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, 1868INTRODUCTION
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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