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Posicion taxonomica de Lovenella gracilis Clarke, 1882 (Lovenellidae, Hydrozoa): nuevas evidencias de microanatomia justifican su permanencia en el genero Lovenella Hincks, 1868.

Taxonomic position of Lovenella gracilis Clarke, 1882 (Lovenellidae, Hydrozoa): new evidences of microanatomy justify its maintenance in the genus Lovenella Hincks, 1868

INTRODUCTION

Leptothecate hydrozoans of the family Lovenellidae Russell, 1953 have a troubled taxonomical history (Calder, 1991; Bouillon et al., 2004). They have a metagenetic life cycle and, like in many other hydroid taxa, the parallel and independent use of the morphological characters of polyps and medusae eventually generated a dual classification, with different understandings and diagnoses for the, presumably, same genera.

The genus Lovenella was proposed by Hincks (1868), based on the polyp stage, and assigned to the family Campanulariidae. Russell (1953), based on the medusa stage, proposed the family Lovenellidae including the genera Eucheilota McCrady, 1859 and Lovenella Hincks, 1868. The family Lovenellidae, as described by Russell (1953), includes medusae without marginal cirri, excretory pores or peduncle, with gonads on four simple radial canals, and with lateral cirri. Kramp (1959) proposed the genus Cirrholovenia as a third genus for the Lovenellidae, based on the presence of marginal cirri in the medusa, amending the original diagnosis of the family. Other disputable arrangements have also been proposed for the family, even comprising genera traditionally included in the family Haleciidae Hincks, 1868, such as Campalecium Torrey, 1902 and Hydranthea Hincks, 1868.

The genus Lovenella Hincks, 1868, type species Lovenella clausa (Loven, 1836), comprises 14 nominal species (Tables 1 and 2) distributed worldwide (Figs. 1 and 2). Only L. chiquitita Millard, 1957 and L. corrugata Thornely, 1908 were recorded for the South Atlantic hitherto. The main diagnostic characters of Lovenella are the medusae with indefinite number of statocysts and polyp with hydrotheca well demarcated by a basal line, separating it from the operculum (Fraser, 1944; Kramp, 1959; Bouillon et al., 2004).

Oddly, the polyp of Lovenella gracilis Clarke, 1882 is defined by the operculum being a continuation of the hydrothecal wall, therefore lacking a basal line separating operculum and hydrotheca (Calder, 1971, 1975). Based on this unique character, Calder (1991) proposed the resurrection of the genus Dipleuron Brooks, 1882 in order to encompass L. gracilis. Bouillon & Boero (2000) and Bouillon et al. (2004) did not agree with this proposal, arguing that the medusa of L. gracilis presents the typical characters of the genus and that the diagnostic characters of the polyps of "lovenellid" are puzzling, since the opercular structure can be variable within the family, and even within the same genus. No other addition was made to the knowledge of the morphology of L. gracilis, and the taxonomical "status" of the species remains doubtful.

The aim of this study is to redescribe in detailed morphology the polyp of L. gracilis, based on the first material of the species sampled for the South Atlantic, and reveal new data corroborating its maintenance in the genus Lovenella.

MATERIALS AND METHODS

The material studied was collected in the intertidal zone of Tibau Beach (Tibau, State of Rio Grande do Norte, Brazil) and Bombas Beach (Bombinhas, State of Santa Catarina, Brazil). The colonies were fixed in 92.5% ethanol and 4% formaldehyde solution. We have studied the morphology, morphometry and cnidome of all specimens. Morphological details were studied in scanning electronic microscopy (SEM), following routine protocol (Migotto & Marques, 1999). The cnidome was studied with squashed preparations of the fixed material, in light microscopy. Studied material has been deposited in the Cnidarian Collection of the Museu de Zoologia of the Universidade de Sao Paulo (MZUSP), Sao Paulo, Brazil.

RESULTS

Lovenella gracilis Clarke, 1882 (Figs. 3a-3d; 4a-4f).

Lovenella gracilis Clarke, 1882, p. 139, pl. 9, fig. 25-39; Fraser, 1944, p. 174, pl. 31, fig. 147; Calder, 1971, p. 61, pl. 4, fig. h, pl. 8, fig. b-c; 1975, p. 298, fig. 3c.

Dipleuronparvum Brooks, 1882, p. 135, 139-140.

Lovenella clausa -Fraser, 1910, p. 364, fig. 26a-d; 1912, p. 45 [non Lovenella clausa (Loven, 1836)].

Dipleuron gracilis-Huve, 1952, p. 389, fig. 1a-b, 2a-b; Calder, 1991, p. 3.

Material examined. Santa Catarina, Bombinhas, Bombas Beach (27.131[degrees]S 48.514[degrees]W, 2 m, 3.xii.2006)--MZUSP4242, in formaldehyde 4%, without gonophores, on rock and Sargassum sp.; MZUSP4260, in ethanol 92.8%, without gonophores, on rock; MZUSP4263, in ethanol 92.8%, with gonophores, on rock; MZUSP4266, in formaldehyde 4%, with gonophores, on rock and Sargassum sp. Rio Grande do Norte, Tibau, Tibau Beach (4.835[degrees]S, 37.247[degrees]W, intertidal zone, on Donax striatus, in ethanol 92.8%)MZUSP5356, with gonophores, 5.vi.2004; MZUSP 5357, MZUSP5358, with gonophores, 15.ix.2004; MZUSP5359, with gonophores, 9.iii.2004.

Description. Colonies stolonal or erect, up to 19 mm (n = 10) in height, arising directly from creeping hydrorhiza 80-240 [micro]m (n = 10) in diameter. Hydro-caulus monosiphonic, with 0-6 annulations (n = 10) at the proximal region, branched or unbranched, divided into internodes by transverse septa at more or less regular intervals. Perisarc of main stem moderately thick, thinner at secondary branches and pedicels. Internode length 930-6640 [micro]m (n = 10), diameter 87.5-160 [micro]m (n = 10), with 1-7 septa (n = 10), supporting hydrothecal pedicel arising from distal apophysis. Apophyses alternate; branches or additional pedicels, when present, arising laterally to the apophysis. Pedicels either annulated throughout or with 2-11 (n = 10) distal annulations, length 110-820 [micro]m (n = 10), diameter 75-120 [micro]m (n = 10). Hydrotheca campanulate, 350-740 [micro]m (n = 10) deep from rim to base, 215-340 [micro]m (n = 10) wide at margin, 100-180 [micro]m (n = 10) wide at diaphragm; diaphragm thin, transversal; operculum with 8-11 triangular to pentagonal valves (n = 10), apparently as folded continuation of hydrothecal wall, but with discrete line demarcating operculum from hydrotheca (only in SEM). Gonothecae inverted cone-shaped, length 620-1180 [micro]m (n = 10), diameter at margin 150-270 [micro]m (n = 10), diameter at base 100-200 [micro]m (n = 10); walls smooth, distal region of gonothecae deepened, with a central aperture. Gonothecal pedicels short, length 60-300 [micro]m (n = 10), diameter 60-120 [micro]m (n = 10), with 2-8 annulations (n = 10) throughout, arising near base of hydrothecal pedicels or directly from hydrorhiza; several medusa buds in each gonotheca, but some gonothecae empty. Nematocysts of one type: small microbasic mastigophores, dimensions 6-7 [micro]m X 1.5-2 [micro]m (n = 10, undischarged capsules).

Distributional range. North Atlantic (Clarke, 1882; Brooks, 1882; Fraser, 1910, 1912, 1944; Calder, 1971, 1975; Manning & Lindquist, 2003; Bouillon et al., 2004; Dougherty & Russell, 2005), Gulf of Mexico (Calder & Cairns, 2009), Caribbean Sea (Bandel & Wedler, 1987), Mediterranean Sea (Huve, 1952; Picard, 1958; Bouillon et al., 2004).

DISCUSSION

Clarke (1882: 139) described the polyp and medusa of Lovenella gracilis for Chesapeake Bay, uncertain of its "relationships and systematic position" when compared to L. clausa (Loven, 1836). Indeed, Fraser (1910, 1912) mistakenly assigned North Carolina and Massachusetts specimens of L. gracilis to L. clausa; but he corrected himself after examining further material, noting that both species are distinct and that "the European species L. clausa has not been observed in the Western Atlantic" (Fraser, 1944: 174).

Concomitantly to Clarke's description of L. gracilis, Brooks (1882) described the new genus Dipleuron, and its type-species D. parvum, based on a medusa found at North Carolina coast. Huve (1952), based on Mediterranean material, considered L. gracilis and D. parvum similar, adopting the name Dipleuron gracilis because Lovenella would not be a valid genus since the type species L. clausa was linked to the medusa of Eucheilota hartlaubi by Russell (1936a). However, as explained by Calder (1971: 64) "Eucheilota and Lovenella are not congeneric, and the medusa E. hartlaubi has since been shown to be a Lovenella [ ... ]".

Life cycle studies of L. gracilis eventually revealed that its medusa stage is indistinguishable from D. parvum as described by Brooks (1882) (Calder, 1971). Then, Dipleuron was reaffirmed as junior synonym of Lovenella, with the actual name L. gracilis Clarke, 1882 having priority over Dipleuron parvum Brooks, 1882. However, Calder (1991) reconsidered this synonymy when referring to L. gracilis, arguing that Dipleuron and Lovenella would be distinct because of "differences in the morphology of their opercula" (Calder, 1991: 3).

Under light microscopy, the operculum of L. gracilis is a continuation of the hydrothecal wall, without demarcation (cf. Calder, 1971, 1975; Bouillon et al., 2004). The original definition of the genus Lovenella has no mention to a basal line demarcating the operculum (Hincks, 1868), therefore potentially accommodating L. gracilis. Amending diagnoses, however, have defined Lovenella by the presence of this line demarcating the operculum (Calder, 1991; Cornelius, 1995), a notable characteristic of most of the species of the genus (Millard, 1957; Cornelius, 1995). Based on this pattern, the absence of the demarcation in L. gracilis justified its transference to Dipleuron (Calder, 1991).

A refinement of the morphological study was necessary. We have found specimens representing the first record of L. gracilis for the South Atlantic and Brazilian coast [cf. Migotto et al., 2002; even though Stechow (1914) recorded Gonothyraea (?)nodosa, a disputable and inconclusive similar hydroid for Rio de Janeiro coast, to which we prefer not to make inferences about its taxonomic status]. Scanning electron microscopy of this Brazilian L. gracilis revealed the presence of a tenuous line separating the operculum from the hydrotheca (Fig. 4), making it clear it is a Lovenella species. Therefore, considering the troubled taxonomy of the family Lovenellidae and the new evidence presented herein, we propose the maintenance of the genus Dipleuron Brooks, 1882 as a junior synonym of Lovenella Hincks, 1868.

DOI: 10.3856/vol41-issue2-fulltext-7

ACKNOWLEDGEMENTS

We would like to thank Dr. Helena Matthews-Cascon (Universidade Federal do Ceara) for providing material from Rio Grande do Norte, Dr. Peter Schuchert (Museum d'Histoire Naturelle) and Dr. Yayoi Hirano (Chiba University) for the help with literature, and Enio Mattos (Universidade de Sao Paulo) for technical support. This study was supported by CAPES Procad, Prodoc e Pro-Equipamentos 1887/ 2007, CNPq (Proc. 490348/2006-8, 304720/2009-7, 304720/2009-7, 562143/2010-6, 563106/2010-7) and FAPESP (Proc. 2004/09961-4, 2006/58226-0, 2010/ 06927-0). This is a contribution of NP-BioMar, USP.

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Received: 16 May 2011; Accepted: 22 October 2012

Thais Pires Miranda (1), Amanda Ferreira Cunha (1) & Antonio C. Marques (1)

(1) Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo Rua do Matao Trav. 14, 101, 05508-090, Sao Paulo, Brazil

Corresponding author: Antonio C. Marques (marques@ib.usp.br)

Table 1. Comparison of the diagnostic characters of species of
Lovenella with polyp stage recorded in literature.

Species          Shape       Growth     Septa per   Annulations
                   of                   intemode    on pedicel
                 colony

L. chiquitita   Stolonal   Sympodial       --       Throughout
Millard, 1957   or erect

L. clausa       Stolonal   Sympodial       --         2-5 on
(Loven, 1836)   or erect                             proximal
                                                    and distal
                                                      regions

L. corrugata     Erect     Sympodial       --         3-6 on
Thornely, 1908                                       proximal
                                                    and distal
                                                      regions

L. gracilis      Erect     Sympodial       3-4        1-2 on
Clarke, 1882                                         proximal
                                                    and distal
                                                      regions

L. granais       Erect     Sympodial       --           Not
Nutting, 1901                                        specified

L. nodosa        Erect     Sympodial       --         2-3 on
Fraser, 1938                                          distal
                                                      region

L. paniculata    Erect     Sympodial       --           On
(G.O. Sars,                                          proximal
1874)                                                 region

L. producia     Stolonal   Monopodial      --         3-8 on
(G.O. Sars,                                          proximal
1874)                                               and distal
                                                     regions,
                                                    sometimes,
                                                      in the
                                                      middle

L. rugosa       Stolonal   Sympodial       --       Throughout
Fraser, 1938    or erect

Species         Hydrothecae    Frequency of    Shape of    Number of
                               hydrothecae    operculum    operculum
                               regeneration     valves      valves

L. chiquitita      Deep-       Occasionally   Triangular     8-10
Millard, 1957   campanulate                      with
                                               rounded
                                                 base

L. clausa       Cylindrical,    2-3 times     Triangular     8-10
(Loven, 1836)     widening                       with
                  distally                     rounded
                                                 base

L. corrugata     Deep-camp      2-4 times     Triangular     8-12
Thornely, 1908    anuiate,                       with
                 corrugated                    rounded
                 proximally                      base

L. gracilis     Campanulate         --         Conical         8
Clarke, 1882

L. granais      Cylindrical,        --        Triangular     10-12
Nutting, 1901     widening                       with
                  distally                     rounded
                                                 base

L. nodosa       Cylindrical,        --        Triangular       8
Fraser, 1938      widening                       with
                  distally                     rounded
                                                 base

L. paniculata   Cylindrical,        --         Conical        --
(G.O. Sars,       widening
1874)             distally

L. producia     Cylindrical,        --         Conical     Up to 12
(G.O. Sars,       widening
1874)             distally

L. rugosa          Deep-            --        Triangular     9-10
Fraser, 1938    campanulate                      with
                                               rounded
                                                 base

Species          Diaphragm    Nematophores   Gonothecae   Arising of
                                                          gonothecae

L. chiquitita     Oblique        Absent       Smooth,        From
Millard, 1957                                elongated,   hydrorhiza
                                              widening
                                              distally

L. clausa       Transversal      Absent       Smooth,      Axillary
(Loven, 1836)                                elongated,      from
                                              widening     pedicel
                                              distally

L. corrugata      Oblique        Absent       Ringed,      Axillary
Thornely, 1908                                spindle        from
                                               shaped      pedicel

L. gracilis     Transversal      Absent       Smooth,      Axillary
Clarke, 1882                                  clavate        from
                                                           pedicel

L. granais          --           Absent       Smooth,      Axillary
Nutting, 1901                                elongated,      from
                                              widening     pedicel
                                              distally

L. nodosa           --           Absent       Smooth,      Axillary
Fraser, 1938                                  tubular,       from
                                              widening     pedicel
                                              distally

L. paniculata     Oblique        Absent          --           --
(G.O. Sars,
1874)

L. producia     Transversal     Present          --           --
(G.O. Sars,
1874)

L. rugosa         Oblique        Absent       Smooth,        From
Fraser, 1938                                 elongated,   hydrorhiza
                                              widening
                                              distally

Species         References

L. chiquitita   Millard (1957,
Millard, 1957   1975)

L. clausa       Garcia Corrales et
(Loven, 1836)   al. (1979);
                Cornelius (1995)

L. corrugata    Vervoort (1959);
Thornely, 1908  Millard (1980)

L. gracilis     Clarke (1882);
Clarke, 1882    Calder (1971, 1975)

L. granais      Nutting (1901);
Nutting, 1901   Fraser (1941)

L. nodosa       Fraser (1938)
Fraser, 1938

L. paniculata   Sars (1874)
(G.O. Sars,
1874)

L. producia     Sars (1874);
(G.O. Sars,     Cornelius (1995)
1874)

L. rugosa       Fraser (1938)
Fraser, 1938

Table 2. Comparison of diagnostic characters of species of
Lovenella with medusa stage recorded in literature.

Species             Umbrella        Velum       Manubrium     Number
                                                             of gonads

L. annae (von        Broader          --         Small,          4
Lendenfeld,         than high                   globular,
1884)                                            with 4
                                               interradial
                                                  spots

L. assimilis         Broader        Narrow        Short          4
(Browne, 1905)      than high

L. bermudensis     Wide, low,         --        Short and        4
(Fewkes, 1883)       without                      wide
                     raised
                      apex

L. chiquitita        Higher           --          Short         --
Millard, 1957       than wide

L. clausa            Hemis-       ca. 1/4 of    Short and        4
(Loven, 1836)       pherical       the bell       small
                                    radius

L. gracilis          Hemis-          Wide         Short          4
Clarke, 1882        pherical

L.                   Hemis-           --          Short          4
haichangen-sis      pherical
Xu & Huang,
1983

L. sinuosa        Flatter than        --            --           4
Lin, Xu, Huang    hemispherical
& Wang, 2009

Species            Position of     Morphology of    Number of
                      gonads           gonads       tentacles

L. annae (von      Near radial          Oval            8
Lendenfeld,           canals
1884)

L. assimilis      On distal end         Oval            4
(Browne, 1905)      of radial
                      canals

L. bermudensis      On stomach       Spherical         4-8
(Fewkes, 1883)

L. chiquitita           --               --             8
Millard, 1957

L. clausa         On distal end       Oval and        16-24
(Loven, 1836)       of radial      longitudinally
                      canals          divided

L. gracilis       On midway of 2     Spherical         21
Clarke, 1882      radial canals

L.                On 2/3 of the    Slender linear       8
haichangen-sis    distal portion
Xu & Huang,         of radial
1983                  canals

L. sinuosa         On proximal        Linear,           4
Lin, Xu, Huang      portion of     longitudinally
& Wang, 2009        manubrium         divided

Species           Number of    Number of   Number of   Number of
                  statocysts   marginal    marginal      cirri
                               vesicles      bulbs

L. annae (von         --           -           8       2 clusters
Lendenfeld,
1884)

L. assimilis          --           -          5-7      2 clusters
(Browne, 1905)

L. bermudensis        --           -           4           2
(Fewkes, 1883)

L. chiquitita         --           8           8           --
Millard, 1957

L. clausa             --         16-23                    1-3
(Loven, 1836)

L. gracilis           --          33           4           --
Clarke, 1882

L.                    --          16          24       1-2 pairs
haichangen-sis
Xu & Huang,
1983

L. sinuosa           7-8          --           4       8-10 pairs
Lin, Xu, Huang
& Wang, 2009

Species           Position of      References
                     cirri

L. annae (von       On each       Kramp (1961)
Lendenfeld,         side of
1884)              tentacles

L. assimilis        On each      Browne (1905);
(Browne, 1905)      side of      Hirano & Yamada
                   tentacles         (1985)

L. bermudensis      On each      Fewkes (1883);
(Fewkes, 1883)      side of       Kramp (1959)
                   tentacles

L. chiquitita         --         Millard (1975)
Millard, 1957

L. clausa           On each       Kramp (1959);
(Loven, 1836)       side of         Cornelius
                   tentacles         (1995)

L. gracilis           --         Clarke (1882);
Clarke, 1882                      Calder (1971)

L.                  On each     Xu & Huang (1983)
haichangen-sis      side of
Xu & Huang,        tentacles
1983                  and
                   marginal
                     bulbs

L. sinuosa          On each     Lin et al. (2009)
Lin, Xu, Huang      side of
& Wang, 2009       tentacles
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Title Annotation:articulo en ingles
Author:Pires Miranda, Thais; Ferreira Cunha, Amanda; Marques, Antonio C.
Publication:Latin American Journal of Aquatic Research
Date:Apr 1, 2013
Words:3958
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