Altuna, Á., F.J. Murillo, and D.R. Calder. 2013. Aglaopheniid hydroids (Cnidaria: Hydrozoa: Aglaopheniidae) from bathyal waters of the Flemish Cap, Flemish Pass, and Grand Banks of Newfoundland (NW Atlantic). Zootaxa 3737: 501–537.

Altuna, Á., F.J. Murillo, and D.R. Calder. 2013. Aglaopheniid hydroids (Cnidaria: Hydrozoa: Aglaopheniidae) from bathyal waters of the Flemish Cap, Flemish Pass, and Grand Banks of Newfoundland (NW Atlantic). Zootaxa 3737: 501–537.

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   Accepted by A. Collins: 29 Oct.. 2013; published: 22 Nov. 2013    Licensed under a Creative Commons Attribution License ZOOTAXA ISSN 1175-5326 (print edition)ISSN   1175-5334   (online edition) Copyright © 2013 Magnolia Press Zootaxa  3737 (5): 501  –  537 / zootaxa  / Article   501 Aglaopheniid   hydroids (Cnidaria: Hydrozoa: Aglaopheniidae) from bathyal waters of the Flemish Cap, Flemish Pass, and Grand Banks of Newfoundland (NW Atlantic) ÁLVARO ALTUNA 1 , FRANCISCO J. MURILLO 2  & DALE R. CALDER  3 1  INSUB, Museo de Okendo, Zemoria, 12, Apartado 3223, 20013 Donostia-San Sebastián, Spain. E-mail:  2  Instituto Español de Oceanografía, Centro Oceanográfico de Vigo, Programa de Pesquerías Lejanas, Apartado 1552, 36280 Vigo, Spain. E-mail: 3  Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, Canada M5S 2C6.  E-mail: 1 Corresponding author Abstract Five species of aglaopheniid hydroids (  Aglaophenopsis cornuta , Cladocarpus diana , C. formosus , C. integer , and  Nema-tocarpus ramuliferus ) were collected from the Flemish Cap, Flemish Pass, and Grand Banks of Newfoundland during sur-veys with bottom trawls, rock dredges, and scallop gear. All are infrequently reported species, with C. diana  being discovered for the first time since its srcinal description from Iceland. We document here the southernmost collections of C. diana  and  N. ramuliferus , both previously unknown in the western Atlantic. Each of the five species is described and illustrated based on fertile material, a key is provided for their identification, and bathymetric distributions are noted. Known depth ranges are extended for  A. cornuta , C. diana , and C. integer .  Aglaophenopsis  and  Nematocarpus  are recog-nized as genera distinct from the polyphyletic Cladocarpus,  based on the unique structure of the phylactocarp in the for-mer, and the existence of appendages with nematothecae (ramuli) on almost all thecate internodes of hydrocladia in the latter. These appendages occur even in the absence of gonothecae, and are here considered defensive structures that protect the hydranths. In differing from typical phylactocarps, we accept the contention that they are characters of generic value. Key words:  Aglaophenopsis , Cladocarpus ,  Nematocarpus , deep-sea, northwestern Atlantic Introduction The hydrozoan family Aglaopheniidae Marktanner-Turneretscher, 1890 is a speciose taxon comprising around 248 valid species worldwide (see Bouillon et al  . 2006). Division of genera within the group is based mainly on the gonosome, because reliable differential characters are lacking in the trophosome (Leloup 1932). Aglaopheniids have been reported infrequently in the western Atlantic northwards of the 40ºN parallel, and especially so in waters off the east coast of Canada. While the hydroid fauna of shallow waters off the Atlantic coast of Canada is relatively well known (Fraser 1944), that of deeper neritic and bathyal waters has received little attention and is consequently poorly known. At such depths off Newfoundland, only two species of aglaopheniids have been recorded previously.  Aglaophenopsis cornuta  (Verrill, 1879) was reported from two locations north of the island (53°34’N–52°01’W, ca. 1792 m; 52°25’N–52°12’W, ca. 296 m) by Jäderholm (1919: 11), while  A. cornuta  and Cladocarpus integer  (G.O. Sars, 1874) were found in collections from a station to the northeast of Newfoundland (51°22’30”N–50°31’30”W, 457 m) by Calder (1970: 1540). Two other aglaopheniid species penetrating into bathyal waters,  Lytocarpia myriophyllum  (Linnaeus, 1758) and Cladocarpus formosus  Allman, 1874, range both north and south of  Newfoundland along the North American Atlantic coast (Fraser 1944, 1946) but have yet to be reported from that locale.  ALTUNA  ET AL. 502   ·   Zootaxa  3737 (5) © 2013   Magnolia Press FIGURE 1 . Distribution of the five aglaopheniid species in the study area. A)  Aglaophenopsis cornuta  (Verrill, 1879). B) Cladocarpus diana  Broch, 1918. C) Cladocarpus formosus Allman, 1874. D) Cladocarpus integer  (G.O. Sars, 1874). E)  Nematocarpus ramuliferus  (Allman, 1874). FC, Flemish Cap; FP, Flemish Pass; GB, Grand Banks. In a literature review, we found records of only eight species of aglaopheniids, referable to the genera  Aglaophenopsis  Fewkes, 1881, Cladocarpus  Allman, 1874, and  Lytocarpia  Kirchenpauer, 1872, in the region between 40ºN (New York Bight east of New Jersey) and Baffin Bay (including the west coast of Greenland) (Table 1). Some have been recorded only a few times and appear to be quite rare, while another is probably not valid. Scarcity of aglaopheniids in the northwestern North Atlantic may be attributed in part to the fact that they are predominantly a warm-water group (Calder 1997a: 40).    Zootaxa  3737 (5) © 2013 Magnolia Press   ·   503 AGLAOPHENIID HYDROIDS FROM THE NW ATLANTIC  ALTUNA  ET AL. 504   ·   Zootaxa  3737 (5) © 2013   Magnolia Press According to the generic diagnoses of aglaopheniids in Bouillon (1985) and Bouillon et al.  (2006), all species of the family known to occur between Baffin Bay and the Maritime Provinces of Canada are referable to Cladocarpus , with the exception of  Lytocarpia myriophyllum . Cladocarpus  was srcinally established by Allman (1874) to accommodate C. formosus , a species first described from the Faroe-Shetland Channel in the northeastern  North Atlantic. Its primary distinguishing characters, according to Allman, were differences in morphology of the reproductive structures: gonothecae were solitary and protected by phylactocarps (‘gonangia not included in corbulae’) arising as appendages of an unmodified hydrocladium. Bouillon et al  . (2006) listed 66 species of Cladocarpus  worldwide, most of them from deep-water. However, the genus has essentially become a collective group, and a revision is needed. Some genera merged with it appear distinctive in characters of their phylactocarps and, in our opinion, are valid (see also Ramil & Vervoort 1992). One of these is  Aglaophenopsis , with several representatives in the western North Atlantic. Fraser (1944) included four species in that genus from the Atlantic coast of North America, with three of them being seldom recorded and poorly known.During 2006–2010, the Spanish Institute of Oceanography, in collaboration with several other institutions, carried out a number of research surveys on the Flemish Cap, the Flemish Pass, and the “tail” and “nose” of the Grand Banks of Newfoundland (Figure 1). Abundant material of sessile benthic fauna was collected over a wide range of depths, mainly in the bathyal zone (see Murillo et al  . 2011). Hydroids comprise a significant part of those catches, and numerous colonies of aglaopheniids referable to five species were collected. All five are described and illustrated in this paper. Two of them are rare and previously unknown in the western Atlantic, one of which is reported for the first time since its srcinal description. The generic value of certain gonosomal characters is discussed, with special emphasis on the genera  Aglaophenopsis  and  Nematocarpus  Broch, 1918. Material and methods  Material examined during this study came from two research programs. Part was obtained during groundfish bottom trawl surveys carried out by the Spanish Institute of Oceanography (Instituto Español de Oceanografía, IEO) and the European Union (EU) on board the Spanish R/V “Vizconde de Eza”. The remainder came from rock dredge and scallop gear samples taken by the Spanish R/V “Miguel Oliver,” operated by the Spanish General Secretariat of the Sea (Secretaría General del Mar) under the NEREIDA project. Groundfish bottom trawl surveys covered the “tail” of the Grand Banks of Newfoundland between 40 and 1500 m, the Flemish Cap between 130 and 1450 m, and the Flemish Pass between 110 and 1450 m. These surveys were carried out annually between spring and summer using a random stratified sampling design with standardized 30-min tows and vessel speed of three knots. Campelen 1800 bottom trawl gear was used in the Flemish Pass and the Grand Banks of Newfoundland, whereas Lofoten bottom trawl gear was used at Flemish Cap. NEREIDA surveys were undertaken in the Flemish Pass, Flemish Cap, and slope of the Grand Banks of Newfoundland at depths between 700 and 2000 m. These surveys were carried out during spring and summer using a rock dredge and scallop gear. Tows of 15-min were made at a vessel speed of about 1.5 knots. NEREIDA is a multidisciplinary research project involving scientists from Instituto Español de Oceanografía (IEO), Fisheries and Oceans Canada (DFO), Natural Resources Canada (NRCAN), Centre for the Environment, Fisheries and Aquaculture Science (Cefas), Instituto de Investigaciones Marinas (CSIC), Secretaría General del Mar (SGM), Polar Research Institute of Marine Fisheries and Oceanography (PINRO), and the P.P. Shirshov Institute of Oceanology (RAS). Our study is based on material from groundfish bottom trawl surveys undertaken on the “tail” of the Grand Banks and Flemish Cap during 2007 and on the “nose” of the Grand Banks and Flemish Pass from 2006–2010, as well as from NEREIDA surveys during 2009 and 2010. Samples were fixed on board in 70 % ethanol. Voucher material of each species has been deposited in collections of the Invertebrate Zoology Section, Department of Natural History, Royal Ontario Museum (Canada), the Oceanographic Center of Vigo (IEO, Spain), and the Okendo Museum, Donostia-San Sebastián (Spain).Depths in Levinsen (1893) and Broch (1918) were given in Danish fathoms (  favne ), equivalent to 1.883 m. Fathoms in other papers have been taken to equal 1.828 m.    Zootaxa  3737 (5) © 2013 Magnolia Press   ·   505 AGLAOPHENIID HYDROIDS FROM THE NW ATLANTIC TABLE 2 . Species of aglaopheniid hydroids reported from the western North Atlantic between 40°N (New York Bight) and Baffin Bay (including western and southwestern Greenland) but not recorded during the present study. Literature records are not intended to be exhaustive. ResultsClass Hydrozoa Owen, 1843Subclass Leptolina Haeckel, 1879 Order Leptothecata Cornelius, 1992Suborder Conica Broch, 1910Family Aglaopheniidae Marktanner-Turneretscher, 1890  Aglaophenopsis cornuta  (Verrill, 1879) (Figs. 1A, 2A–G, tables 1, 3, 9–12) Cladocarpus cornutus  Verrill, 1879: 310.—Levinsen 1893: 208, pl. 8, fig. 19–22.—Vanhöffen 1897: 246.—Jäderholm 1909: 110.—Kramp 1913: 28.—Kramp 1914: 1059.—Fraser 1946: 417.—Calder 1970: 1540, pl. 8, fig. 7.—Peña Cantero & García Carrascosa 1999: 214.—Schuchert 2001: 137, fig. 117A–E.—Bouillon et al  . 2006: 283.  Aglaophenia cornutus : Bonnevie 1899: 94.  Aglaophenopsis cornuta : Nutting 1900: 120, pl. 30, fig. 6–9.—Whiteaves 1901: 28.—Broch 1910: 234.—Kindle & Whittaker 1917: 232.—Broch 1918: 77, fig. 39a–b, fig. 40a–d, fig. 41.—Jäderholm 1919: 11.—Fraser 1921: 177, fig. 102.—Kramp 1932a: 56.—Kramp 1932b: 20.—Kramp 1943: 44.—Fraser 1944: 395, pl. 85, fig. 382.—Vervoort 1972: 204.—Calder 1997b: 88.—Henry 2001: 163. Material examined . FN3L06 L51, 2 fertile colonies, largest one 9.5 cm high; FN3L06 L85, one fertile colony 7.0 cm high; FN3L06 L88, one fertile colony 10.5 cm high; FN3L06 L92, one fertile colony 15.0 cm high; PLA07 L61, one sterile colony 8.4 cm high; PLA07 L99, one sterile colony 10.0 cm high; PLA07 L107, one fertile colony 6 cm high; FC07 L74, one sterile colony 7.2 cm high; FC07 L133, one sterile colony 4.3 cm high; FC07 L153, one sterile colony 13.0 cm high; FC07 L171, one sterile fragment 1.6 cm high; FC07 L177, one sterile colony 5.5 cm  Aglaophenopsis bonnevieae  (Jäderholm, 1909): 60º16’5N–47º48’W and 60º22’N–47º27’W (southwest Greenland, 135 m and 120 m) [Kramp 1932a, as  Aglaophenopsis compressa (Bonnevie, 1899)]. No further records in the western Atlantic.  Aglaophenopsis verrilli  Nutting, 1900: 40º34’N–66º04’W (off Georges Bank, 3184 m) [Nutting (1900) and Fraser (1944)] = Cladocarpus verrilli  in Bouillon et al  . (2006) and Schuchert (2012b). Cladocarpus campanulatus  Ritchie, 1912: 60º22’N–47º27’W (southwest Greenland, 120 m) (Kramp 1932a). No further records in the western Atlantic. Cladocarpus crenatus  (Fewkes, 1881): 41º25’N–65º35’W (2270 m) [Fewkes (1881), as  Aglaophenia crenata )], Nutting (1900) as  Aglaophenia  ? crenata referring to Fewkes (1881), east of Nantucket (Fraser 1944), referring to Fewkes (1881), as Cladocarpus  ? crenatus  with unknown gonosome. Likely coterminous with Cladocarpus    formosus . Cladocarpus flexilis  Verrill, 1885: southern coast of New England (Verrill 1885), seven stations in the Georges Bank area at different depths (  Fish Hawk   and  Albatross  stations) (Nutting 1900), Martha’s Vineyard and several stations in the Georges Bank area (Fraser 1944, some stations referring to Nutting 1900).  Lytocarpia myriophyllum  (L., 1758): Massachusetts Bay, Mingan Islands (Quebec), Eastport, Maine (A. Agassiz 1865 as Sertularia myriophyllum ; Whiteaves 1901 and Fraser 1918 as Thecocarpus myriophyllum ), 41°44’N–64°36’W (Le Have Bank, Nova Scotia, ca. 110 m) (Smith & Harger 1874 as  Aglaophenia myriophyllum ; Whiteaves 1901 and Fraser 1918 as T. myriophyllum ), off Cape Gaspé (Gulf of St. Lawrence, Quebec, ca. 55 m) (Whiteaves 1901 and Fraser 1918 as T. myriophyllum ), eastern Canada [(Kindle & Whittaker 1917, ca. 55–110 m, as T. myriophyllum , based on literature records)].
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