Cah. Biol. Mar. (2004) 45 : 313-324

Cladoplumaria anomala gen. nov., sp. nov., a new and species of the family (: ) from the Chesterfield Islands region (Pacific Ocean)

Jose A. ANSÍN AGÍS1, Francisco RAMIL1 and Willem VERVOORT2 (1) Departamento de Ecoloxía e Bioloxía , Facultad de Ciencias, Universidade de Vigo, Campus Lagoas- Marcosende, 36200 Vigo, Spain. E-mail: [email protected] and [email protected] (2) Nationaal Natuurhistorisch Museum, P.O. Box 9517, 2300 RA, Leiden, The Netherlands. E-mail: [email protected]

Abstract: This paper is the first result of the study of large collections of (Cnidaria, Hydrozoa, Leptolida) collected in the seas surrounding New Caledonia (Pacific Ocean) by French expeditions. A new genus of the family Halopterididae is described and its relations with other representatives of this family are discussed. The gonosome in its shape approaches the type of phylactocarp also met with in Aglaopheniidae; while the gonothecae (female with nematothe- cae and male without nematothecae) are nearer to those of the family Halopterididae.

Résumé : Cladoplumaria anomala gen. nov. sp. nov., un nouveau genre et une nouvelle espèce de la famille des Halopterididae (Cnidaria, Hydrozoa) des Iles Chesterfield (océan Pacifique). Dans ce travail, nous présentons les premiers résultats de l’étude d’une importante collection d’Hydroïdes appartenant à la superfamille des Plumularioidea (Cnidaria, Hydrozoa, Leptolida) recueillis par des expéditions françaises dans la région de la Nouvelle Calédonie (Océan Pacifique). Un nouveau genre de la famille des Halopterididae est décrit, et ses relations avec les autres genres de la famille sont dis- cutées. La présence de phylactocarpes protégeant les gonothèques est une caractéristique typique des Aglaopheniidae, mais la présence de nématothèques sur les gonothèques, au moins sur les femelles, est un caractère propre de la famille des Halopterididae.

Keywords: Cnidaria, Hydrozoa, Leptolida, Cladoplumaria anomala sp. nov., Chesterfield Islands, Pacific Ocean.

Introduction Pacific Ocean between 10°-30°S and 160°-175°E. During these expeditions large and interesting collections of Since 1978 the «lnstitut Français de recherche scientifique Leptolida (Cnidaria, Hydrozoa) have been obtained that so pour le développement en coopération (ORSTOM), Centre de Nouméa, NouveIle Calédonie» has sampled the marine far have been partly studied. The results of a detailed study fauna of the New Caledonian region in the south-western of two leptolid families, Thyroscyphidae and Sertulariidae, have so far been published (Vervoort, 1993). The present paper is based on the study of a part of the large collection Reçu le 19 février 2004 ; accepté après révision le 27 septembre 2004. of the superfamily Plumularioidea and contains the descrip- Received 19 February 2004; accepted in revised form 27 September 2004. tion of a new species of the family Halopterididae. 314 CLADOPLUMARIA ANOMALA GEN. NOV., SP. NOV.

Material Table 1. Measurements of Cladoplumaria anomala in µm. Tableau 1. Biométrie de Cladoplumaria anomala en µm. Material examined Chesterfield Islands. CHALCAL 1: Stn CP 5, 19°29.10’S, Chalcal 1 CP 5 158°37.63’E, 290 m depth, 16.07.1984: fragments of sever- al large colonies of which many with gonosome (‘phylacto- Height of (fragmented) colony (in mm) 8-196 Diameter Stem 55-100 carp’ and male and female gonothecae). Holotype, a Hydrotheca, length abcauline wall 190-210 fragmented colony to be deposited in the Muséum National length adcauline wall 210-220 d’Histoire Naturelle, Paris under no. Hy 1293. diameter at rim 70-90 Mesial nematothecae, length 75-100 Results Diameter at rim 25-30 Lateral nematothecae, length 65-80 Diameter at rim 20-30 Genus Cladoplumaria gen. nov. Female Gonothecae, length 440-500 Diagnosis Maximum diameter 290-320 Robust colonies composed of a rigid, ramified polysiphonic Phylactocarp, length 1830-2140 Female Gonothecal nematothecae, length 350-500 axis (hydrocaulus); branches polysiphonic, always origina- Diameter at rim 40-60 ting from a hydrocladium. Hydrocladia alternately arranged Male gonothecae, length 290-380 along axis and branches, originating from primary tube and Diameter at rim 150-190 alternately directed left and right, placed on distinct apo- physis provided with well-developed mamelon. Hydrothecae on basal parts of the branches and on the hydrocladia, tubu- lar, with smooth rim. All nematothecae conical, bithalamic axis arises. All material was detached, but the flattened and movable. Gonothecae protected by a phylactocarp; at mass of hydrorhizal fibers suggests that it developed on least the female gonothecae bearing nematothecae. firm substrate. Colony irregularly branched, with a tenden- Type species: Cladoplumaria anomala sp. nov. cy towards alternate disposition of branches. Primary axial tube of main stem undivided, with a longi- Etymology tudinal series of hydrocladial apophyses, distal parts may The name of the genus, Cladoplumaria, bas been derived show irregular division into internodes by means of straight from Cladocarpus Allman, 1888 and Pseudoplumaria septa. Apophyses alternately directed left and right; two or Ramil & Vervoort, 1992, two genera of Plumularioidea three nematothecae between two successive apophyses. with which the new genus shows superficial resemblance. Each apophysis with one well developed mamelon on The gender is female. upper surface and with two axillary nematothecae, one on back of apophyses near its insertion, second on axis above Cladoplumaria anomala sp. nov. (Figs 1-4, Table 1) apophysis. Secondary tubes running parallel with primary tube, develop from hydrorhizal mass leading towards Diagnosis strong polysiphony of stem and branches; stem basally 1.5 Cladoplumaria anomala sp. nov. is characterized by polysi- phonic, ramified colonies, apophyses with a well developed mm diameter. Secondary tubes with many nematothecae; mamelon, hydrothecae placed on hydrocladia and basal part all nematothecae bithalamic and movable. of branches (Fig. 3A, C). The gonothecae are protected by a Branches originating from hydrocladia, becoming poly- dichotomously branched phylactocarp (Fig. 2D, 4D); the siphonic by apposition of secondary tubes. Basal part of female gonothecae have long nematothecae (Fig. 2 A-C), primary tube of branch with a linear series of hydrothecae these are lacking in the male gonothecae (Fig. 4A-C). flanked by nematothecae in same position as those on the hydrocladia. Series of hydrothecae followed by apophyses Etymology supporting hydrocladia, alternately directed left and right; The species name ‘anomala’ is derived from the greek the, originally axillary, hydrothecae gradually change into adjective ‘anomalus’, abnormal, referring to the condition mamelons. The ultimate structure of the branches is as that of the gonosome. of the main stem. Hydrocladia inserting on apophyses, alternately directed Description left and right, undivided or with an occasional oblique node. Colony large, about 200 mm high, composed of a dense Hydrothecae tubular, narrow and deep; adcauline wall fully mass of hydrorhizal fibers from which the polysiphonic adnate, abcauline wall moderately concave, hydrothecal rim J. A. ANSÍN AGÍS, F. RAMIL, W. VERVOORT 315

Figure 1. Cladoplumaria anomala sp. nov. A. Hydrocladial internodes, lateral view, x 102.5. B. Hydrocladial internodes, frontal view, x 102.5. C. Regenerated hydrocladia after damage with two extra nematothecae, x 77.5. D. Hydrocladium with phylactocarp, x 38.3. Figure 1. Cladoplumaria anomala sp. nov. A. Articles d’un hydroclade, vue latérale, x 102,5. B. Articles d’un hydroclade, vue frontale, x 102,5. C. Régénération d’un hydroclade endommagé avec deux nématothèques supplementaires, x 77,5. D. Hydroclade avec un phylactocarpe, x 38,3. 316 CLADOPLUMARIA ANOMALA GEN. NOV., SP. NOV.

Figure 2. Cladoplumaria anomala sp. nov. A-C. Female gonothecae with nematothecae, x 61. D. Hydrocladium with phylactocarp and two female gonothecae, x 51.6. E. Phylactocarp with two female gonothecae and a hydrotheca in the last segment of a branch, x 51.6. Figure 2. Cladoplumaria anomala sp. nov. A-C. Gonothèque femelle avec nématothèques, x 61. D. Hydroclade avec phylactocarpe et deux gonothèques femelles, x 51,6. E. Phylactocarpe avec une hydrothèque sur le dernier segment d’une branche, x 51,6. J. A. ANSÍN AGÍS, F. RAMIL, W. VERVOORT 317

Figure 3. Cladoplumaria anomala sp. nov. A. Basal portion of a branch, x 26,6. B. Transition from hydrotheca to mamelon, x 51. C. Basal portion of a branch to show the transition from hydrotheca to mamelon as well as the arrangement of hydrocladia, x 35. Figure 3. Cladoplumaria anomala sp. nov. A. Partie basale d’une branche, x 26,6. B. Transformation d’une hydrothèque en un mamelon, x 51. C. Partie basale d’une branche avec la disposition typique des hydroclades et la transformation d’une hydrothèque en un mamelon, x 35. 318 CLADOPLUMARIA ANOMALA GEN. NOV., SP. NOV.

Figure 4. Cladoplumaria anomala sp. nov. A-C. Male gonothecae, x 170. D. Phylactocarp with a male gonotheca and a hydrotheca in the last segment of a branch, x 105. Figure 4. Cladoplumaria anomala sp nov. A-C. Gonothèque mâle, x 170. D. Phylactocarpe avec une gonothèque mâle et une hydrothèque sur le dernier segment d’une branche, x 105. J. A. ANSÍN AGÍS, F. RAMIL, W. VERVOORT 319 smooth. Pair of flanking nematothecae inserting at slightly also excludes its inclusion in the family Kirchenpaueriidae different height near hydrothecal rim, projecting above Stechow, 1921. The two remaining families of hydrotheca. Two median nematothecae between two Plumulariodea, the Plumulariidae McCrady, 1859, and consecutive hydrothecae, when internode is present one of Halopterididae Millard, 1962 have recently been discussed these becomes infracalycine, the second inserts close to basal by Calder (1997) and the Halopterididae were reviewed by node. Nematothecae as those on stem and branches. Schuchert (1997), but the limits between Plumulariidae and Gonosome shaped as ramified phylactocarp, inserting at Halopterididae are not clearly determined. Schuchert base of first hydrocladial hydrotheca and composed of (1997) considers that the most important diagnostic charac- forked basal part supporting two prongs that give rise to ter of Halopterididae is the presence of hydrothecae on the secondary and tertiary branches. In angle between branches hydrocaulus, and that the absence of hydrothecae on the a mamelon with distal, circular aperture; in addition each caulus (or their reduction to a “mamelon”) must be seen as segment with two or three nematothecae; distal segments a synapomorphy that unites in a clade the families with increased number of nematothecae, pointed. Plumulariidae, Kirchenpaueriidae and Aglaopheniidae. Gonothecae globular, shortly stalked, top flattened with cir- Contrary to Schuchert’s opinion, Calder (1997) considers cular lid. Gonothecae inserting under a mamelon in wider the presence of the large mamelons on the cauline apophy- part of segments. Each female gonotheca with three to five ses (regarded as atrophied hydrothecae) and the presence of long, tubular nematothecae inserting slightly under its mid- nematothecae on the gonothecae as typical characters of dle; male gonothecae without nematothecae. Halopterididae. We have here largely followed Calder Perisarc moderately developed, fairly strong on stems, (1997) in his views concerning the allocation of genera to branches and hydrocladia, thinning out along abcauline both families, pending a phylogenetic revision of the wall of hydrotheca; gonothecae with thin walls. Polyps Halopterididae referred to by Schuchert (1997). badly preserved, nematocysts could not be studied. To the Plumulariidae are here referred the following genera: Nemertesia Lamouroux, 1812 (= Antennularia Variability Lamarck, 1816; Antennopsis Allman, 1877; Nemertella Occasionally three nematothecae are present between two Stechow, 1923), Plumularia Lamarck, 1816, Hippurella consecutive hydrothecae. New branches and hydrocladia, Allman, 1877 (= Antomma Stechow, 1919), Callicarpa developing after damage and subsequent regeneration, usu- Fewkes, 1881, Sciurella Allman, 1883, Monotheca Nutting, ally have internodes separated by nodes. Sometimes the 1900, Sibogella Billard, 1911 (= Stechowia, Nutting, 1922), phylactocarp inserts under the second or third hydrotheca Dentitheca Stechow, 1920 (= Sphaerocystis, Fraser, 1943), of a hydrocladium. In two phylactocarps the last segment of MonothecelIa Stechow, 1923, and Cladacanthella Calder, a branch is provided with a hydrotheca flanked by two lat- 1997 (= Acanthella Allman, 1883). The taxonomic position eral nematothecae (Figs. 2E, 4D). of Plumella Stechow, 1920 is left undecided for the moment. So far gonothecae with nematothecae have only Discussion been observed in Plumularia wasini Jarvis, 1922. Phylactocarps or phylactocarp-like structures protecting the Cladoplumaria gen. nov. is characterized by large, polysi- gonothecae have been observed in certain genera phonic, ramified colonies, apophyses with a well developed (Hippurella Allman, 1877 and Callicarpa Fewkes, 1881). mamelon, hydrothecae placed on hydrocladia and basal In the Halopterididae, we here place the following gene- part of branches and accompanied by four bithalamic and ra: Antennella Allman, 1877, Allman, 1877 (= movable nematothecae (two laterals and two median infra- Heteroplon Allman, 1883; Acladia Marktanner- calycine), while the gonothecae are inserted on a dichoto- Turneretscher, 1890; Heterotheca Stechow, 1921), mously branched phylactocarp; the female having long Monostaechas Allman, 1877, Schizotricha Allman, 1883, nematothecae. These characters are also found in the Polyplumaria G.O. Sars, 1879, Calvinia Nutting, 1900, plumularioid families Halopterididae Millard, 1962, and Nuditheca Nutting, 1900, Allman, 1885 (= Plumulariidae McCrady, 1859. The presence of phylacto- Paragattya Warren, 1908), Antennellopsis Jäderholm, carps is characteristic of many genera of Aglaopheniidae 1896, Anarthrocladia Naumov, 1955, Astrolabia Naumov, Marktanner-Turneretscher, 1890, but the morphology of the 1955, Pentatheca Naumov, 1955, Corhyza Millard, 1962, bithalamic, movable nematothecae excludes the genus Pseudoplumaria Ramil & Vervoort, 1992, and Cladoplumaria from the Aglaopheniidae with monothala- Diplopteroides Peña Cantero & Vervoort, 1999 (= mic, immovable nematothecae, those in the cormidia being Diplopteron sensu Nutting, 1900). Nematothecae on the at least partly coalescent with the hydrotheca. The gonothe- gonothecae occur in many genera and insertion of gonothe- cae, in this family, have no nematothecae (Calder, 1997). cae on a phylactocarp is found in one (Diplopteroides). The The morphology of the nematophores and nematothecae arrangement of the genera in Plumulariidae or 320 CLADOPLUMARIA ANOMALA GEN. NOV., SP. NOV.

Table 2. Genera of the family Halopterididae and principal differentiating characters. Tableau 2. Genres de la famille Halopterididae et principaux caractères distinctifs. Stem Hydrocladia Hydrotheca Nematothecae Gonothecae Species Remarks

Antennella Flexuous, monosi- Only exceptionally One to each Bithalamic, movable, Unprotected, 16 Colonies occasio- Allman, 1877 phonic, heterome- present. hydrothecate not fused to hydrothe- ovoid to saccate, nally like those of rously segmented, Hydrotheca with internode (or ca or internode, with with flattened top, Halopteris or internodes occasio- one or two pairs of more on fused exception of unpaired inserting on the- Monostaechas. nally fused, frequent- lateral (‘flanking’) internodes), cam- nematotheca under cate internode ly forming tufts with nematothecae; panulate to cylin- hydrotheca that may under hydrotheca; adnate stems, unbran- varied number of drical; rim become fixed; reduced nematothecae ched (can be conside- nematothecae bet- smooth. nematotheca behind may be present. red hydrocladia rising ween hydrothecae. hydrotheca may be directly from stolon). present.

Halopteris Flexuous or rigid, Heteromerously One to each Bithalamic, movable, Unprotected, 32 Probably a poly- Allman, 1877 usually monosiphonic segmented, rarely hydrothecate not fused to hydrothe- generally ovoid to phyletic group of and segmented, occa- branched. internode of stem ca or internode, with globular, inserting species (cf. sionally branched, Secondary tubes, or hydrocladium, exception of unpaired on thecate inter- Schuchert, 1997). internodes with when present, may campanulate to nematotheca under node under Species of hydrothecae and develop hydrothe- cylindrical, rim hydrotheca that may hydrotheca; Antennella occa- alternate or opposite cae and hydrocla- smooth or with become fixed; reduced nematothecae sionally resemble apophyses supporting dia. one abcauline nematotheca behind may be present. monosiphonic hydrocladia, forming cusp. hydrotheca may be Halopteris spe- pinnate colonies. present. cies.

Monostaechas Stem, if present, poly- Heteromerously One to each Bithalamic, movable, Unprotected, 5 Species of Allman, 1877 siphonic and compo- segmented, bran- hydrothecate not fused to hydrothe- ovoid or globular, Antennella occa- sed of intercommuni- ching characteris- internode of ca or internode, with with flattened top, sionally resem- cating tubes; hydro- tic: successive hydrocladium, exception of unpaired inserting on the- bling cladia may also rise branches arising campanulate to nematotheca under cate internode Monostaechas in directly from stolon. from posterior sur- cup-shaped; no hydrotheca that may under hydrotheca, mode of bran- face of previous hydrothecae on become fixed; reduced with nematothe- ching hydrocladium, for- stem (if present). nematotheca behind cae. ming helicoid or Rim smooth. hydrotheca may be scorpioid sympo- present. dia. Schizotricha Erect, mono- or poly- Heteromerously One hydrotheca Bithalamic, movable, Unprotected, 15 Allman, 1883 siphonic, branched or segmented, usually on each stem not fused to hydrothe- pear-shaped with unbranched; primary sympodially bran- internode; one on ca or internode; one flattened top to axial tube homome- ched, branches each hydrothecate pair of ‘flanking’ elongated sack- rously segmented, inserting below internode of nematothecae placed shaped, inserting internodes each with hydrotheca, on hydrocladium, near hydrothecal rim on thecate inter- one hydrotheca and anterior or lateral campanulate to node under one apophysis, alter- surface. cup-shaped; rim hydrotheca, with nately directed left or circular. Smooth. nematothecae. right; colony pinnate. Antennellopsis Stem either ‘hydro- Unbranched, either Hydrotheca deep One pair of lateral Unprotected, 1 or 2 Either one species Jäderholm, 1896 cladia’ rising directly directly inserted on cup-shaped; nematothecae and two elongated ovoid, with two differing from stolon or a poly- stolon or on tubes adcauline wall unpaired between suc- with flattened top, modes of growth siphonic stem com- from polysiphonic completely fused cessive hydrothecae, inserted directly (age-related?) or posed of stolon-like stem, basal part to internode, on bithalamic, immova- below hydrotheca, two separate spe- tubes. ahydrothecate, front of hydrocla- ble. with nematothe- cies. remainder hydro- dium. cae thecate and usually unsegmented. Corhiza Millard, Erect, branched or Unbranched, basal Hydrotheca cam- Bithalamic, one or two Unprotected, 9 Probably poly- 1962 unbranched, compo- part ahydrothecate; panulate to cylin- pairs of lateral (‘flan- ovoid with flatte- phyletic; some sed of intercommuni- thecate part hetero- drical, adcauline king’) nematothecae ned top, inserted species quite near cating tubes forming merously segmen- wall with free and a varied number directly below to Antennella. hydrothecate hydro- ted. part. Rim smooth, on rest of hydrothecate hydrotheca, with cauli with pinnately circular or sinua- and ahydrothecate nematothecae arranged hydrocladia, ted. internodes. or directly hydrocla- dia. J. A. ANSÍN AGÍS, F. RAMIL, W. VERVOORT 321

Gattya Allman, Erect, branched or Unbranched, Hydrotheca campa- Bithalamic and mova- Unprotected, 8 1886 unbranched, mono- homo- or hetero- nulate to cup-sha- ble; one pair of lateral pear-shaped, or polysiphonic; merously segmen- ped; part of adcauli- (‘flanking’) nematothe- inserted directly when monosiphonic ted, arrangement ne wall free; rim cae, one (unpaired) below hydrotheca, with hydrothecae and sub-opposite to with several cusps. above, one under with nematothe- hydrocladia, when alternate. Abcauline hydrothe- hydrotheca; one nema- cae. polysiphonic hydro- cal wall in some totheca on ahydrothe- cladia, hydrothecae species with cate internode (if pre- and nematothecae inwardly projecting sent) only on main tube. ledge.

Calvinia Nutting, Erect, polysiphonic, Homomerously Hydrotheca cup- Bithalamic and mova- Unprotected, glo- 1 Characters of 1900 pinnately branched, segmented; seg- shaped, with ble, a pair of lateral bular, inserted at genus based on ultimate parts mono- mented, nemato- smooth, circular (‘flanking’) nematothe- base of nemato- single species siphonic, with hydro- theca-bearing pro- rim. cae near hydrothecal phorous process; known thecae. cess at base of rim and others distri- with nematothe- each hydrotheca. buted over internode cae. and nematophorous process.

Nuditheca Erect, polysiphonic, Pinnately arranged Hydrotheca deep Bithalamic, unmova- Unprotected, 3 Nutting, 1900 branched or unbran- along stem and cup-shaped, adcau- ble; one pair of lateral elongated sack- ched, ultimate parts branches, typically line wall adnate to (‘flanking’) nematothe- shaped, inserting monosiphonic, with branched, homo- internode; rim cae and one unpaired on internode hydrothecae. merously segmen- smooth, circular or nematotheca under under hydrotheca; ted; occasionally sinuated. hydrotheca. with nematothe- ahydrothecate cae internode present.

Astrolabia Erect, polysiphonic Branched, pinnate- Hydrotheca elonga- Bithalamic, unmova- Unprotected, 2 Naumov, 1955 basally, monosipho- ly arranged; bran- ted cup-shaped, ble, two pairs of lateral elongated sack- nic in upper parts ching at top of adcauline wall (‘flanking’) nematothe- shaped, inserted and there segmented, first hydrocladial adnate to internode; ca: one near hydrothe- on hydrocladia each segment with internode; seg- rim smooth though cal rim, one at middle under hydrotheca, big apophysis and mentation homo- sinuated. Axillary of adcauline wall of apparently modified hydrotheca merous though hydrotheca on stem hydrotheca; single without nemato- in its axil; apophyses first internode dif- internode modified, (unpaired) nematothe- thecae and with alternately directed fering in shape (by more or less tubu- ca under hydrotheca. lateral, slit-like left and right. branching!) and lar; slightly curved. mouth. arrangement of nematothecae.

Anarthrocladia Erect, branched, Unbranched, long, Hydrotheca campa- Lateral (‘flanking’) Unprotected, big, 1 Characters of Naumov, 1955 polysiphonic, basally in two closely nulate with slightly nematothecae modi- elongated ovoid, genus based on thick, gradually nar- packed opposite everted, smooth fied, scale-shaped, with single basal single species rowing towards rows, division into rim, adcauline wall directed anterior. nematotheca and known. monosiphonic apical internodes indis- fully adnate to inter- Unpaired nematotheca big, laterally parts bearing axial tinct to completely node. at hydrothecal base, directed aperture. hydrothecae. absent. bithalamic.

Pentatheca Erect, occasionally Unbranched, Hydrotheca campa- Bithalamic though Unprotected, big, 1 Characters of Naumov, 1955 branched, polysipho- arranged along nulate with slightly immovable; one pair elongated ovoid, genus based on nic, gradually narro- both sides of stem, everted, smooth, on each side of hydro- with basal pair of single species wing apically homomerously circular rim; adcau- thecal rim, a second nematothecae; known. towards monosipho- segmented over line wall fully adna- pair basally on interno- aperture big, late- nic, hydrothecate whole length, each te to internode. de under base of rally directed. part. internode with one hydrotheca. Unpaired hydrotheca and nematotheca absent. two pairs of nema- tothecae. 322 CLADOPLUMARIA ANOMALA GEN. NOV., SP. NOV.

Pseudoplumaria Erect, branched and Unbranched, Cup-shaped to All bithalamic, Unprotected, pear- 2 Ramil & Vervoort, polysiphonic, gra- homomerously seg- cylindrical, about movable or immo- shaped to elongated 1992 dually thinning and mented (except for half of adcauline vable. sack-shaped, inser- becoming monosi- short ahydrothecate wall free from ting on apophyses, phonic; branches ori- basal internode), internode; rim cir- with 4-13 nematothe- ginating from secon- with one hydrothe- cular, smooth. cae, occasionally dary tubes, alterna- ca having one or considerable elonga- ting or in opposite two pairs of lateral ted. pairs. Internodes of (‘flanking’) nema- monosiphonic parts tothecae and a long, with nematothe- varied number of cae and apophyses movable or immo- alternately directed vable bithalamic left and right; with nematothecae. big, reduced, axillary hydrotheca (mame- lon) or occasionally normal hydrotheca. Polyplumaria G.O. Erect, polysiphonic, Typically with Cup-shaped to All bithalamic and Pear-shaped, inser- 6 Sars, 1874 pinnately branched, secondary, occasio- cylindrical, large movable. ting on apophysis, branches formed from nally with tertiary portion of adcau- with six to nine basal secondary tube. hydrocladia, homo- line wall free nematothecae. Monosiphonic, ulti- merously segmen- from internode; mate parts indistinctly ted (except for rim circular, segmented, with apo- short ahydrothecate smooth. physes alternately basal internode), directed left and right, with one hydrothe- having a modified ca, a pair of lateral axillary hydrotheca in nematothecae and the shape of a big three unpaired mamelon. ones, one below, two above hydro- theca of which one occasionally on separate internode.

Diplopteroides Erect, unbranched, Unbranched, undi- Elongated cup- Bithalamic, mova- ‘Protected’ by phy- 3 Characters of Peña Cantero & polysiphonic; mono- vided or irregularly shaped to tubular, ble, not fused to lactocarp-like struc- gonosome only Vervoort, 1999 siphonic ultimate divided into inter- only fraction of hydrotheca or ture composed of for- based on type of parts hydrothecate, nodes; hydrothecae adcauline wall internode; one pair ked nematophorous genus; two remai- with apophyses with two pairs of free. Rim sinua- of ‘flanking’ nema- and occasionally ning species with having a hydrotheca lateral (‘flanking’) ted. tothecae placed hydrothecate bran- unknown gonoso- at their base. nematothecae and near hydrothecal ches inserting at base me. additional nemato- rim; a second pair of first hydrotheca of thecae between halfway hydrothe- hydrocladium; gono- hydrothecae. cal border thecae globular with flattened top inserted on arms of ‘phylacto- carp’ and having a few nematothecae. Cladoplumaria Erect, polysiphonic, Unbranched, irre- Tubular, adcauli- Bithalamic and ‘Protected’ by phy- 1 Characters of gen. nov. with alternate bran- gularly divided into ne wall complete- movable lactocarp-like, dicho- genus based on ches derived from internodes with ly adnate to inter- tomously branched single species hydrocladia becoming several hydrothe- node; rim smooth, nematophorous struc- known. covered by secondary cae; each hydrothe- slightly sinuated ture inserting at base tubes; ultimate parts ca with pair of late- of first hydrocladial monosiphonic, irregu- ral (‘flanking’) hydrotheca. larly segmented, with nematothecae and Gonothecae globular nematothecae and two median nema- with flattened top, apophyses alternately tothecae, one inserted on ‘arms’ of directed left and right, above, one below ‘phylactocarp’, each having a modified hydrotheca. with 3-5 elongated axillary hydrotheca or nematothecae mamelon. J. A. ANSÍN AGÍS, F. RAMIL, W. VERVOORT 323

Halopterididae presented here differs from views expressed Challenger, Zoology, 7: 1-55. by Ansín Agís et al. (2001) in a recent paper and is the Allman G.J. 1885. Description of Australian, Cape, and other result of weighing the evidence presented by Calder (1997) Hydroida, mostly new, from the collection of Miss H. Gatty. and the comparison with the morphology of the present Joumal of the Linnean Society Zoology, 19: 132-161. Ansin Agis J., Ramil F. & Vervoort W. 2001. new species. Atlantic Leptolida (Hydrozoa, Cnidaria) of the families Aglaopheniidae, Cladoplumaria anomala sp. nov., in colony structure, Halopterididae, Kirchenpaueriidae and Plumulariidae collec- arrangement of hydrothecae and accompanying nematothe- ted during the CANCAP and Mauritania-Il expeditions of the cae shows affinities with Polyplumaria and National Museum of Natural History. Zoologische Pseudoplumaria, from which genera it differs by the pre- Verhandelingen Leiden, 333: 1-268. sence of phylactocarps and the pattern of ramification, as in Bedot M. 1921. Notes systématiques sur les plumularides. 2ème Polyplumaria and Pseudoplumaria the branches arise from partie. Revue suisse de Zoologie, 29: 1-40. secondary tubules and not from a hydrocladium. The pres- Billard A. 1911. Note sur un nouveau genre et une nouvelle ence of phylactocarps in the family Halopterididae only espèce d’hydroïde: Sibogella erecta. Archives de Zoologie occurs in the genus Diplopteroides Peña Cantero & expérimentale et générale, (5)6, notes et revue, 108-109. Vervoort, 1999 (= Diplopteron sensu Nutting 1900), but in Bouillon J. 1985. Essai de c1assification des Hydropolypes- Hydroméduses (Hydrozoa-Cnidaria). Indo-Malayan Zoology, this genus the colonies are unbranched, there is a cauline 1: 29-243. hydrotheca at base of each hydrocladium and the gonothe- Calder D.R. 1997. Shallow-water hydroids of Bermuda: cae are without nematothecae. The various genera of (Superfamily Plumularioidea). Life Sciences Contributions of Halopterididae and their principal characters are listed in the Royal Ontario Museum, 161: 1-85. Table 2. Fewkes J.W. 1881. Report on the Acalephae. In: Reports on the The phylactocarp of Cladoplumaria anomala sp. nov. results of dredging, under the supervision of Alexander could also be conceived as a branched hydrocladium as Agassiz, in the Caribbean Sea, in 1878, 1879, and along the occurs in species of Schizotricha, the forked basal portion Atlantic coast of the United States, during the summer of 1880, of the phylactocarp representing the first internode of a pri- by the U.S. Coast Survey Steamer Blake. Bulletin of the mary hydrocladium with its mamelon, giving rise to Museum of comparative Zoology at Harvard College, 8: 127- 140. hydroc1adia of lower order. This suggestion seems to be Fraser C. McLean 1943. Distribution records of same hydroids strengthened by the occasional occurrence of a hydrotheca in the collection of the Museum of Comparative Zoology at flanked by two lateral nematothecae on one of the branches. Harvard College, with description of new genera and new species. Proceedings of the New England zoological Club, 22: Aknowledgements 75-98. Jäderholm E. 1896. Ueber aussereuropäische Hydroiden des zoologischen Museums der Universitat Upsala. Bihang till The authors wish to express their gratitude to Dr. A. Kungliga svenska Vetenskaps-Akademiens Handlingar, 21, Crosnier (Muséum National d’Histoire Naturelle, Paris, Afd. 4: 1-20. France) for providing the material for this study. We extend Jarvis F.E. 1922. The hydroids from the Chagos, Seychelles and our gratitude to Dr. D.R. Calder and Dr A. Peña Cantero for other islands and from the coasts of British East Africa and their suggestions about the position of the new genus in Zanzibar. 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