 mm in length in Caribbean specimens (Canu & Bassler 1928), a thin-walled proximally incomplete peristome surrounding the orifice, and a large proximal suboral avicularium placed laterally on the rim of the peristome. Canu & Bassler (1928) noted that the budded zooids of M. tubulosa are distinct from those of other species of Holoporella Waters, 1909 (= Celleporaria Lamouroux, 1821). They noted similarities with the peristome of Coleopora species, which lack attachment marks around the operculum. They considered their material as an undetermined genus owing to the absence of ovicells and the growth mode of the colonies in which frontal budding is lacking occur. However, their plate clearly illustrates ovicells (Canu & Bassler 1928: pl. 26, fig. 6), the presence of which we confirmed by the examination of a paratype of Holoporella (?) tubulosa deposited in the United States National Museum (USNM 7532). The examination of this material also revealed that in some zooids there is a proximal suboral avicularium placed laterally on the peristome, occupying one third of it, and globular ovicells opening inside the peristome, as in the Brazilian material figured herein (Figures 13– 14). Prior to the original description of the genus Coleopora, Canu & Bassler (1925) described the Austrian fossil species Coleopora tubulosa, later regarded by David & Pouyet (1974) as a junior synonym of Coleopora insignis (Reuss, 1874). Teuchopora sp. recorded by Banta & Carson (1977) from Costa Rica resembles Marcusadorea tubulosa in general dimensions, frontal-shield formation, orifice, and the hemispherical aperture of ovicell opening into the base of the peristome, but it apparently lacks avicularia. Among the type material of M. tubulosa were found erect colonies, comprising cylindrical bifurcated branches, figured by Canu & Bassler (1928: pl. 24, figs 1, 4–5). Erect cylindrical colonies have also been collected from Bahia state, northeastern Brazil (Figure 15). The detached erect colony has irregular to oblong autozooids, somewhat erratically disposed along the axis; the avicularia are unusual (V. E.S. Almeida pers. comm. 2009), as in specimens from Albatross Station 2319 (J. Sanner pers. comm. 2009), and are infrequent, as in encrusting specimens collected in Alagoas. Both forms of colonies found in Brazil have similar-sized zooids, orifices and avicularia. Distribution. West Atlantic – Brazil: Bahia and Alagoas states (present study); Colombian Caribbean (Montoya-Cadavid et al. 2007); Gulf of Mexico and Cuba (Canu & Bassler 1928); USA: Florida (Winston 1986).

Marcusadorea corderoi (Marcus, 1949) n. comb. (Figures 6–11; Table 2) Coleopora corderoi Marcus 1949: 18, figs 23–25; Vieira, Migotto & Winston 2008: 30. Material examined. MZC Hassler Expedition Box 33, 36; Coleopora corderoi Marcus; off North of Rio de Janeiro, Brazil. Description. Colonies encrusting, unilaminar. Autozooids large (1.251 mm long, 0.766 mm wide for MCZ Hassler colonies), irregularly polygonal to rectangular, separated by distinctly raised lateral wall junctions. Frontal shield convex, surface granular with small scattered pores, these becoming obscured with a tuberculated frontal wall in increasing calcification; some larger marginal areolar pores remain visible. Primary orifice bell-shaped, wider proximally, anter arched, U-shaped, separated from broad and shallow poster by barely projecting condyles (mean 0.140 mm long, 0.178 mm wide for MCZ Hassler colonies). No oral spines. Peristome tubular, entire. Secondary orifice mostly rounded to oblong, with U- to V-shaped proximal edge (0.464 mm long, 0.401 mm wide for MCZ Hassler colonies). Suboral avicularium occasionally present, positioned laterally on inner side of peristome, in V-shaped portion, with curved and cuspidate mandible; none was present on the two colony fragments measured. Vicarious avicularia lacking. Ooecia not observed, but Marcus (1949) described them as prominent, globular, with porous surface, opening inside peristome. Ancestrula not observed. Remarks. When introducing the new species Coleopora corderoi, Marcus (1949) distinguished it from other Coleopora species by features of the peristome, absence of tubes on frontal wall, presence of avicularia with curved cuspidate mandible, and prominent ooecium opening inside the peristome. He described very large zooids, reaching 1.4–1.8 mm in length and 0.7 mm in width, while his drawing shows zooids about 1.2 mm long (see Marcus 1949: fig. 23). It was not possible to reexamine Marcus’ specimens as the holotype of C. corderoi is missing, along with other Brazilian specimens identified by Ernst Marcus that were deposited in the Museum of Natural History (UK) (M. Spencer Jones pers. comm., 2009) and Museu de Zoologia of Universidade de São Paulo (Brazil). Some of the colonies of M. corderoi collected north of Rio de Janeiro by the Hassler Expedition have smaller zooids (MCZ Hassler Box 33), more like those of Marcusadorea jamaicensis specimens, but colonies with large zooids (MCZ Hassler Box 36) as described by Marcus (1949) were also found. The Brazilian colonies found in MCZ collection (Figures 1–6) lack avicularia and ovicells. Other authors have already described further species with large zooids, raised peristomes, irregularly porous frontal shields, and raised lateral walls adjacent to the areolar pores, which we consider as the diagnostic features of the new genus Marcusadorea. The species M. tubulosa (Canu & Bassler, 1928) differs from M. corderoi by having smaller zooids and suboral avicularia with curved spatulate mandibles; colonies in which suboral avicularia are lacking might be difficult to identify; however, colonies of Marcusadorea without avicularia may be distinguished by features of the peristome and orifice. Marcusadorea corderoi has an almost rounded secondary orifice at the end of the peristome in most zooids. In the other three species of the genus, the peristome of most zooids is incomplete; there is a characteristically proximally incomplete Ushaped slit in Marcusadorea jamaicensis n. sp., a broader proximal V-shaped to U-shaped gap in M. tubulosa, and a quadrangular to subtriangular one in M. efatensis (Tilbrook, 2006). Distribution. Western Atlantic – Brazil: Rio de Janeiro (present study) and Espírito Santo states (Vieira et al. 2008).

Marcusadorea jamaicensis n. sp. (Figures 1–5, 16–17; Table 1) Coleopora corderoi: Winston 1986: 16, figs 68–69. Non Coleopora corderoi Marcus 1949. Holotype. VMNH 13359; Marcusadorea jamaicensis; Rio Bueno, Jamaica, 10 m, coll. J.B.C. Jackson, 1978. Paratypes. VMNH 13660–13662; Marcusadorea jamaicensis; Rio Bueno, Jamaica, 10 m, coll. J.B.C. Jackson, 1978, 3 colonies. Description. Colonies encrusting, unilaminar; living colonies bright orange. Autozooids large (averaging 0.975 mm long, 0.615 mm wide), irregularly polygonal to rectangular, separated by distinctly raised lateral wall junctions. Frontal shield convex, surface granular with small scattered pores, these becoming obscured by a tuberculated frontal wall with increasing calcification; some larger marginal areolar pores remaining visible. Primary orifice bell-shaped, wider proximally, anter arched, U-shaped, separated from broad and shallow poster by barely projecting condyles (mean, N=3, 0.140 mm length by 0.178 mm width of illustrated specimen VMNH 13662, Figure 17, primary orifice 0.172 mm length by 0.210 mm width). No oral spines. Peristome tubular, entire or proximally incomplete. Secondary orifice mostly rounded to oblong, with U- to Vshaped proximal edge (0.262 mm length by 0.227 mm width). Suboral avicularium occasional, positioned laterally on inner side of peristome, in pseudosinus, with curved and cuspidate mandible; rostrum triangular, hooked distally, directed frontally, crossbar complete. Vicarious avicularia lacking. Ooecium prominent, globular (0.239 mm long, 0.439 mm wide), with porous surface, opening inside peristome. Embryos cherry red. Polypides with 20–22 tentacles. Ancestrula not observed. Remarks. As noted below, some of the colonies of M. corderoi collected north of Rio de Janeiro by the Hassler Expedition have smaller zooids (MCZ Hassler Box 33), more similar to the Jamaican specimens, but colonies with larger zooids (MCZ Hassler Box 36) as described by Marcus (1949) also were found. The morphology of the two species is very similar, and the zooid length of Jamaican specimens overlaps (barely) with those from Brazil (MCZ Hassler Box 33). Since M. corderoi is a species that shows considerable variability in zooid size and shape, this could be considered geographic variation within one species. However, the dimensions of the secondary orifice and its proportion relative to zooid length raise difficulties that are best resolved by considering the Jamaican material as belonging to a second, previously undescribed species. This species, described under the name of Marcusadorea jamaicensis, has a zooid length only 81% and 76% of the M. corderoi specimens from MCZ Box 33 and Box 36 respectively. The length of the secondary orifice in M. jamaicensis is 60% and 51% of M. corderoi from MCZ Box 33 and MCZ Box 36 respectively. Moreover, the ratio of zooid length to secondary orifice length is about 3.7–1 for M. jamaicensis specimens, versus 2.4–1 and 2.9–1 for Box 33 and Box 36, respectively. Marcusadorea jamaicensis is characterized by an almost cleithridiate secondary orifice in most zooids; rarely the peristome may be proximally incomplete as a U-shaped slit. Distribution. Western Atlantic – Jamaica. Cryptic reef habitats, 10– 20 m., Published as part of Vieira, Leandro M., Migotto, Alvaro E. & Winston, Judith E., 2010, Marcusadorea, a new genus of lepralioid bryozoan from warm waters, pp. 57-68 in Zootaxa 2348 (1) on pages 59-61, DOI: 10.11646/zootaxa.2348.1.4, http://zenodo.org/record/5307125, {"references":["Winston, J. E. (1986) An annotated check-list of coral-associated bryozoans. American Museum Novitates, 2859, 1 - 39.","Marcus, E. (1949) Some Bryozoa from the Brazilian coast. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 3 (53), 1 - 33."]}

Zyzzyzus robustus (Petersen, 1990) Table 4 Zyzzyzus robustus Petersen, 1990: 183, figs. 32���33.��� Schuchert, 2001: 42 (footnote). Material examined. Preserved: Greenland: off Akuliaruseq (60 ��09���N. 44 �� 10 ���W) (140 m), 12 Jul 1970, polyps on steep rocky slope (type with sponge fibers in the hydrorhiza); Cape Farewell Expedition st. 93; southwest Qeqertat (59 �� 52.8`N, 43 �� 31.5`W) (225 m) (no date), 10 polyps without gonophores, on encrusting bryozoans, ZMUC (without collection number). Description. Hydroids 7.3 ���29.0 mm in height, covered by thick, heavy perisarc, growing on rock, sponge, and encrusting bryozoans. Hydrorhiza swollen, with 1���3 large, strong, club-shaped processes, covered by stiff perisarc. Hydranth large, oblong; hydranth body 1.4���3.7 mm high, 0.7���3.5 mm in diameter at base. Oral tentacles filiform, ca. 40 in number, arranged in five whorls, adnate to hypostome around mouth, tentacles laterally flattened in cross section; most proximal whorl arising at right angles to hypostome, most distal whorl reaching no more than one-third of hypostomial length, 0.8 ��� 0.14 mm long, 0.06���0.21 mm in diameter. Aboral tentacles filiform, 28���30 in number, tentacular bases contiguous to each other at base of hydranth body and in one whorl, tentacles laterally flattened in cross section, 0.8���3.8 mm long, 0.08���0.30 mm in diameter. Fertile hydranths with 21 blastostyles, ca. 1 mm in length, arranged in one imperfect or irregular whorl above aboral tentacles, each blastostyle with 8���14 spherical cryptomedusoid gonophores. Hydrocaulus trunk-like, bent from median to distal parts, cylindrical and covered by heavy, stiff perisarc at basal part, distal part of hydrocaulus narrowing, marked by end of stiff perisarc, surrounded by thinner perisarc. Transition between hydranth base and hydrocaulus smooth, without marked groove at region of perisarc secretion. Cnidome. Large stenoteles 10.2���13.5 x 8.2���12.2 ��m (11.2 �� 0.7 x 9.1 �� 0.8, n = 55); basitrichous isorhizae 5.3���6.8 x 1.8���2.5 ��m (6.2 �� 0.6 x 2.2 �� 0.3, n = 11) (aboral tentacles). Additional data. ���Hydrocaulus filled with parenchymatic endoderm, 20���30 peripheral longitudinal canals of equal diameter. [���] Blastostyles consisting of stout shaft distally branching into seven to eight very short, stout branches, each bearing a gonophore��� (Petersen 1990: 183���184). Colour. Hydranth and hydrocaulus yellowish-brown in alcohol-preserved material. Type. One polyp ca. 36.0 mm high (50 mm after Petersen 1990: 184). Hydrorhiza 6.5���6.8 mm in length, 2.3���2.8 mm in diameter. Hydrocaulus 25.8 mm in total length. Hydranth 3.7 mm in length, ca. 3.5 mm in diameter at base, with 40 filiform oral tentacles 0.9���1.4 mm in length, 0.12���0.21 mm in diameter; and 30 filiform aboral tentacles 2.9���3.8 mm in length, 0.18���0.35 mm in diameter. Fertile hydranth with 21 blastostyles, 1 mm in length, with several spherical cryptomedusoid incompletely developed. Hydrocaulus 5.8 mm in length with one primary polyp 7.3 mm in length; hydranth body 1.4���1.5 mm in length, 0.7���0.9 mm in diameter, ZMUC (without collection number). Additional material. One polyp from southwest Qeqertat (59 �� 52.8`N, 43 �� 31.5`W) (225 m) (no date), without gonophores, hydranth ca. 2.3 mm in length, ca. 2.6 mm in diameter at base, with detached oral and aboral tentacles. Hydrocaulus with two newly settled small polyps, 11 mm and 3 mm in height, respectively, ZMUC (without collection number). x �� SD (in ��m) Range (in ��m) n Hydrorhiza Number of processes 2 �� 1 1���3 1 Length - - - Diameter 1967 �� 678 992���2815 12 Hydrocaulus Length 11222 �� 9787 5857���25875 4 Proximal diameter 1771 �� 954 603���3376 27 Median diameter 1537 �� 1051 492���3089 26 Distal diameter 948 �� 542 457���2367 30 Hydranth Total length 2254 �� 954 1455���3744 8 Length from aboral tentacle bases to hypostomial tip 1431 �� 495 926���2163 8 Diameter at base 1845 �� 1073 687���3506 12 Oral tentacles Number 40 - 1 Length 586 �� 400 883���1451 13 Diameter 137 �� 53 61���214 40 Aboral tentacles Number 29 �� 1 28���30 2 Length 1754 �� 873 813���3817 12 Diameter 191 �� 78 77���352 60 Gonosome Total number - - - Gonophore Number 11 �� 2 8���14 2 Length - - - Diameter 190 �� 38 149���270 19 Blastostyles Number 21 - 1 Length 1047 1047 1 Diameter - - - Remarks. We corroborate the main morphological characters described by Petersen (1990). The diagnostic characters of Z. robustus are: absence of a groove at region of secretion of perisarc, long and monopodial blastostyles, and distance between adjacent aboral tentacles inferior to diameter of tentacular base. Zyzzyzus robustus is morphologically unique compared with other species of the genus, but Petersen (1990: 184) remarked ���the general morphology of Z. robustus nevertheless agrees so closely with that of the other species that it would seem unreasonable to erect a separate genus for it���. Morphologically it resembles species of Bouillonia Petersen, 1990, including its larger dimensions and its bathyal habit (cf. Petersen 1990; Schuchert 2001, Svoboda et al. 2006). Additional phylogenetic studies are necessary to establish the taxonomic affinities of the species. Petersen (1990: 184) considered young specimens of Z. robustus to be similar to Z. warreni, especially the thinner perisarc, noting that ���as the hydroid grows to a size about 10-15 mm [���] becomes thicker and harder, preventing further expansion of the tubers and the lower part of the hydrocaulus; growth lengthwise is still possible���. In fact, the specimens of Z. robustus reach a larger size than other species of Zyzzyzus. A synopsis of differences among species of Zyzzyzus is given in Table 7. Distribution. Northwestern Atlantic: off Akuliaruseq (60 ��09`N, 44 �� 10`W), southwest Qeqertat (59 �� 52.8`N, 43 �� 31.5`W) (Petersen 1990). Ecology. Substratum. The largest polyp in type material has two meshes of sponge fibres on its hydrorhiza, indicating that it was embedded in sponges, but dislodged from them during or after the collecting process. Remaining materials are attached to rocks and encrusting bryozoans. One newly-settled polyp from type material has a colony of the hydroid Clytia sp. growing along the hydrocaulus extension, arising from small hydrorhizal pedicels and with badly preserved hydrothecae and gonothecae. Depth. The uniqueness of Z. robustus is also reflected in its ecology. It is the only non-littoral species, having been collected at depths from 140 to 225 m off Greenland. Other specimens tentatively assigned to the genus Zyzzyzus from deep-water (110���400 m) are those described by Hirohito (1988: 26). He identified them as ��� Zyzzyzus (?) sp.��� but noted: ���as well-developed gonophores can not be observed, I feel some hisitation (sic) in referring it to the genus Zyzzyzus ���., Published as part of Campos, Carlos J. A., Marques, Antonio C. & Migotto, Alvaro E., 2007, A taxonomic revision of the genus Zyzzyzus Stechow, 1921 (Cnidaria: Hydrozoa: Tubulariidae), pp. 1-22 in Zootaxa 1627 on pages 10-12, DOI: 10.5281/zenodo.273951, {"references":["Petersen, K. W. (1990) Evolution and taxonomy in capitate hydroids and medusae. Zoological Journal of the Linnean Society, 100, 1 - 231.","Schuchert, P. (2001) Hydroids of Greenland and Iceland (Cnidaria, Hydrozoa). Meddelelser om Gronland, 53, 1 - 184.","Svoboda, A., Stepanjants, S. D. & Ljubenkov, J. (2006) The genus Bouillonia (Cnidaria: Hydrozoa: Anthoathecata). Three species from the northern and southern hemispheres, with a discussion of bipolar distribution of this genus. Zoologische Mededelingen, Leiden, 80 - 4 (14), 185 - 206."]}

Zyzzyzus warreni Calder, 1988 Figure 1, Tables 5���7 Tubularia solitaria Warren, 1906: 83, pls. 10���11, figs. 1���26 (junior primary homonym of Tubularia solitaria Rapp, 1829).��� Warren, 1908: 280.��� Millard, 1957: 179; 1966: 434.���de Kruijf, 1977: 19. Zyzzyzus solitarius;��� Stechow, 1921: 249; 1923: 50.��� Kramp, 1949: 198.��� Millard & Bouillon, 1974: 2.��� Millard, 1975: 40, fig. 16 A���B.��� Petersen, 1979: 121.��� Wedler & Larson, 1986: 72 ���73, fig. 1 A, pl. 1, fig. 3.��� Migotto & Silveira, 1987: 104 ���106, figs. 4���5.���Hirohito, 1988: 24���26, fig. 7. Corymorpha solitaria;��� Kramp, 1933: 12. Zyzzysus solitarius Bandel & Wedler, 1987: 39 [incorrect subsequent spelling]. Zyzzyzus warreni Calder, 1988: 49 ���50, figs. 38���40; 1991: 2068.��� Migotto, 1996: 25. Zyzzyzus calderi Petersen, 1990: 180 ���181: fig. 30 A���C.��� Calder, 1993: 66; 1998: 1850.���Marques & Migotto, 2001: 469 [new synonymy]. Material examined. Preserved: South Africa: Bat���s Cave Rocks, East London (33 ��01���S, 27 �� 54 ���E), 30 Dec 1948, three hydroids without gonophores, embedded in sponge, SAM H��� 3589; several hydroids with gonophores in sponge, SAM H��� 2648 cp 326 D; Langebaan Lagoon (33 ��09���S, 18 ��03.4���E), 15 Jul 1946, several hydroids with gonophores, embedded in sponge, SAM H��� 2649; Oudekraal (33 �� 58.5 ���S, 18 �� 22.2 ���E), 15 Mar 1934, several hydroids without gonophores, embedded in sponge, SAM H��� 2647; Oudekraal (33 �� 58.5 ���S, 18 �� 22.2 ���E), 15 Mar 1934, microslide with five hydroids without gonophores, SAM H��� 2647 A 122; Cape Peninsula (ca. 34 �� 10 ���S, 18 �� 30 ���E), microslide with three hydroids without gonophores, SAM H��� 2648 cp 326 D; St. James False Bay (ca. 34 �� 10 ���S, 18 �� 30 ���E), 1912, coll. and det. K.H. Barnard, several hydroids with gonophores, embedded in sponge, SAM H��� 398. Cape Verde Archipelago: S��o Vicente (ca. 16 �� 54 ���N, 25 ��00���W), 30 Jul 1904, microslide with detached gonophores, coll. and det. J. Ritchie, NMS 1959.33. 39. Bermuda: Flatt���s Inlet (ca. 32 �� 19 ���N, 64 �� 44 ��� 30 ���W), 1-2 m, 13 Sep 1977, several hydroids up to 5 mm high, some with developing blastostyles, embedded on Tedania ignis, coll. and det. D. Calder, ROMIZ B 133; Flatt���s Inlet (ca. 32 �� 19 ���N, 64 �� 44 ��� 30 ���W), 0 2 Aug 1982, 3 m, on underside of flat rock, several hydroids up to 10 mm high, with developing blastostyles, embedded in sponge and Eudendrium carneum, coll. and det. D. Calder, ROMIZ B 147; Castle Grotto, Castle Harbour (ca. 32 �� 21 ���N, 64 �� 42 ���W), 20 Jul 1982, 1 m, 25 m inside cave entrance, several hydroids up to 11 mm high, with developing blastostyles, embedded in sponge, coll. and det. D. Calder, ROMIZ B 165. Brazil: S��o Paulo state, S��o Sebasti��o, Ponta do Jarob��, (ca. 20 �� 45 ���S, 45 �� 26 ���W), 2 m, 0 3 May 1985, coll. A.E. Migotto, several hydroids without gonophores, embedded on Mycale laxissima, MZUSP 1987; S��o Sebasti��o, Ponta do Jarob��, (23 �� 45 ���S, 45 �� 26 ���W), 1.5m, 26 Mar 2002, on Mycale angulosa, coll. C. Campos, MZUSP 2000, 1998 and 1996; S��o Sebasti��o, Ponta do Jarob��, (23 �� 45 ���S, 45 �� 26 ���W), 13 May 2001, on Mycale angulosa, coll. C. Campos, MZUSP 1993; S��o Sebasti��o, Ponta do Jarob��, (23 �� 45 ���S, 45 �� 26 ���W), 27 May 2001, on Mycale angulosa and Hypnea cervicornis, coll. O.M. Oliveira, MZUSP 1991; S��o Sebasti��o, Ponta do Jarob��, (23 �� 45 ���S, 45 �� 26 ���W), 1m, 26 Mar 2002, on Mycale angulosa, coll. C. Campos, MZUSP 1989; S��o Sebasti��o, Ponta do Jarob��, (23 �� 45 ���S, 45 �� 26 ���W), 11 Jan 1988, on Mycale laxissima,, coll. A. E. Migotto, MZUSP 1985; S��o Sebasti��o, Ponta do Jarob��, (23 �� 45 ���S, 45 �� 26 ���W), 4.5m, 24 Jan 2002, coll. C. Campos, MZUSP 1982; Laje dos Moleques (ca. 23 �� 49 ��� 72 ���S, 45 �� 24 ��� 78 ���W), 4 m, 14 May 1987, several hydroids without gonophores, on Mycale laxissima, coll. A.E. Migotto, MZUSP 1986; S��o Sebasti��o, test panel at Ponta do Jarob��, (23 �� 49 ��� 42 ���S, 45 �� 25 ��� 16 ���W), 18 Feb 1992, several hydroids without gonophores, on Hypnea cervicornis and Eudendrium sp., coll. A.E. Migotto, MZUSP 1988; S��o Sebasti��o, Ponta do Jarob��, (23 �� 45 ���S, 45 �� 26 ���W), 1m, 24 Jan 2002, in sponge, coll. C. Campos, MZUSP 1999; S��o Sebasti��o, test panel at Ponta do Jarob��, 0 6 Oct 1989, several polyps with gonophores, on Clavelina oblonga, coll. A.E. Migotto, MZUSP 1995; S��o Sebasti��o, test panel at Ponta do Jarob��, (23 �� 49 ��� 42 ���S, 45 �� 25 ��� 16 ���W), 2 m, 13 May 1991, several hydroids with gonophores, embedded on Mycale angulosa, coll. A.E. Migotto, MZUSP 1994; S��o Sebasti��o, test panel at Ponta do Jarob��, (23 �� 49 ��� 42 ���S, 45 �� 25 ��� 16 ���W), 15 Jan 1985, several hydroids without gonophores, embedded on Mycale angulosa, coll. A.E. Migotto, MZUSP 1981; Ponta do Jarob�� (ca. 20 �� 45 ���S, 45 �� 26 ���W), 27 Nov 1984, 2.5m, several detached hydroids with gonophores, coll. F.L. da Silveira, MZUSP 1992; Ponta do Baleeiro (ca. 23 �� 49 ��� 45 ``S, 45 �� 25 ��� 24 ���W), 27 Aug 1985, 0.5m, several hydroids with gonophores, on Zoanthus sp., coll. A.E. Migotto, MZUSP 1990; Ponta do Jarob�� (ca. 20 �� 45 ���S, 45 �� 26 ���W), 27 Nov 1984, 2.5m, several hydroids with gonophores, embedded on Mycale angulosa, coll. F.L. da Silveira, MZUSP 001984; Live. S��o Sebasti��o, test panel at Ponta do Jarob��, (23 �� 49 ��� 42 ���S, 45 �� 25 ��� 16 ���W), 13 Oct 2001, several hydroids with gonophores, coll. C. Campos, MZUSP 1983; S��o Sebasti��o, test panel at Ponta do Jarob��, (23 �� 49 ��� 42 ���S, 45 �� 25 ��� 16 ���W), 25 Jul 2001, several hydroids without gonophores, on Hypnea cervicornis, coll. A.E. Migotto, MZUSP 1980; S��o Sebasti��o, test panel at Ponta do Jarob��, (23 �� 49 ��� 42 ���S, 45 �� 25 ��� 16 ���W), 23 Jan 2002, 1.5m, several hydroids with gonophores, on colonial ascidians, coll. A.E. Migotto, MZUSP 1997. Description. Hydroids 2.8���12.1 mm in height, extent of hydrocaulus embedded in sponges varied, also on tunicates, algae, and other cnidarians. Hydrorhiza with one to several bulbous, finger-shaped or clubshaped processes, 1.8���4.4 mm in length, 0.4���1.4 mm in diameter. Hydrorhizal processes grouped together at most basal part of hydrorhiza or spread around median parts of hydrocaulus, forming anchoring processes. Hydranth vasiform, hydranth body 0.4���1.8 mm high, 0.2 ���1.0 mm in maximum diameter. Oral tentacles filiform, 7���18 in number, in one whorl, adnate to hypostome around mouth, tentacles circular in cross section, 0.17���0.62 mm long, 0.02���0.12 mm in maximum diameter. Aboral tentacles filiform, 16���24 in number, in one whorl, evenly spaced at base of hydranth body, space between adjacent tentacular bases equal to diameter of tentacles, tentacles ovate in cross section near base, circular in cross section at distal free part, 0.4 ���2.0 mm long, 0.03���0.15 mm in maximum diameter. Fertile hydranths with 3���10 short, dichotomously ramified blastostyles, in one whorl above aboral tentacles. Hydranths monoecious, each blastostyle with 1���8 cryptomedusoid gonophores, male and female gonophores on different blastostyles; gonophores oval, thickened distal end forming vault around orifice, 4 short, laterally compressed crests at distalmost end. Developed gonophores with large central spadix; male gonophores 0.14���0.72 mm long, 0.14���0.38 mm in maximum diameter; female gonophores 0.7���0.9 mm long, 0.5���0.9 mm in maximum diameter. Completely developed female gonophores with up to 4 actinulae. Actinulae 0.40 mm in height, 0.45 mm in maximum diameter, with 6���16 slightly capitate aboral tentacles. Hydrocaulus flexible, sometimes twisted or bent in several directions, covered by thin perisarc, slightly thickened at basal part of hydrocaulus; hydrocaulus widening proximally, distal part of hydrocaulus cylindrical. Some hydroids growing horizontally on algae with median part of hydrocaulus widened. Perisarc secreted at region above circular groove, between hydrocaulus and hydranth base. Coenosarc of hydrocaulus with 7���10 longitudinal canals, 0.023���0.091 mm in diameter, uniformly aligned at most distal part, irregularly anastomosed along hydrocaulus length. Living hydroids with hydrocaulus well demarcated from hydrorhiza by perisarc thickness, near basal embedding region. Cnidome. Large stenoteles 9.2���13.7 x 8.2���13.2 ��m (11.5 �� 1.1 x 10.6 �� 1.3, n = 51); small stenoteles 5.1��� 7.1 x 3.9���5.9 ��m (5.9 �� 0.4 x 4.8 �� 0.5, n = 100); basitrichous isorhizae 5.7���7.6 x 1.8���3.2 ��m (6.7 �� 0.4 x 2.3 �� 0.2, n = 55); heterotrichous microbasic euryteles 8.0��� 11.8 x 3.7���6.7 ��m (9.8 �� 1.2 x 5.3 �� 0.9, n = 11); desmonemes 3.4���4.7 x 2.4���3.6 ��m (4.0 �� 0.3 x 3.0 �� 0.3, n = 45). Homotrichous microbasic euryteles 6.8���8.4 x 4.9���6.5 ��m (7.6 �� 0.4 x 5.5 �� 0.4, n = 26) (hydranth); merotrichous isorhizae 6.8 ���8.0 x 3.4���4.4 ��m (7.6 �� 0.3 x 3.9 �� 0.3, n = 35) (only seen on gonophores) (see Table 6 for detailed distribution of basitrichous isorhizae and desmonemes). Additional data. Warren (1906) described the development of a new hydranth from an adult hydrocaulus. Such regenerative potential under adverse conditions would be associated with additional nutrients stored in the hydrorhiza. Migotto & Silveira (1987: 106, fig. 4) also observed and figured the ���budding of a new trophosome starting from a hydrocaulus of a well developed trophosome��� (original text in Portuguese). We believe that these observations are correct because occasional growth of some new hydroids, budding off basally in the same hydrocaulus, were also observed by us. However, it is not clear if this is a result of aggregated settlement of actinulae or some kind of regenerative process (or asexual reproduction) of the polyps. Colour. Tentacles, hypostome, and hydrocaulus translucent to white near most proximal part; hydranth base bright orange; cryptomedusoids milky white with reddish spadix. Type. South Africa: St. James False Bay (ca. 34 �� 10 ���S, 18 �� 30 ���E), 1912, several hydroids without gonophores, embedded in sponge; hydrocauli 1.2���4.2 mm in length; hydranth 0.8���1.7 mm in length, 0.4 ���1.0 mm in maximum diameter; aboral tentacles 1.2 ���2.0 mm in length, 0.1���0.2 mm in maximum diameter, coll. K.H. Barnard, SAM H��� 398. Additional material assigned to Z. warreni [not studied by us]. Japan: Sagami Bay, hydroids with gonophores, Hydr. 3304���3329 (Hirohito 1988). Brazil: Pernambuco state, hydroids collected along a mangrove swamp on the rivers Porto Alegre and Ariquind��, deposited at ROM (Calder & Ma��al 1998). Puerto Rico: La Parguera, 1 m, hydroids with gonophores, on Tedania sp. and on Rhizophora sp. roots, USNM 60726, (Wedler & Larson 1986). South Africa: Oudekraal (33 �� 58.5 ���S, 18 �� 22.2 ���E), 15 Mar 1934, west coast of Cape Peninsula, SAM A��� 122; west coast of Langebaan Lagoon (33 ��09.0���S, 18 ��03.4���E), 15 Jul 1946, 7 m, coll. Day, 1959, SAM LB ��� 166; west coast of Saldanha Bay (33 ��02.5���S, 18 ��02���E), Sep 1957, SAM SB ��� 153 U; south coast of East London (33 ��01���S, 27 �� 54 ���E), Jul 1937, SAM L��� 172 (Millard 1966). Remarks. The diagnostic characters for this species are the two different classes of homotrichous microbasic euryteles, the presence of merotrichous isorhizae (though this character is uncertain for Z. floridanus because of the scarcity of fertile material); male and female gonophores present on different blastostyles (again inconclusive for Z. floridanus because of the scarcity of fertile material), and the distal end of the gonophore with four crests. The cnidome as described herein differs from previous accounts by Calder (1988), Migotto & Silveira (1987), and Millard (1975), who did not report heterotrichous microbasic euryteles, homotrichous microbasic euryteles [probably referred by Calder (1988) as ���?mastigophore���] and merotrichous isorhizae [probably referred by Calder (1988) as ���ovate isorhiza���]. Access to large quantities of polyps during this study likely favoured our conclusions. Our observations on Z. warreni corroborate most of previous reports regarding a single whorl of oral tentacles, morphology of oral and aboral tentacles, place and structure of perisarc secretion, number of blastostyles, and morphology of gonophores. However, some important differences were also noted, such as number of tentacles, internal and external morphology of the hydrocaulus, structure of blastostyles, and nematocyst composition. We discuss these differences below. Reports on the numbers of oral and aboral tentacles have been quite variable: 16���21 oral, 16���21 aboral (Warren 1906); 16���21 oral, 15���34 aboral (Millard 1975); 16���20 oral, 18���25 aboral (Wedler & Larson 1986); 12 oral and 24 aboral (Hirohito 1988); 15���20 oral, 22���25 aboral (Calder 1988); 15���21 oral, 15���34 aboral (Petersen 1990). However, hydranths of Z. warreni may develop new tentacles during ontogeny (pers. obs.), and such differences likely reflect tentacle counts that were made on different developmental stages of preserved materials. This also explains the different morphometric values given in Migotto & Silveira (1987). Petersen (1990) described Z. calderi based on part of the material ROMIZ B 370, previously described as Z. warreni by Calder (1988) from the Bermuda. Petersen (1990: 181) justified the new species based on a series of characters, viz. ���the extreme relative length of the neck region, the lower number of endodermal canals, the shape of the hydrocaulus, and the hydrorhiza which in Z. calderi consists of two to three small, club-shaped tubers with finger-like distal processes; the hydrorhiza in Z. warreni consists of large, flesh storage tubers and thin supporting rootlets. Desmonemes in Z. calderi measure 3.7 x 2.8���4.2 x 2.9 ��m, in Z. warreni 4.5 x 3.6���6.3 x 3.6 ��m, and the isorhizas in Z. calderi measure 6.4 x 1.9���7.3 x 2.1 ��m, those of Z. warreni measure 8.1 x 3.6 ���9.0 x 3.6 ��m.��� However, as seen in Table 7, the nematocysts of the two species have different proportions. Although Petersen did not provide morphometric values to document the ���extreme relative length of the neck region���, he described the ���neck region [of Z. calderi] long and slim, four to five times as long as hydrocaulus proper��� (Petersen 1990: 180). Meanwhile, Petersen (1990: 182) described Z. warreni with ���neck region nearly as long as hydrocaulus proper���. However, we maintain that there is no clear morphological differentiation between distal and median regions of the hydrocaulus (see discussion of Z. floridanus above). The existence of ���about 16 longitudinal canals of equal size��� (Petersen 1990: 182, for Z. warreni) is not corroborated by our observations, although the reported number is quite varied (���about 16 ��� for Millard 1975; ���about ten��� for Hirohito 1985). We counted numbers of canals in living polyps and measured histological cross sections at proximal, median, and distal parts of the hydrocaulus. Our results clearly show that canals are open distally, totally closed proximally, and have several anastomoses along the hydrocaulus; moreover, some canals are wider than others and are present in varied numbers (7���10). Combining our data with literature reports, we conclude that the number of canals varies, from 7 (in Brazilian specimens) to 16 (in South African specimens). The proposed diagnostic feature of Z. calderi falls within this range. The supposed differences in dimensions of desmoneme and basitrichous isorhiza nematocysts are not upheld in a broader, non-regional analysis. Pooling data from various regions (South Africa, Bermuda, and Brazil, see Table 5), the ���gap��� between the putative species disappeared and the conclusion of Petersen (1990) of differences between them is no longer valid. Petersen (1990: 181) described the hydrorhiza of Z. calderi as ���consisting of one to three short, finger- or club-shaped tubers which may split up into finger shaped processes��� and that of Z. warreni as ���hydrorhiza developed as slender rootlets and swollen tubers of varying size and shape��� (Petersen 1990: 182). Warren (1906: 85) was the first to describe ���branches of two kinds: thin rootlets���and thickened fleshy structures���, also observed by Hirohito (1988: 24) (���tuberous root of sweet potatoes in shape, branched or not branched, usually producing several slender processes���). We believe that the morphology of the hydrorhiza should be adopted carefully as a diagnostic character, as it is variable and dependent on the substratum type or host structure. We are convinced that this character is not reliable in describing new species within this genus. Conspicuous reproductive characters that might differentiate Z. warreni from Z. calderi were not listed by Petersen (1990). He (page 181) described the gonophores of Z. calderi as follows: ���gonophores developed [���] male and females on same blastostyle��� and those of Z. warreni as: ���male and female gonophores produced by same hydroid on separate blastostyles��� (Petersen 1990: 182). However, Calder (1988: 50) described the same materials as ���in examined specimens, gonophores incompletely developed���, as represented in his figure 39. Although we could confirm the information for Z. warreni, we did not check the only specimens assigned to Z. calderi. Attempts were made by D.R. Calder to locate the type material of Z. calderi at the ROM, without success. For these reasons, we regard Z. calderi as a junior synonym of Z. warreni. More studies on material from other regions is needed to enhance the knowledge of this genus and species. Distribution. Northwestern Atlantic: P, Published as part of Campos, Carlos J. A., Marques, Antonio C. & Migotto, Alvaro E., 2007, A taxonomic revision of the genus Zyzzyzus Stechow, 1921 (Cnidaria: Hydrozoa: Tubulariidae), pp. 1-22 in Zootaxa 1627 on pages 12-20, DOI: 10.5281/zenodo.273951, {"references":["Calder, D. R. (1988) Shallow-water hydroids of Bermuda: the Athecatae. Royal Ontario Museum Life Sciences Contributions, 148, 1 - 107.","Warren, E. (1906) On Tubularia solitaria sp. nov., a hydroid from the Natal coast. Annals of the Natal Museum, 1, 83 - 96.","Rapp, W. (1829) Untersuchungen uber den Bau einiger Polypen des mittellandischen Meeres. Nova Acta Physico-Medica Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum, 14, 643 - 658.","Warren, E. (1908) On a collection of hydroids, mostly from the Natal coast. Annals of the Natal Museum, 1, 269 - 355.","Millard, N. A. H. (1957) The Hydrozoa of False Bay, South Africa. Annals of the South African Museum, 41, 173 - 243.","de Kruijf, H. A. M. (1977) Bursting pacemaker activity in the solitary hydroid Tubularia solitaria. Journal of Experimental Biology, 68, 19 - 34.","Stechow, E. (1921) Neue genera und species von hydrozoen und anderen evertebraten. Archiv fur Naturgeschichte, 87, 248 - 265.","Kramp, P. L. (1949) Origin of the hydroid Family Corymorphidae. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kobenhavn, 111, 183 - 215.","Millard, N. A. H. & Bouillon, J. (1974) A collection of hydroids from Mocambique, East Africa. Annals of the South African Museum, 65 (1), 1 - 40.","Millard, N. A. H. (1975) Monograph on the Hydroida of southern Africa. Annals of the South African Museum, 68, 1 - 513.","Petersen, K. W. (1979) Development of coloniality in Hydrozoa. In: Larwood, G. & Rosen, B. R. (Eds), Biology and Systematics of Colonial Animals. Academic Press, New York, pp. 105 - 139.","Wedler, E. & Larson, R. (1986) Athecate hydroids from Puerto Rico and the Virgin Islands. Studies on Neotropical Fauna and Environment, 21 (1 - 2), 69 - 101.","Migotto, A. E. & Silveira, F. L. (1987) Hidroides (Cnidaria, Hydrozoa) do litoral sudeste e sul do Brasil: Halocordylidae, Tubulariidae e Corymorphidae. Iheringia, Serie Zoologia, 66, 95 - 115.","Kramp, P. L. (1933) The Scoresby Sound Committee's 2 nd East Greenland Expedition in 1932 to King Christian IX's Land. Coelenterata, Ctenophora and Chaetognatha. Meddelelser om Gronland, 104 (11), 1 - 20.","Bandel, K. & Wedler, E. (1987) Hydroid, amphineuran and gastropod zonation in the littoral of the Caribbean Sea, Colombia. Senckenbergiana Maritima, 19 (1 - 2), 1 - 129.","Migotto, A. E. (1996) Benthic shallow-water hydroids (Cnidaria, Hydrozoa) of the coast of Sao Sebastiao, Brazil, including a checklist of Brazilian hydroids. Zoologische Verhandelingen, Leiden, 306, 1 - 125.","Petersen, K. W. (1990) Evolution and taxonomy in capitate hydroids and medusae. Zoological Journal of the Linnean Society, 100, 1 - 231.","Calder, D. R. (1993) Local distribution and biogeography of the hydroids (Cnidaria) of Bermuda. Caribbean Journal of Science, 29 (1 - 2), 61 - 74.","Calder, D. R. & Mayal, E. M. (1998) Dry season distribution of hydroids in a small tropical estuary, Pernambuco, Brazil. Zoologische Verhandelingen, Leiden, 323, 69 - 78.","Millard, N. A. H. (1966) The Hydrozoa of the south and west coasts of South Africa. Part III, The Gymnoblastea and small families of Calyptoblastea. Annals of the South African Museum, 48 (18), 427 - 487.","Calder, D. R. (1991) Associations between hydroid species assemblages and substrate types in the mangal at Twin Cays, Belize. Canadian Journal of Zoology, 69, 2067 - 2074.","Rees, W. J. & Thursfield, S. (1965) The hydroid collections of James Ritchie. Proceedings of the Royal Society of Edinburgh, Biology, 69 (2), 34 - 43.","Watson, J. E. (1978) New species and new records of Australian athecate hydroids. Proceedings of the Royal Society of Victoria, 90 (2), 301 - 314.","Marques, A. C. (1993) Sistematica dos Eudendriidae L. Agassiz, 1862 (Cnidaria, Hydrozoa) do Litoral Paulista. Thesis, Instituto de Biociencias, Universidade de Sao Paulo, Sao Paulo, 168 pp.","Migotto, A. E. (1993) Hidroides (Hydrozoa, Cnidaria) Marinhos Bentonicos da Regiao Costeira do Canal de Sao Sebastiao, SP, Thesis, Instituto de Biociencias, Universidade de Sao Paulo, Sao Paulo, 258 pp.","Guerrazzi, M. C. (1999) Padroes Populacionais e Ecologia do Forrageamento de Echinaster brasiliensis Muller & Troschel (Echinodermata: Asteroidea) no Canal de Sao Sebastiao, SP. Thesis, Instituto de Biociencias, Universidade Estadual Paulista, Sao Paulo, 77 pp."]}

Zyzzyzus floridanus Petersen, 1990 Table 2 Zyzzyzus floridanus Petersen, 1990: 178, fig. 29. Material examined. Preserved: United States of America: Florida, Miami (ca. 25 �� 45 ���N, 80 �� 15 ���W), west of Long Reef (2 m), 27 Jan 1965, coll. C. Nielsen, one hydroid, with one fully developed gonophore (type); one hydroid, without gonophores; several hydroids, without gonophores; embedded in the sponge Callyspongia vaginalis (paratypes), ZMUC (without collection number). Description. Hydroids 2.2���4.4 mm high, with hydrocaulus stiffly embedded in sponge (Callyspongia vaginalis). Hydrorhiza with up to four rounded bulbs. Hydranth vasiform; hydranth body 0.5���0.9 mm high, 0.3���0.5 mm in maximum diameter. Oral tentacles short, filiform, 8���15 in number, arranged in two close whorls, with both whorls adnate to hypostome around mouth, 0.15���0.28 mm long, 0.03���0.05 mm in maximum diameter. Aboral tentacles long, filiform, 15���20 in number, evenly spaced at base of hydranth body in one whorl, space between adjacent tentacular bases equal to diameter of tentacles; tentacles ovate in cross section near base, circular in cross section at distal free part, 0.7���2.1 mm long, 0.03���0.08 mm in maximum diameter. Fertile hydranths with short blastostyles in one whorl above aboral tentacles, each blastostyle with cryptomedusoid gonophores. Only female gonophores observed. Developed female gonophores oval, widening distally, as long as hydranth, reaching tips of oral tentacles, distal end smooth around orifice, 0.47���0.50 mm long, 0.28���0.30 mm in maximum diameter. Gonophores when completely developed containing one actinula. Hydrocaulus flexible, sometimes twisted or bent in several directions, covered by thin perisarc at median and distal parts, basal part of hydrocaulus embedded in sponge with thickened perisarc; hydrocaulus widening proximally, distal part of hydrocaulus cylindrical, basal and median parts ovate in cross section. Perisarc secreted at region above circular groove, between hydrocaulus and hydranth base; preserved material with hydrocaulus having folded perisarc. Coenosarc of hydrocaulus with 8���11 basal longitudinal canals, anastomosed in 4���6 distal longitudinal canals. Cnidome. Large stenoteles 11.1 x 10.0 ��m (n = 1); small stenoteles 5.3 ���6.0 x 4.0��� 5.3 ��m (5.6 �� 0.17 x 4.4 �� 0.35, n = 16); basitrichous isorhizae 5.3���6.1 x 2.0��� 2.5 ��m (5.5 �� 0.26 x 2.2 �� 0.13, n = 10); heterotrichous microbasic euryteles 7.4���8.4 x 4.2 ���5.0 ��m (7.8 �� 0.31 x 4.6 �� 0.26, n = 16); desmonemes 3.1���3.6 x 2.1��� 3.1 ��m (3.4 �� 0.21 x 2.6 �� 0.25, n = 11) (hydranth). Additional data. Petersen (1990: 178���179) described ���about 14 tentacles��� in the actinula of the type material and concentrations of nematocysts ���at tip and along aboral side��� in the oral tentacles and ���along aboral surface��� in the aboral tentacles [pseudofiliform type defined by Pr��vot (1959)]. Male gonophores are unknown. Colour. Transparent white in alcohol-preserved material (Petersen 1990, our observations). Type. One hydroid removed from sponge, up to 2.3 mm high, with 15 short filiform oral tentacles and 16 filiform aboral tentacles. Hydranth supporting one completely developed gonophore containing one actinula and 5 short incompletely developed blastostyles, arising distal to aboral tentacles in a circular whorl, ZMUC (without collection number) (Petersen 1990: 179). Additional material. Several hydroids without gonophores, embedded in sponge (C. vaginalis), ZMUC (without collection number). x �� SD (in ��m) Range (in ��m) n Hydrorhiza Number of processes 4 - 1 Length 1247 �� 27 1224���1276 3 Diameter 357 �� 82 274���466 6 Hydrocaulus Length 1494 �� 612 469���2185 6 Proximal diameter 375 �� 114 250���586 15 Median diameter 283 �� 98 147���461 14 Distal diameter 273 �� 58 177���353 20 Hydranth Total length 651 �� 116 504���901 12 Length from aboral tentacle bases to hypostomial tip 467 �� 92 373���648 10 Diameter at base 425 �� 68 320���547 16 Oral tentacles Number 10 �� 3 8���15 6 Length 219 �� 50 149���284 11 Diameter 39 �� 5 29���49 21 Aboral tentacles Number 17 �� 2 15���20 8 Length 1413 �� 420 685���2103 24 Diameter 59 �� 12 31���77 28 Gonosome Total number 1 - 1 Male gonophore Number - - - Length - - - Diameter - - - Female gonophore Number 1 - 1 Length 479 �� 16 466���497 3 Diameter 282 �� 6 276���288 3 Blastostyles Number 1 - 1 Length - - - Diameter - - - Remarks. The diagnostic characters of Z. floridanus are the barrel-shaped hydranth, the two whorls of oral tentacles, and the smooth surface of the female gonophore around orifice. The study of the material above corroborated most details of the general description by Petersen (1990). However, our observations do not coincide with his descriptions of the hydranth ("slimly vasiform") and the aboral tentacles (���rectangular cross-section���) (Petersen 1990: 178). Although no morphometric data were given by Petersen (1990), the increasing number of oral tentacles might cause a projection enlarging the hypostome, changing the whole morphology of the hydranth. As for aboral tentacles being ���rectangular in cross-section," there are no other accounts of this kind of tentacular morphology among tubulariid species. Instead, we adopted the general nomenclature and definitions following Pr��vot (1959). Petersen (1990: 156) observed in several tubulariids the existence of a ���reversed pear-shaped neck region, which in Zyzzyzus is very prolonged and constitutes about half of the hydrocaulus.��� From our observations, Z. robustus is the only species ever assigned to the genus with a narrowed distal hydrocaulus, clearly distinguishable by perisarc thickness. Therefore, we did not establish any morphological differentiation between distal and median regions in the hydrocaulus of Z. floridanus [which Petersen (1990) considered as having a ���ringshaped swelling���] or in any other species of the genus, and we discard the idea of distinguishing a ���neck region" in the hydrocaulus of species of Zyzzyzus. We (and Petersen, 1990) observed only female gonophores, and male gonophores of the species are unknown. Consequently, the nature of the blastostyles (whether mono- or dioecious) has not yet been determined for this species. Distribution. Northwestern Atlantic: shallow-waters off west of long reef, Miami, Florida, United States of America (ca. 25 �� 45 ���N, 80 �� 15 ���W) (Petersen 1990). Ecology. Substratum. The thickness of the perisarc on hydrorhizal processes suggests that the species is strongly embedded on volcano-like projections of the sponge Callyspongia vaginalis. Seasonality. January (one actinula present in type material) (Petersen 1990). Depth. 2 m at low tide (Petersen 1990)., Published as part of Campos, Carlos J. A., Marques, Antonio C. & Migotto, Alvaro E., 2007, A taxonomic revision of the genus Zyzzyzus Stechow, 1921 (Cnidaria: Hydrozoa: Tubulariidae), pp. 1-22 in Zootaxa 1627 on pages 5-7, DOI: 10.5281/zenodo.273951, {"references":["Petersen, K. W. (1990) Evolution and taxonomy in capitate hydroids and medusae. Zoological Journal of the Linnean Society, 100, 1 - 231.","Prevot, E. (1959) Morphologie et evolution des structures tentaculaires chez les hydraires gymnoblastes Capitata. Recueil des Travaux de la Station Marine D'Endoume, 29, 91 - 126."]}

Zyzzyzus spongicolus (von Lendenfeld, 1884) Table 3 Tubularia spongicola von Lendenfeld, 1884: 597, pl. 26, fig. 50. Zyzzyzus spongicolus;��� Watson, 1978: 303, fig. 1, C���M; 1982: 85, pl. 7.4, fig. 4.6 a.��� Petersen, 1990: 178.���Marques & Migotto, 2001: 469. Zyzzyzus spongicola;��� Rudman, 1981: 421, pl. 1, fig. 1.��� Edmunds, 1987: 189 [incorrect subsequent spelling]. Material examined. Preserved: Australia: Victoria, Port Phillip Bay, Popes Eye Reef (ca. 38 ��S, 144 �� 50 ���E), 10 m, 29 Jan 1999, about 40 polyps with gonophores, embedded in sponges, in two samples, coll. and det. J.E. Watson, MZUSP 1978 and MZUSP 1977; Port Phillip Bay, Point Wilson (ca. 38 ��S, 144 �� 50 ���E), 0 6 Apr 1999, about 125 polyps with gonophores, embedded in sponge, coll. and det. J.E. Watson, MZUSP 1976. Description. Hydroids 1.4 ���4.0 mm in height; hydrocaulus stiff, embedded on horny and soft sponges by expanded hydrorhiza. Hydrorhiza with several bulbous or finger-shaped processes, varied in morphology and length. Hydranth vasiform, hydranth body 0.7 ���1.0 mm high, 0.2 ���1.0 mm in maximum diameter. Oral tentacles filiform, 9���12 in number, in one whorl, adnate to hypostome around mouth; tentacles circular in cross section throughout. Aboral tentacles filiform, 11���17 in number, evenly spaced in one whorl at base of hydranth body; tentacles ovate in cross section at bases, circular in cross section at distal free part, 0.7���1.8 mm long, 0.03���0.11 mm in maximum diameter. Fertile hydranths with 4���7 blastostyles, in one whorl above aboral tentacles. Hydranths monoecious, each blastostyle with 2���6 cryptomedusoid gonophores in different developmental stages; male and female gonophores on same blastostyles. Developing male gonophores oval, almost spherical in later developmental stages, with visible spadix, 0.27���0.51 mm long, 0.03���0.11 mm in maximum diameter. Female gonophores with raised collar around orifice at distal region in later developmental stages, enlarged, with up to 3 actinulae. Hydrocaulus flexible, sometimes twisted or bent in several directions, covered by thin perisarc, slightly thickened at basal part of hydrocaulus. Perisarc secreted at region above circular groove, between hydrocaulus and hydranth base. Hydrocaulus sometimes with small tubular hydrorhizal processes at most proximal region and marked by sets of transverse grooves at distal region. Coenosarc of hydrocaulus with endodermal canals, anastomosed along hydrocaulus length at region above thickened perisarc. Cnidome. Large stenoteles 8.5���11.8 x 7.3���10.8 ��m (10.4 �� 0.97 x 9.5 �� 1.06, n = 14); small stenoteles 5.3���5.5 x 4.3���4.6 ��m (5.4 �� 0.15 x 4.5 �� 0.25, n = 2); basitrichous isorhizae 6.1���7.6 x 2.2���3.7 ��m (6.6 �� 0.49 x 2.7 �� 0.49, n = 7); heterotrichous microbasic euryteles 6.5���8.6 x 3.3���4.3 ��m (7.3 �� 0.73 x 3.7 �� 0,31, n = 15); desmonemes 3.2���4.6 x 2.6 ���3.0 ��m (3.6 �� 0.39 x 2.8 �� 0.12, n = 9) (hydranth). Additional data. Watson (1978) described 12 endodermal canals communicating with the gastral cavity of the hydranth. Main body of actinulae 0.25���0.3 mm in diameter, supporting 10���12 aboral tentacles (Watson 1978). Colour. Hydranth and tentacles translucent white, main body of the hydranth red, actinulae white, spadix of well developed gonophores orange (Watson 1978). Type. Tubularia spongicola von Lendenfeld (1884), AM G 10809 and AM G 10789 [cf. von Lendenfeld (1884)]. Additional material. Australia: Victoria, Port Phillip Bay, Popes Eye Reef (ca. 38 ��S, 144 �� 50 ���E), about 30-40 polyps embedded in sponge, in two samples; about 125 polyps embedded in sponge; New South Wales, Port Jackson, AM G 10809, AM G 10798. Remarks. The diagnostic characters of Z. spongicolus are male and female gonophores growing on the same blastostyle (although this character is not known for Z. floridanus because of the scarcity of fertile material), and the raised collar around the orifice in female gonophores. In fact, the species is quite similar to Z. warreni. The tenuous diagnosis of the species was noted by Calder (1988: 50), who stated: ��� Z. warreni differs from Z. spongicolus in having male and female gonophores on separate blastostyles instead of on the same ones [���]. From the existing descriptions, the two seem otherwise remarkably similar and may prove conspecific.��� Reports of monoecy in Zyzzyzus are somewhat confused in the literature. Calder (1988) defined the genus as monoecious, and Marques & Migotto (2001) extended this interpretation to Z. spongicolus, Z. calderi, and Z. warreni in their study based both on Petersen���s (1990) data and on the description of Z. warreni by Migotto & Silveira (1987). Our observations from extensive collections demonstrate that species of Zyzzyzus are monoecious, with gonophores of different sexes either developing on the same or on different blastostyles (uncertain for Z. floridanus; see above). From our observations, Zyzzyzus spongicolus is the only species of the genus in which male and female gonophores are both present on the same blastostyle. Von Lendenfeld (1884) described Tubularia spongicola from hydroids collected in Port Jackson, New South Wales, Australia (material AM G 10809 and AM G 10789). Watson (1978) re-examined von Lendenfeld���s material, remarking on some imprecise observations regarding the morphology of oral tentacles and branching of blastostyle, and complemented von Lendenfeld���s description concerning the number of aboral tentacles and structure of gonophores and hydrorhizal processes. Our study of Australian material agrees with the general descriptions of von Lendenfeld (1884) and Watson (1978) regarding morphometric features, morphology of the hydranth, region of perisarc secretion and hydrocaulus, and morphology of blastostyles and gonophores. However, we observed some slight differences in composition and dimensions of nematocysts. Watson (1978) described large stenoteles 7.5 ���9.0 x 9.5 ��m; small stenoteles, 5.5 x 3.0 ��m; basitrichous isorhizae, 8.0 x 3.5 ��m and heterotrichous anisorhizae, 3.0 x 5.0 ��m and the number of oral and aboral tentacles. She also reported 12���16 aboral tentacles, and 10���12 oral tentacles. However, she studied live material, which may account for part of the difference. Zyzzyzus spongicolus presents great variability in the morphology of hydrorhizal processes, apparently as a phenotypic response to the skeletal structure of the sponge. Similar observations have been made for Z. warreni. x �� SD (in ��m) Range (in ��m) n Hydrorhiza Number of processes - - - Length - - - Diameter - - - Hydrocaulus Length 1777 �� 599 722���2918 9 Proximal diameter 347 �� 155 210���755 25 Median diameter 315 �� 86 191���464 22 Distal diameter 281 �� 76 118���395 22 Hydranth Total length 869 �� 202 679���1075 4 Length from aboral tentacle bases to hypostomial tip 523 �� 145 350���653 4 Diameter at base 534 �� 187 262���1047 17 Oral tentacles Number 11 �� 1 9���12 19 Length - - - Diameter - - - Aboral tentacles Number 13 �� 2 11���17 19 Length 1162 �� 427 696���1770 8 Diameter 70 �� 20 32���110 35 Gonosome Total number - - - Male gonophore Number 4 �� 2 2���6 5 Length 381 �� 99 275���514 5 Diameter 285 �� 78 212���413 5 Female gonophore Number - - - Length - - - Diameter - - - Blastostyles Number 6 �� 1 4���7 5 Length - - - Diameter - - - Distribution. Eastern Indian Ocean: shallow-waters off Port Phillip Bay (ca. 38 ��S, 144 �� 50 ���E), Victoria, Australia. Southwest Pacific: Port Jackson (ca. 33 �� 50 ���S, 151 �� 15 ���E), New South Wales, Australia (Watson 1978); Montague island (36 �� 16 ���S, 150 �� 12 ���E), south of Narooma, New South Wales, Australia; Green Point (33 �� 55 ���S, 151 �� 10 ���E), Sydney Harbour, New South Wales, Australia (Rudman 1981). Ecology. Substratum. Species with ���fuzzy growth��� on orange-coloured sponges in sheltered ocean bays (Watson 1982: 85). Seasonality. From April to December (Watson 1978, 1982). Associations. Z. spongicolus is a preferred food item of the nudibranch Cuthona kuiteri, whose cerata developed processes acting as a kind of mimetic ���special resemblance��� with Z. spongicolus tentacles (Edmunds 1987: 191)., Published as part of Campos, Carlos J. A., Marques, Antonio C. & Migotto, Alvaro E., 2007, A taxonomic revision of the genus Zyzzyzus Stechow, 1921 (Cnidaria: Hydrozoa: Tubulariidae), pp. 1-22 in Zootaxa 1627 on pages 7-10, DOI: 10.5281/zenodo.273951, {"references":["von Lendenfeld, R. (1884) Das system der medusen. Zoologischer Anzeiger, 8, 425 - 429.","Watson, J. E. (1978) New species and new records of Australian athecate hydroids. Proceedings of the Royal Society of Victoria, 90 (2), 301 - 314.","Petersen, K. W. (1990) Evolution and taxonomy in capitate hydroids and medusae. Zoological Journal of the Linnean Society, 100, 1 - 231.","Rudman, W. B. (1981) Polyp mimicry in a new species of aeolid nudibranch mollusc. Journal of the Zoological Society of London, 193, 421 - 427.","Edmunds, M. (1987) Color in opisthobranchs. American Malacological Bulletin, 5 (2), 185 - 196.","Stepanjants, S. D. & Svoboda, A. (1999) Genera and species of Antarctic Corymorphinae with the discussion concerning structure of this subfamily. Proceedings of the Zoological Institute of the Russian Academy of Sciences, 281, 47 - 54.","Calder, D. R. (1988) Shallow-water hydroids of Bermuda: the Athecatae. Royal Ontario Museum Life Sciences Contributions, 148, 1 - 107.","Migotto, A. E. & Silveira, F. L. (1987) Hidroides (Cnidaria, Hydrozoa) do litoral sudeste e sul do Brasil: Halocordylidae, Tubulariidae e Corymorphidae. Iheringia, Serie Zoologia, 66, 95 - 115.","Watson, J. E. (1982) Hydroids (Class Hydrozoa). In: Shepherd, S. A. & Thomas, I. M. (Eds), Marine Invertebrates of Southern Australia. Part I, Handbooks Committee on Behalf of the South Australian Government. Published by Favor of the Honourable the Premier (D. O. Tonkin, M. P.), pp. 77 - 115."]}

Zyzzyzus Stechow, 1921 Table 1 Zyzzyzus Stechow, 1921: 249; 1923: 49.��� Kramp, 1933: 12, 1949: 198.��� Millard, 1975: 38.��� Watson, 1978: 302.��� Petersen, 1979: 121.��� Bouillon, 1985: 112.��� Wedler & Larson, 1986: 72.��� Migotto & Silveira, 1987: 104.��� Calder, 1988: 49.���Hirohito, 1988: 22.��� Petersen, 1990: 177.��� Migotto, 1996; 25.���Marques & Migotto, 2001: 465. Zyzzygus;��� Neave, 1940: 712 [incorrect subsequent spelling]. Zyzzyzuz;��� Bouillon, 1985: 243 [incorrect subsequent spelling]. Description. Hydroid solitary. Hydranth vasiform or oblong. Oral tentacles filiform, short, in one or two whorls, circular in cross-section, adnate to hypostome. Aboral tentacles in one whorl, short and slightly capitate in juvenile polyps, long and filiform in mature ones, flattened in cross-section proximally, circular in cross-section distally. Hydrocaulus long, cylindrical, slightly widening basally, covered by thin perisarc originating from circular groove immediately below hydranth base. Space of coenosarc of hydrocaulus filled by large vacuolated gastrodermal cells, peripheral region with longitudinal endodermal canals or ridges formed by columnar epithelium. Distal portion of endodermal canals not continuously circular but opened in ridges, fusing proximally to form true canals, some wider than others. Anastomoses of canals occurring at any level of hydrocaulus. Basal region of hydrorhiza forming swollen structures of variable morphology: flattened base resembling basal disc, supporting small finger-shaped processes to rounded bulbs or anchoring processes of varied length. Hydranth with male and female gonophores on the same or on separate blastostyles. Gonophores cryptomedusoid, supported by short, dichotomously developed blastostyles. Mature female gonophores containing up to four actinulae. Type species. Zyzzyzus warreni Calder, 1988, nomen novum for Zyzzyzus solitarius (Warren, 1906); originally described as Tubularia solitaria Warren, 1906 (Millard 1975, Migotto & Silveira 1987, Calder 1988). Taxonomic history. The genus Zyzzyzus was established by Stechow (1921) for a single species, Tubularia solitaria Warren, 1906. However, that species name is a permanently invalid junior primary homonym of Tubularia solitaria Rapp, 1829 (ICZN 1999; Art. 57.2), and Calder (1988) proposed the binomen Z. warreni as a replacement name. The genus remained monospecific for many years, exclusively for Z. warreni, but it now includes five nominal species, viz. Zyzzyzus warreni Calder, 1988, the most widespread species; Zyzzyzus spongicolus (von Lendenfeld, 1884); Zyzzyzus calderi Petersen, 1990; Zyzzyzus floridanus Petersen, 1990; and Zyzzyzus robustus Petersen, 1990. Historically, Watson (1978) transferred the Australian coastal species Tubularia spongicola von Lendenfeld, 1884 to the genus Zyzzyzus, as Z. spongicolus (von Lendenfeld, 1884). Zyzzyzus calderi, a coastal species of Bermuda, was described by Petersen (1990) from part of the material previously described by Calder (1988) as Z. warreni. Zyzzyzus floridanus is also known from the north Atlantic region, but restricted to the southeastern coast of the United States of America (Petersen 1990). Lastly, Z. robustus is the only nominal species of the genus recorded from deep-water (Petersen 1990). Hirohito (1988: 24) suggested that Corymorpha iyoensis Yamada, 1959 should be included in Zyzzyzus. His proposal has not been adopted in subsequent papers, and we also prefer to exclude it from the genus. Studies of type material of C. iyoensis, or examinations of fertile hydroids of the species from Japan, are needed to clarify its taxonomic affinities. The systematic position of the genus has also been debated. Initially, Stechow (1923) included Zyzzyzus in the subfamily Corymorphinae Allman, 1872, within the family Tubulariidae. Kramp (1933) regarded Zyzzyzus as a synonym of Corymorpha M. Sars, 1835 at first, but later (Kramp 1949) regarded it as congeneric with Tubularia Linnaeus, 1758, after noting morphological and ecological similarities between the two (viz. association with sponges; perisarc structure; hydrorhiza not homologous with root filaments of Corymorpha; and production of an actinula larva in its life cycle). He therefore returned Zyzzyzus to the Tubulariidae. Millard (1975) kept Zyzzyzus in the Tubulariidae based mainly on the existence of an actinula. She nevertheless recognized resemblances of the genus with some species of Corymorphidae (viz. coenosarc of the hydrocaulus with longitudinal endodermal canals; basal portion of the hydranth thickened and forming a diaphragm; and tentacular endoderm separated from the hypostomial endoderm by a mesogleal lamella). She also interpreted the hydrorhizal processes of Zyzzyzus as adaptations for living embedded in sponges, and not as being homologous with the filaments of corymorphids. Millard���s arguments were followed by Watson (1978), Petersen (1979), Migotto & Silveira (1987), Hirohito (1988), and Bouillon (1985), noting that ���some characters from the Corymorphidae ��� are present in the species of Zyzzyzus. Calder (1988) presented alternative arguments about the taxonomic position of Zyzzyzus. Although acknowledging resemblances to Tubularia and Tubulariidae, he described the existence of a well-developed gastrodermic diaphragm (Calder 1988, fig. 38) between basal and hypostomial portions of the hydranth of Z. warreni which, associated with the solitary condition and both internal and external morphology of the hydrocaulus, were considered reasons for including the genus in the Corymorphidae. During the 1990 s, the taxonomic position of Zyzzyzus was finally discussed within a phylogenetic framework. Petersen (1990) considered both Tubulariidae and Corymorphidae to be referable to different superfamilies, Tubularioidea and Corymorphoidea. He (Petersen 1990: 121���122) proposed that the Corymorphidae had the following synapomorphies differentiating the family from tubulariids: ���direct development of polyp through encysted gastrula stage, [���] perisarc gelatinous, sheath-like, secreted from just under the aboral tentacles, [���] aboral end of hydrocaulus pointed, [���] statocysts present in the papillae or rooting filaments developed from the hydrocaulus of the hydroids���. Moreover, he noted that Tubulariidae had the following synapomorphies differentiating the family from corymorphids: ���development from egg over sterrogastrula [sic] to actinula, [���] hydranth vasiform, with broad base carrying the long aboral tentacles, and slender distal portion with short oral tentacles, [���] parenchymatic cushion under the aboral tentacles enclosed by mesolamella���. In common, both superfamilies would have (page 121) the ���development from egg to polyp without a planula stage���. This group was later supported in molecular analyses using mitochondrial 16 S and 28 S rDNA and was named Aplanulata, a putative clade uniting Hydridae with Candelabridae, Corymorphidae, and Tubulariidae (Collins et al. 2005, 2006). However, as noted by Collins et al. (2006: 111), many taxa have yet to be sampled (e.g. Acaulidae, Margelopsidae, Paracorynidae, and Tricyclusidae) to confirm the monophyly of Aplanulata and to understand the relationship among its families and genera (including Zyzzyzus). Distribution. North and south Atlantic, Indian, Pacific., Published as part of Campos, Carlos J. A., Marques, Antonio C. & Migotto, Alvaro E., 2007, A taxonomic revision of the genus Zyzzyzus Stechow, 1921 (Cnidaria: Hydrozoa: Tubulariidae), pp. 1-22 in Zootaxa 1627 on pages 2-5, DOI: 10.5281/zenodo.273951, {"references":["Stechow, E. (1921) Neue genera und species von hydrozoen und anderen evertebraten. Archiv fur Naturgeschichte, 87, 248 - 265.","Kramp, P. L. (1933) The Scoresby Sound Committee's 2 nd East Greenland Expedition in 1932 to King Christian IX's Land. Coelenterata, Ctenophora and Chaetognatha. Meddelelser om Gronland, 104 (11), 1 - 20.","Millard, N. A. H. (1975) Monograph on the Hydroida of southern Africa. Annals of the South African Museum, 68, 1 - 513.","Watson, J. E. (1978) New species and new records of Australian athecate hydroids. Proceedings of the Royal Society of Victoria, 90 (2), 301 - 314.","Petersen, K. W. (1979) Development of coloniality in Hydrozoa. In: Larwood, G. & Rosen, B. R. (Eds), Biology and Systematics of Colonial Animals. Academic Press, New York, pp. 105 - 139.","Bouillon, J. (1985) Essai de classification des hydropolypes-hydromeduses (Hydrozoa-Cnidaria). Indo-Malayan Zoology, 2 (1), 29 - 243.","Wedler, E. & Larson, R. (1986) Athecate hydroids from Puerto Rico and the Virgin Islands. Studies on Neotropical Fauna and Environment, 21 (1 - 2), 69 - 101.","Migotto, A. E. & Silveira, F. L. (1987) Hidroides (Cnidaria, Hydrozoa) do litoral sudeste e sul do Brasil: Halocordylidae, Tubulariidae e Corymorphidae. Iheringia, Serie Zoologia, 66, 95 - 115.","Calder, D. R. (1988) Shallow-water hydroids of Bermuda: the Athecatae. Royal Ontario Museum Life Sciences Contributions, 148, 1 - 107.","Petersen, K. W. (1990) Evolution and taxonomy in capitate hydroids and medusae. Zoological Journal of the Linnean Society, 100, 1 - 231.","Migotto, A. E. (1996) Benthic shallow-water hydroids (Cnidaria, Hydrozoa) of the coast of Sao Sebastiao, Brazil, including a checklist of Brazilian hydroids. Zoologische Verhandelingen, Leiden, 306, 1 - 125.","Neave, S. A. (1940) Nomenclator Zoologicus, Vo l. 4, A List of the Names of Genera and Subgenera in Zoology From the Tenth Edition of Linnaeus 1758 to the End of 1935. Zoological Society of London, London, 758 pp.","Warren, E. (1906) On Tubularia solitaria sp. nov., a hydroid from the Natal coast. Annals of the Natal Museum, 1, 83 - 96.","Rapp, W. (1829) Untersuchungen uber den Bau einiger Polypen des mittellandischen Meeres. Nova Acta Physico-Medica Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum, 14, 643 - 658.","von Lendenfeld, R. (1884) Das system der medusen. Zoologischer Anzeiger, 8, 425 - 429.","Stechow, E. (1923) Zur kenntnis der hydroidenfauna des mittelmeeres, amerikas und anderer gebiete. II. Teil. Zoologische Jahrbucher, Abteilung fur Systematik, Okologie und Geographie der Tiere, 47, 29 - 270.","Allman, G. J. (1872) A Monograph of the Gymnoblastic or Tubularian Hydroids, Conclusion of Part I and Part II, Containing Descriptions of the Genera and Species of the Gymnoblastea. Ray Society, London, 450 pp.","Kramp, P. L. (1949) Origin of the hydroid Family Corymorphidae. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kobenhavn, 111, 183 - 215.","Collins, A. G., Winkelman, S., Hadrys, H. & Schierwater, B. (2005) Phylogeny of Capitata and Corynidae (Cnidaria, Hydrozoa) in light of mitochondrial 16 S rDNA data. Zoologica Scripta, 34, 91 - 99.","Collins, A. G., Schuchert, P., Marques, A. C., Jankowski, T., Medina, M. & Schierwater, B. (2006) Medusozoan phylogeny and character evolution clarified by new large and small subunit rDNA data and an assessment of the utility of phylogenetic mixture models. Systematic Biology, 55 (1), 97 - 115."]}

Acryptolaria gracilis (Allman, 1888) Cryptolaria gracilis: Farquhar 1896, p 461 (non-record; reference to Allman's record). Cryptolaria gracilis: Hutton 1904, p 320 (non-record). Lafoea (Cryptolaria) gracilis: Billard 1906b, p 330 (doubtful record; neither description nor figures. Mazagham, Morocco). Lafoea gracilis: Billard 1906c, p 157, 178 [doubtful record; neither description nor figures. Mazagham (5 El Jadida), Morocco, 33 ° 119N, 11 ° 199W, 550 m (5 Acryptolaria conferta minor after Ramil and Vervoort 1992, p 43)]. Osaealdaria gracilis: Stechow 1923a, p 147 (non-record; change of genus). Acryptolaria gracilis: Ralph 1958, p 314, Figure 3 b–d (doubtful record; unspecified locality). Acryptolaria gracilis: Patriti 1970, p 30, Figure 34 (same locality as the record by Billard 1906c). Acryptolaria gracilis: Dawson 1992, p 15 (non-record). Cryptolaria gracilis: Stranks 1993, p 7 (non-record; catalogue). Acryptolaria gracilis: Watson and Vervoort 2001, p 157 –159, Figure 4 a–c (doubtful record, considered so by the authors. Clearly different from A. gracilis. Tasmanian seamounts, 44.19– 44.22 ° S, 147.02– 147.05 ° E, 936–1018 m). Acryptolaria gracilis: Vervoort and Watson 2003, p 46 –47, Figure 6D, E (non-record; revision of the type material)., Published as part of Peña Cantero, Alvaro L., Marques, Antonio C. & Migotto, Alvaro E., 2007, Revision of the genus Acryptolaria Norman, 1875 (Cnidaria, Hydrozoa, Lafoeidae), pp. 229-291 in Journal of Natural History 41 (5 - 8) on pages 288-289, DOI: 10.1080/00222930701228132, http://zenodo.org/record/4582056

Acryptolaria angulata (Bale, 1914) Acryptolaria angulata: Blackburn 1942, p 111 (non-record; systematic list). Acryptolaria angulata: Vervoort and Watson 2003, p 41 –43, Figure 3 C–F (doubtful record. Probably the record includes material of Bale's species, but according to the figures material of other species too. Far offshore of New Zealand: Chatham Rise, Hikurangi Trough and near Kermadec Ridge, 145–913 m). Cryptolaria angulata: Stranks 1993, p 7 (non-record; catalogue of recent cnidarian types in the Museum of Victoria)., Published as part of Peña Cantero, Alvaro L., Marques, Antonio C. & Migotto, Alvaro E., 2007, Revision of the genus Acryptolaria Norman, 1875 (Cnidaria, Hydrozoa, Lafoeidae), pp. 229-291 in Journal of Natural History 41 (5 - 8) on page 281, DOI: 10.1080/00222930701228132, http://zenodo.org/record/4582056

Acryptolaria corniformis Naumov and Stepanjants, 1962 Acryptolaria corniformis: Gravier-Bonnet 1979, p 22 (non-record, discussion of the species). Acryptolaria corniformis: Vervoort and Watson 2003, p 44, Figure 4D, E (doubtful record, clearly belonging to a different species. Macquarie Gap, 51 ° 06.009S, 167 ° 48.509E, 216 m)., Published as part of Peña Cantero, Alvaro L., Marques, Antonio C. & Migotto, Alvaro E., 2007, Revision of the genus Acryptolaria Norman, 1875 (Cnidaria, Hydrozoa, Lafoeidae), pp. 229-291 in Journal of Natural History 41 (5 - 8) on page 286, DOI: 10.1080/00222930701228132, http://zenodo.org/record/4582056

Acryptolaria operculata Stepanjants, 1979 Acryptolaria operculata: Blanco 1994a, p 162 (non-record; identification key). Acryptolaria operculata: Blanco 1994b, p 188 –189 (non-record; reference to other authors). Acryptolaria operculata: Blanco et al. 1994, p 10 –12, Figures 7, 8 (non-record; repetition of Stepanjants' description). Acryptolaria operculata: Genzano and Zamponi 1997, p 290 (non-record; list of species)., Published as part of Peña Cantero, Alvaro L., Marques, Antonio C. & Migotto, Alvaro E., 2007, Revision of the genus Acryptolaria Norman, 1875 (Cnidaria, Hydrozoa, Lafoeidae), pp. 229-291 in Journal of Natural History 41 (5 - 8) on page 290, DOI: 10.1080/00222930701228132, http://zenodo.org/record/4582056

Acryptolaria flabellum (Allman, 1888) Osaealdaria flabellum: Stechow 1923a, p 147 (non-record; change of genus). Acryptolaria flabellum: Fraser 1944, p 212, Plate 41 Figure 191 (non-record; reference to Allman's material). Cryptolaria flabellum: Naumov 1957, p 41 (non-record). Cryptolaria flabellum: Naumov 1960, p 278 –279, Figure 168, Plate 1 Figure 2 (doubtful record; with description and figure. From the Sea of Okhotsk, 628–2400 m). Cryptolaria flabellum: Filatova and Barsanova 1964, p 90 (?). Acryptolaria flabellum: Vervoort 1968, p 99 (non-record; checklist—reference to other authors). Cryptolarella flabellum: Vervoort 1972, p 47 –49, Figure 13a, b [5 Cryptolarella abyssicola (Allman, 1888)]. Cryptolaria flabellate: Belousov 1975a, p 206 (non-record). Cryptolaria flabellum: Belousov 1975b, p 655, Figure 1 no. 1 (non-record). Cryptolarella cf. flabellum: Sheiko and Stepanjants 1997, p 439 (doubtful record; list of species. Commander Islands)., Published as part of Peña Cantero, Alvaro L., Marques, Antonio C. & Migotto, Alvaro E., 2007, Revision of the genus Acryptolaria Norman, 1875 (Cnidaria, Hydrozoa, Lafoeidae), pp. 229-291 in Journal of Natural History 41 (5 - 8) on page 288, DOI: 10.1080/00222930701228132, http://zenodo.org/record/4582056

Acryptolaria longitheca (Allman, 1877) Cryptolaria longitheca: Fewkes 1881, p 128 (doubtful record; neither description nor figures. Caribbean Sea: Dominica, 76 fathoms; Martinique, 334 fathoms; Barbados, 103 fathoms). Cryptolaria longitheca: Stechow 1913a, p 30 (non-record; reference to other authors concerning the coppinia). Lafoea (Cryptolaria) longitheca: Bonnevie 1899, p 63 (non-record; reference to the species). Lafoea (Cryptolaria) longitheca: Billard 1906c, p 178 (non-record; reference to the species). Osaealdaria longitheca: Stechow 1923a, p 147 (non-record; change of genus). Acryptolaria longitheca: Fraser 1943, p 78, 90, Plate 16 Figure 5, Plate 17 Figure 5 (doubtful record; with description and figures. With coppinia. 24 ° 189N, 80 ° 589300W, south of Florida Keys, 324 fathoms; 13 ° 119540N, 59 ° 389450W, off Barbados, 73 fathoms). Acryptolaria longitheca: Fraser 1944, p 212, Plate 41 Figure 192 (non-record; reference to previous records). Acryptolaria longitheca: Deevey 1954, p 270 (non-record; checklist—reference to other authors). Acryptolaria longitheca: Vervoort 1968, p 99 (non-record; checklist—reference to other authors). Acryptolaria longitheca: Vervoort 1972, p 45, Figure 12b, c (doubtful record. Closer to A. corniformis both in the shape and size of the hydrothecae. Western Atlantic, east of South Carolina, 32 ° 349N, 74 ° 21.59W, 4681 m). Acryptolaria longitheca: Cairns et al. 1991, p 24 (non-record; list of common and scientific names). Acryptolaria longitheca: Calder 1996, p 1723, Figure 2 d–f (doubtful record. Bermuda Pedestal, 32 ° 35.09N, 64 ° 54.99W, 3550 m; 32 ° 34.39N, 64 ° 54.79W, 3011 m, on hexactinellid sponges). Acryptolaria longitheca: Calder 1997, p 87 (non-record; reference to other authors). Acryptolaria longitheca: Calder and Vervoort 1998, p 22 –25, Figures 9–11 (doubtful record. Mid-Atlantic Ridge: 15 ° 35.339N, 46 ° 45.069W, 3410 m; 15 ° 28.569N, 46 ° 34.159W, 3000 m, coppinia in March, lacking modified hydrothecae; 15 ° 28.759N, 46 ° 33.569W, 3289 m; 15 ° 30.859N, 46 ° 38.249W, 3902 m; 23 ° 19.679N, 45 ° 14.659W, 2500 m). Acryptolaria cf. longitheca: Vervoort and Watson 2003, p 47, Figure 4F, G (doubtful record; closer to A. flabellum. 37 ° 15.209S, 176 ° 51.209E, Raukumara Plain, east of Coromandel Peninsula, New Zealand, 130 m). Acryptolaria longitheca: Vervoort 2006, p 229 –230, Figures 11e, 12a (doubtful record; off Azores, 37– 38 ° N, 25– 25.5 ° W, 1370–2850 m)., Published as part of Peña Cantero, Alvaro L., Marques, Antonio C. & Migotto, Alvaro E., 2007, Revision of the genus Acryptolaria Norman, 1875 (Cnidaria, Hydrozoa, Lafoeidae), pp. 229-291 in Journal of Natural History 41 (5 - 8) on pages 289-290, DOI: 10.1080/00222930701228132, http://zenodo.org/record/4582056

Peña Cantero, Alvaro L., Marques, Antonio C., Migotto, Alvaro E. (2007): Revision of the genus Acryptolaria Norman, 1875 (Cnidaria, Hydrozoa, Lafoeidae). Journal of Natural History 41 (5-8): 229-291, DOI: 10.1080/00222930701228132, URL: http://dx.doi.org/10.1080/00222930701228132

Lafoeina amirantensis (Millard & Bouillon, 1973) (Figs. 1–3; Tables 1–3) Egmundella amirantensis Millard & Bouillon, 1973: 40; Millard, 1975: 133; Gibbons & Ryland, 1989: 389 –390; Ramil & Vervoort, 1992: 22 –24. Lafoeina amirantensis: Calder 1991 a: 10; Calder 1991 b: 2068; Calder 1991 c: 223; Nogueira et al., 1997: 368; Calder & Maÿal, 1998: 73; Calder, 1998: 1852; Calder & Vervoort, 1998: 15; Watson, 2000: 5; Calder et al., 2003: 1180. Holotype: Musée Royal de l’Afrique Centrale– MRAC 2665, Egmundella amirantensis Millard & Bouillon, 1973, colony on Dynamena disticha (Bosc), Amirante, Seychelles. Material examined. MZUSP 1409, 23º 49.69 ’S 045º 24.76 ’W, Farol dos Moleques, São Sebastião, Brazil, 3–5m, colony on calcareous algae, 18 February 2001; MZUSP 1410, same data as MZUSP 1409, two 1 ­day­old medusae, 22 February 2001; MZUSP 1411, same data as MZUSP 1409, five 1 ­day­old medusae, 22 February 2001; MZUSP 1412, 23º 46 ’S 045º 40 ’W, Praia da Baleia, São Sebastião, Brazil, colony on calcareous algae and Sargassum sp., 1m, 16 July 2001; Praia de Barequeçaba, São Sebastião, Brazil, colony on calcareous algae, 4 February 2002. Description of polyp stage. Colonies with stolonal growth; perisarc smooth throughout, without annuli (Fig. 1). Hydrothecae and nematophores widely separated from each other, both arising directly from a branching and anastomosing hydrorhiza. Hydrorhiza 16–40 µm in diameter, without internal septa. Hydrotheca 110–250 µm high, 54–110 wide, thin­walled, smooth, tubular, without pedicel, constricted basally and slightly enlarged near base of operculum (Fig. 1 D, E). Operculum thin­walled, not delimited from hydrotheca by a crease line; not segmented, a continuation of the hydrothecal wall which folds upon itself forming 10–14 pleats, becoming conical when hydranth retracts within hydrotheca (Fig. 1 E–G). No visible difference between operculum and hydrotheca wall when hydranth extended. Hydranth extensile, 2.0–4.0 times higher than hydrotheca when fully extended, with 9–14 slender tentacles, no intertentacular web. Basitrichous isorhiza (Fig. 2 C) capsules arranged in circular bands evenly spaced along the length of the tentacle; atrichous isorhiza (Fig. 2 B) on body and tentacles. Tentacles amphicoronate, those of upper whorl 180–380 µm in length, held at 45–60 º angles, longer than those of lower whorl (110–220 µm) which are held horizontally or slightly downward (Fig. 1 A). Hypostome conical. Nematotheca (Fig. 1 B) 30–63 µm high, 15–32 µm in diameter, club­shaped, with a bundle of large, elongated microbasic mastigophore nematocysts (Fig. 2 F) inside. Hydranth column with basitrichous isorhiza, atrichous isorhiza, and scarce microbasic mastigophores identical with the ones present on nematophores. Gonothecae similar to hydrothecae. Character variation of polyp stage is presented in Table 1. Nematocyst size and distribution are shown in Table 2. mean ± SD (range) (n) Hydrotheca Height from base to tip of operculum (m) 188.5 33.9 (110–250) (38) Height from base to margin (m) 148.5 34.5 (90–235) (33) Maximum diameter (m) 72.0 11.4 (54–110) (45) Diameter at diaphragm (m) 41.9 8.5 (23–60) (27) Hydranth Height from diaphragm to tip of hypostome (m) 550.6 115.3 (415–740) (10) Diameter of column (m) 36.0 5.4 (30–50) (14) Diameter at tentacular ring (m) 58.3 15.6 (42–100) (11) Diameter at diaphragm (m) 32.7 11.0 (20–40) (3) Height of hypostome 22.7 1.5 (20–24) (6) Number of tentacles 11 1 (9–14) (21) Length of upward tentacles (m) 259.2 56.9 (180–380) (20) Length of downward tentacles (m) 158.5 33.8 (110–220) (20) Diameter of tentacles (m) 16.2 11.5 (10–50) (13) Nematotheca Height (m) 44.0 7.8 (30–63) (25) Diameter (m) 18.8 3.7 (15–32) (25) Diameter of hydrorhiza (m) 28.5 8.0 (16–40) (27) Description of Medusa. Medusa released in immature condition, transparent, hemispherical, about 0.2–0.4 mm high, slightly wider than high (Fig. 3 A). Mesoglea thin, without apical process and apical canal; exumbrella with scattered nematocysts (microbasic mastigophores, Fig. 2 A). Radial canals four, simple and straight; circular canal narrow. Velum broad, aperture circular. Two opposed perradial marginal bulbs, pyriform or triangular, brown­pigmented, with tentacles (Fig. 3 B). Other perradial bulbs inconspicuous or absent, a few nematocysts marking the site of the future tentacular bulb. Tentacles hollow, filiform, tapering gradually, with two types of nematocysts: basitrichous isorhiza (Fig. 2 D) and atrichous isorhiza (Fig. 2 F). Tentacle tip without terminal swelling. Four interradial statocysts, each with a single statolith (Fig. 3 B–D). With one or two marginal cirri on the sides of each statocyst (Fig. 3 B–D). Cirrus solid, with scattered nematocysts. Lateral cirri absent. Manubrium tubular, short, cruciform in cross­section. Mouth simple, with four oral lips. Stomach broad, quadrate; peduncle lacking (Fig. 3 E). Four incipient gonads on the proximal portion of the radial canals, near stomach. Medusae grew relatively fast; some individuals (Fig. 3 F–G.) had doubled their size after one day. Noteworthy was the development of the other pair of opposed perradial tentacles, which at this stage were shorter and with smaller bulbs than the former two (Fig. 3 F). Tentacles reached 12 mm when fully extended and cirri about 0.4 mm. The number of cirri remained variable, increasing in number (4–8). Exumbellar nematocysts were still present but fewer than in the newly liberated medusae. The gonads remained underdeveloped, but were more visible. The medusae did not accept the copepods and Artemia nauplii offered and died a few days later; during this time no significant morphological changes were seen. Measurements of many young medusae are in Table 3. Nematocysts size and distribution are presented on Table 2. Remarks. SEM images (Fig. 1 E) revealed that the hydrothecal wall has shallow, longitudinal depressions extending from the base of operculum downward to approximately the middle of the hydrotheca. These features are, however, not seen under low or high power light microscopy. Medusae were liberated two and three days after their parent colonies were collected, respectively, at Farol dos Moleques in 18 February 2001 and Praia de Barequeçaba on 4 February 2002. On both occasions gonothecae were noticed only when empty, probably because they resemble hydrothecae in shape and size (as documented in the description and figures of Millard and Bouillon 1973). Care was taken to ensure that other species of hydroids were not present in the culture bowls. After the pieces of colonies were tied on glass slides, the hydrorhiza grew along the glass substratum; a few days later ramifying sparingly and giving rise to new hydranths at regular intervals of about 1–2 mm. Nematophores were scarce or even absent on some sections of the hydrorhiza and often they were covered by debris or unicellular algae. In one of the cultures, the terminal regions of the hydrorhizae gave rise to free planuloids or frustules (Fig. 1 C). These planula­like bodies were flattened, 400–800 µm long, 50–75 µm wide, and had the same nematocyst types as polyps (microbasic mastigophores, basitrichous isorhizas, and atrichous isorhizas). Planuloids lacked external cilia and crept over the glass surface by means of peristaltic waves. Three to four days after release, some planuloids stopped moving, settled onto their sides, and secreted a perisarc. Within about 12 h, primary hydranths with 8–10 tentacles were formed. Additional hydranths (up to three were observed) appeared singly from unbranched hydrorhiza. Nematophores did not develop. The process of formation of planuloids in L. amirantensis is similar to the ones reported for, e.g., Coryne, by Berrill 1953, and Obelia, by Allman 1871, and is usually elicited as a response to rapid changes in temperature (Berrill 1953, Panteleeva 1999). Although with limited means of locomotion which results in settlement near the parent colony, the planuloids increase the possibilities of dispersal of the species.

Lafoeina amirantensis (Millard & Bouillon, 1973) (Figs. 1���3; Tables 1���3) Egmundella amirantensis Millard & Bouillon, 1973: 40; Millard, 1975: 133; Gibbons & Ryland, 1989: 389 ���390; Ramil & Vervoort, 1992: 22 ���24. Lafoeina amirantensis: Calder 1991 a: 10; Calder 1991 b: 2068; Calder 1991 c: 223; Nogueira et al., 1997: 368; Calder & Ma��al, 1998: 73; Calder, 1998: 1852; Calder & Vervoort, 1998: 15; Watson, 2000: 5; Calder et al., 2003: 1180. Holotype: Mus��e Royal de l���Afrique Centrale��� MRAC 2665, Egmundella amirantensis Millard & Bouillon, 1973, colony on Dynamena disticha (Bosc), Amirante, Seychelles. Material examined. MZUSP 1409, 23�� 49.69 ���S 045�� 24.76 ���W, Farol dos Moleques, S��o Sebasti��o, Brazil, 3���5m, colony on calcareous algae, 18 February 2001; MZUSP 1410, same data as MZUSP 1409, two 1 ��day��old medusae, 22 February 2001; MZUSP 1411, same data as MZUSP 1409, five 1 ��day��old medusae, 22 February 2001; MZUSP 1412, 23�� 46 ���S 045�� 40 ���W, Praia da Baleia, S��o Sebasti��o, Brazil, colony on calcareous algae and Sargassum sp., 1m, 16 July 2001; Praia de Bareque��aba, S��o Sebasti��o, Brazil, colony on calcareous algae, 4 February 2002. Description of polyp stage. Colonies with stolonal growth; perisarc smooth throughout, without annuli (Fig. 1). Hydrothecae and nematophores widely separated from each other, both arising directly from a branching and anastomosing hydrorhiza. Hydrorhiza 16���40 ��m in diameter, without internal septa. Hydrotheca 110���250 ��m high, 54���110 wide, thin��walled, smooth, tubular, without pedicel, constricted basally and slightly enlarged near base of operculum (Fig. 1 D, E). Operculum thin��walled, not delimited from hydrotheca by a crease line; not segmented, a continuation of the hydrothecal wall which folds upon itself forming 10���14 pleats, becoming conical when hydranth retracts within hydrotheca (Fig. 1 E���G). No visible difference between operculum and hydrotheca wall when hydranth extended. Hydranth extensile, 2.0���4.0 times higher than hydrotheca when fully extended, with 9���14 slender tentacles, no intertentacular web. Basitrichous isorhiza (Fig. 2 C) capsules arranged in circular bands evenly spaced along the length of the tentacle; atrichous isorhiza (Fig. 2 B) on body and tentacles. Tentacles amphicoronate, those of upper whorl 180���380 ��m in length, held at 45���60 �� angles, longer than those of lower whorl (110���220 ��m) which are held horizontally or slightly downward (Fig. 1 A). Hypostome conical. Nematotheca (Fig. 1 B) 30���63 ��m high, 15���32 ��m in diameter, club��shaped, with a bundle of large, elongated microbasic mastigophore nematocysts (Fig. 2 F) inside. Hydranth column with basitrichous isorhiza, atrichous isorhiza, and scarce microbasic mastigophores identical with the ones present on nematophores. Gonothecae similar to hydrothecae. Character variation of polyp stage is presented in Table 1. Nematocyst size and distribution are shown in Table 2. mean �� SD (range) (n) Hydrotheca Height from base to tip of operculum (m) 188.5 33.9 (110���250) (38) Height from base to margin (m) 148.5 34.5 (90���235) (33) Maximum diameter (m) 72.0 11.4 (54���110) (45) Diameter at diaphragm (m) 41.9 8.5 (23���60) (27) Hydranth Height from diaphragm to tip of hypostome (m) 550.6 115.3 (415���740) (10) Diameter of column (m) 36.0 5.4 (30���50) (14) Diameter at tentacular ring (m) 58.3 15.6 (42���100) (11) Diameter at diaphragm (m) 32.7 11.0 (20���40) (3) Height of hypostome 22.7 1.5 (20���24) (6) Number of tentacles 11 1 (9���14) (21) Length of upward tentacles (m) 259.2 56.9 (180���380) (20) Length of downward tentacles (m) 158.5 33.8 (110���220) (20) Diameter of tentacles (m) 16.2 11.5 (10���50) (13) Nematotheca Height (m) 44.0 7.8 (30���63) (25) Diameter (m) 18.8 3.7 (15���32) (25) Diameter of hydrorhiza (m) 28.5 8.0 (16���40) (27) Description of Medusa. Medusa released in immature condition, transparent, hemispherical, about 0.2���0.4 mm high, slightly wider than high (Fig. 3 A). Mesoglea thin, without apical process and apical canal; exumbrella with scattered nematocysts (microbasic mastigophores, Fig. 2 A). Radial canals four, simple and straight; circular canal narrow. Velum broad, aperture circular. Two opposed perradial marginal bulbs, pyriform or triangular, brown��pigmented, with tentacles (Fig. 3 B). Other perradial bulbs inconspicuous or absent, a few nematocysts marking the site of the future tentacular bulb. Tentacles hollow, filiform, tapering gradually, with two types of nematocysts: basitrichous isorhiza (Fig. 2 D) and atrichous isorhiza (Fig. 2 F). Tentacle tip without terminal swelling. Four interradial statocysts, each with a single statolith (Fig. 3 B���D). With one or two marginal cirri on the sides of each statocyst (Fig. 3 B���D). Cirrus solid, with scattered nematocysts. Lateral cirri absent. Manubrium tubular, short, cruciform in cross��section. Mouth simple, with four oral lips. Stomach broad, quadrate; peduncle lacking (Fig. 3 E). Four incipient gonads on the proximal portion of the radial canals, near stomach. Medusae grew relatively fast; some individuals (Fig. 3 F���G.) had doubled their size after one day. Noteworthy was the development of the other pair of opposed perradial tentacles, which at this stage were shorter and with smaller bulbs than the former two (Fig. 3 F). Tentacles reached 12 mm when fully extended and cirri about 0.4 mm. The number of cirri remained variable, increasing in number (4���8). Exumbellar nematocysts were still present but fewer than in the newly liberated medusae. The gonads remained underdeveloped, but were more visible. The medusae did not accept the copepods and Artemia nauplii offered and died a few days later; during this time no significant morphological changes were seen. Measurements of many young medusae are in Table 3. Nematocysts size and distribution are presented on Table 2. Remarks. SEM images (Fig. 1 E) revealed that the hydrothecal wall has shallow, longitudinal depressions extending from the base of operculum downward to approximately the middle of the hydrotheca. These features are, however, not seen under low or high power light microscopy. Medusae were liberated two and three days after their parent colonies were collected, respectively, at Farol dos Moleques in 18 February 2001 and Praia de Bareque��aba on 4 February 2002. On both occasions gonothecae were noticed only when empty, probably because they resemble hydrothecae in shape and size (as documented in the description and figures of Millard and Bouillon 1973). Care was taken to ensure that other species of hydroids were not present in the culture bowls. After the pieces of colonies were tied on glass slides, the hydrorhiza grew along the glass substratum; a few days later ramifying sparingly and giving rise to new hydranths at regular intervals of about 1���2 mm. Nematophores were scarce or even absent on some sections of the hydrorhiza and often they were covered by debris or unicellular algae. In one of the cultures, the terminal regions of the hydrorhizae gave rise to free planuloids or frustules (Fig. 1 C). These planula��like bodies were flattened, 400���800 ��m long, 50���75 ��m wide, and had the same nematocyst types as polyps (microbasic mastigophores, basitrichous isorhizas, and atrichous isorhizas). Planuloids lacked external cilia and crept over the glass surface by means of peristaltic waves. Three to four days after release, some planuloids stopped moving, settled onto their sides, and secreted a perisarc. Within about 12 h, primary hydranths with 8���10 tentacles were formed. Additional hydranths (up to three were observed) appeared singly from unbranched hydrorhiza. Nematophores did not develop. The process of formation of planuloids in L. amirantensis is similar to the ones reported for, e.g., Coryne, by Berrill 1953, and Obelia, by Allman 1871, and is usually elicited as a response to rapid changes in temperature (Berrill 1953, Panteleeva 1999). Although with limited means of locomotion which results in settlement near the parent colony, the planuloids increase the possibilities of dispersal of the species., Published as part of Migotto, Alvaro E. & Cabral, Andr�� S., 2005, Lafoeina amirantensis (Cnidaria: Hydrozoa, Campanulinoidea), the hydroid stage of the medusa Cirrholovenia tetranema (Cnidaria: Hydrozoa, Lovenelloidea), pp. 1-16 in Zootaxa 919 on pages 3-9, DOI: 10.5281/zenodo.171076, {"references":["Millard, N. A. H & Bouillon, J. (1973) Hydroids from the Seychelles (Coelenterata). Koninklijk Museum voor Midden-Afrika. 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Hydrobiologia, 216 / 217, 221 - 228.","Nogueira, C. C., Grohmann, P. A. & da Silva, V. M. A. P. (1997) Hydroids from the vicinity of a nuclear power plant site (CNAAA - Unidade 1) at Angra-dos-Reis, Rio de Janeiro, southeastern Brazil. In: den Hartog J. C. (Ed) Proceedings of the 6 th International Conference on Coelenterate Biology. Nationaal Natuurhistorisch Museum, Leiden, pp. 365 - 369.","Calder, D. R. & Mayal, E. M. (1998) Dry season distribution of hydroids in a small tropical estuary, Pernambuco, Brazil. Zoologische Verhandelingen, 323, 69 - 78.","Calder, D. R. & Vervoort, W. (1998) Some hydroids (Cnidaria: Hydrozoa) from the Mid-Atlantic Ridge, in the North Atlantic Ocean. Zoologische Verhandelingen, 319, 1 - 65.","Watson, J. E. (2000) Hydroids (Hydrozoa: Leptothecatae) from the Beagle Gulf and Darwin Harbour, northern Australia. The Beagle, Records of the Museum and Art Galleries of the Northern Territory, 16, 1 - 83.","Calder, D. R., Mallinson, J. J., Collins, K. & Hickman, C. P. (2003) Additions to the hydroids (Cnidaria) of the Galapagos, with a list of species from the islands. Journal of Natural History, 37, 1173 - 1218.","Berrill, N. J. (1953) Growth and form in gymnoblastic hydroids. VII. Growth and reproduction in Syncoryne and Coryne. Journal of Morphology, 92 (2), 273 - 302.","Allman, G. J. (1871) On a mode of reproduction of spontaneous fission in the hydroids. Quarterly Journal of microscopical science, n. s., 11, 18 - 21.","Panteleeva, N. N. (1999) Obelia longissima (Pallas, 1766) and Obelia geniculata (L., 1758) (Hydrozoa, Thecaphora, Campanulariidae) in the Barents Sea. Morphology, distribution, ecology and special life history features. Zoosystematica Rossica, suppl. 1, 51 - 65."]}

Marques, Antonio C., Peña Cantero, Alvaro L., Migotto, Alvaro E. (2005): Redescription and systematic status of the Antarctic genus Abietinella Levinsen, 1913 (Lafoeidae, Hydrozoa, Cnidaria). Journal of Natural History 39 (18): 1443-1455, DOI: 10.1080/00222930400005765, URL: http://dx.doi.org/10.1080/00222930400005765

Marques, Antonio C., Peña Cantero, Alvaro L., Migotto, Alvaro E. (2005): Revision of the genus Cryptolarella Stechow, 1913 (Lafoeidae, Leptothecata, Hydrozoa). Journal of Natural History 39 (9): 709-722, DOI: 10.1080/00222930400001467, URL: http://dx.doi.org/10.1080/00222930400001467