Scyphozoa: Coronatae: Nausithoidae
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Contributions to Zoology, 66 (4) 235-246 (1997) SPB Academic Publishing bv, Amsterdam Nausithoe Coronatae: with aurea n. sp. (Scyphozoa: Nausithoidae), a species two pathways of reproduction after strobilation: sexual and asexual Fábio Lang da Silveira & André Carrara Morandini Departamento de Zoología, Instituto de Biociências, Universidade de Sao Paulo, Caixa Postal 11461, 05422-970, SP, Brazil Keywords: Scyphozoa, Coronatae, systematics, life cycle, Nausithoe, new species Abstract and, below, pedalia in equal numbers to the ten- tacles and rhopalia (Franc, 1993; Russell, 1970). Nausithoe of is aurea, a new species scyphozoan Coronatae, The Coronatae are represented mostly by deep- described from São Paulo State, Brazil. The solitary scyphisto- sea medusae (Tardent, 1978) and only few spe- mae, with some zooxanthellae, strobilated producing plan- cies live in shallow waters (Larson, 1990). There- uloids and medusae; this represents an intermediate life cycle fore, their life are still little known (Jarms, between that of metagenetic Nausithoidae and the submarine cycles 1991). Some 40 of coronate medusae are cave-dwelling, reduced medusa stage of Nausithoe planulo- species phora (Werner, 1971). The periderm tube ofthe scyphistomae recognized and the metagenetic life cycles of 9 has internal in The 16 cusps all whorls. medusae present yellow known The life species are (Jarms, pers. comm.). pigment spots in most of their lappets. The early embryonic cycles of 4 additional species without free medu- development is briefly described. Planuloid formation is sae are known (Jarms, 1990; 1991). Although hypothesized as explanation for polyp-stage philopatry. the of they have a metagenetic life cycle, polyps all species of uncertain medusa genera are col- Résumé lectively grouped Stephanoscyphistoma (sensu Jarms, 1990; 1991). Several authors, e.g., Werner On décrit Nausithoe nouvelle de aurea, espèce Scyphozoaire (1971a, b; 1983), Jarms (1990), and Meroz & Ilan (Coronatae) de l’Etat de São Paulo, Brésil. Les scyphistomae (1995), have stressed the importance of life cycle solitaires, contenant des zooxanthelles, produisent des planu- studies for the correct identification of loïdes des méduses ceci species. et par strobilation; représente un cycle The coronate recorded from the Brazilian vital intermédiaire entre celui des Nausithoidae métagénétiques only celui de Nausithoe is the medusa of et planulophora (Werner, 1971) avec son coast stage Nausithoe punctata stade méduse réduit habitant des grottes sous-marines. Le tube Kölliker, 1853, from the north of Bahia State péridermique des scyphistomae présente 16 pointes internes (Goy, 1979). Recently, Silveira & Morandini (in dans chaque spire. Les méduses présentent des taches pigmen- press) redescribed the scyphistoma of a coronate, taires jaunes dans la plupart de leurs lobules. On donne une (Komai, succincte description des phases initiales du développement Stephanoscyphistoma corniformis 1936) On émet c’est la formation de from the Brazilian embryonnaire. l’hypothèse que coast. planuloïdes qui pourrait expliquer la philopatrie du stade polype. Material and methods Introduction The solitary scyphistomae are sampled by SCUBA diving from calcareous debris, mainly the stony coral Mussismilia hispida Coronatae have a with a firm polyp stage peri- (Verrill, 1902) (Scleractinia, Mussidae), in the Sào Sebastiâo derm tube the soft and a medu- enveloping body, Channel (23°50'S 45°25'W). We searched for the polyps on the sa stage with a coronal furrow in the exumbrella calcareous substrata with the aid of a stereomicroscope. The 236 F.L. da Silveira & A.C. Morandini - Nausithoe aurea n. sp. collected in the São Table I. Locality data and variability of total periderm tube length of Nausithoe aurea n. sp. scyphistomae initials stand for the authors ofthis Sebastião Channel. The collector paper. Collector Date of Scyphistomae Total periderm Depth sample number cone length (mm) (m) (initials) 04/VII/1995 6 2.85-6.75 6 FLS/ACM FLS/ACM 06/VII/1995 3 3.41—4.73 3 27/VII/1995 16 1.76-8.25 4 FLS/ACM FLS 3 l/VIII/1995 28 1.35-9.18 6 08/1/1996 9 1.42-5.64 9 ACM 10/1/1996 6 1.89-8.10 6 ACM 11/1/1996 6 1.68-5.17 6 ACM scyphistomae were reared in the Centro de Biología Marinha da Descriptive part Universidade de Säo Paulo (CEBIMar USP), at Säo Sebastiäo, SP. They were transferred into and maintained in small petri Nausithoe and aurea sp. n. dishes. The seawater was changed every day the animals fed other with ofthe clam Pis. were every day homogenized gonad (Figs. 1-7, I-II) Perna In mid- perna (Linnaeus, 1767) (Bivalvia, Mytilidae). at from 8 to 27 July 1995,we made the first cultivation Nacional winter, Material examined. - Holotype: MNRJ 2899, Museu to establish the best room temperature (range 20.7°C-26°C) Universidade Federal do Rio de Janeiro, Brazil, State of Sào In from 31 August to 15 October rearing technique. spring, Paulo, Ilhabela,Praia Grande (23°51'S 45°25'W), 12/X/1995, 1 at 21°C- 1995,we all room temperature(range kept specimens cr medusa reared for 29 days at CEBIMar, FLS/ACM coll. 27.6°C) and from 16 October to 6 November 1995, some ma- Paratypes: MNRJ 2900, Museu Nacional Universidade control- ture medusae and their embryos/planulae were kept at Federal do Rio de Janeiro, Brazil, State of Sao Paulo, Ilhabela, led temperature, 22°C. In summer, from 8 to 28 January 1996, coral, Praia Grande, 5/X/1995, 6 m depth, on dead stony 1 the were kept at controlled temperature, 22°C, and a specimens strobilating chain that originated female medusae, reared for 36 12 hr Table I shows several data of the light/dark regime. days, FLS coll.; MNRJ 2901, Brazil, State of Sâo Paulo, sampled. scyphistomae Ilhabela, Praia Grande, 16/X/1995, 6 m depth, on dead stony The from free-swimming ephyrae, strobilating scyphisto- coral, 1 scyphistoma that originated male medusae, reared for dishes and reared to mature were transferred into single mae, 47 days, FLS coll.; MNRJ 2902, Brazil, State of Säo Paulo, The medusae. were fed with homogenized clam gonads. They Ilhabela, Praia Grande, 16/X/1995, 6 m depth, on dead stony and the medusae were intensively fed, to twice a ephyrae up coral, 1 strobilating scyphistoma, reared for 47 days, in whole- for 37 were immersed in day, as long as days. They individually mount preparation, FLS coll.; MNRJ 2903, Brazil, State ofSào observed under the the homogenized food, stereomicroscope Paulo, Ilhabela, Praia Grande, 1 l/X/1995, 6 ephyrae reared for dish with until their stomachs were full, and removed to a new 2 days at CEBIMar, FLS/ACM coll.; MNRJ 2904, Brazil, State clean filtered seawater. the mature female medusae Thereafter, of Säo Paulo, Ilhabela, Praia Grande, 6/X/1995, 6 planulae released The and female medusa were many oocytes. oocytes a reared for 22 days at CEBIMar, FLS/ACM coll.; MNRJ 2905, combined with male medusa. The fertilized ob- a eggs were Brazil, State ofSào Paulo, Ilhabela,Praia Grande, 15/X/1995, 1 served until early planula FLS/ACM stage. 9 medusa reared for 35 days at CEBIMar, coll.; The from free-swimming planuloids, strobilating scyphi- MNRJ 2906, Brazil, State ofSäo Paulo, Ilhabela, Praia Grande, at were transferred into dishes, 24 hr intervals, stomae, single 15/X/1995, 1 9 medusa reared for 20 days at CEBIMar, FLS/ and were observed from then on.A batch of 24 from planuloids, ACM coll. a single scyphistoma, was maintained in the same dish with the at: The Ontario Additional paratypes are deposited Royal polyp. Museum, Invertebrate Zoology (ROMIZ) B2898, B2899, and fixed and in seawater-formalin were Specimens preserved B2900, Canada; The Zoölogisch Museum, University of Am- for electron (SEM). The prepared scanning microscopy sterdam (ZMA) Coel. 8474, Coel. 8475, and Coel. 8476, The tube sectioned over and below the periderm was transversely Netherlands; Zoologisches Museum, Universität Hamburg middle periderm whorl of internal The tube and the ring cusps. (ZMH) CI 132, CI 133, CI 134, CI 135, CI 136, CI 137, and dried and mounted. fragment were dehydrated, critical-point CI 138, Germany; and The Natural History Museum (NHM) coated with 10 of in Balzers S-SCD 050 They were nm gold a 1996.1076, 1996.1077, 1996.1078, 1996.1079-1084 and coater. examined with a Zeiss DSM sputter Specimens were 1996.1085-1094,United Kingdom. 940 SEM. We studied the cnidome of live specimens and of specimens Etymology. - The specific name is derived from preserved in seawater-formalin. Only undischarged nemato- fern- the Latin adjective aureus (= golden yellow, measured. cysts ofpreserved tissues were Contributions to Zoology, 66 (4) - 1997 237 1. Mature male medusa of Nausithoe the 8 testes after 5 months in seawater- Fig. aurea n. sp. (MNRJ 2899) showing (side view, formalin). Scale bar: 1.3 mm. inine aurea), after the color of the pigment spots ture dioecius medusae with 8 brown round in the medusa lappets. adradial gonads, stomach with 4 pouches and filaments 3 presenting up to 13 gastric (typically Diagnosis. - Metagenic. Polyp solitary with in each quadrant); the central filament of each tube dark with small the is than the oth- periderm to light brown, group, oldest, always longer basal disk for attachment, ± 0.47 mm. Proportions ers. Life cycle: ephyrae retained within periderm (diameter/length) at 2 mm height 0.19 ± 0.03, at cone transform into flagellated planuloids. The ± ± and 5 mm height 0.11 0.02, and total length 0.17 same scyphistoma may produce free medusae 0.06 mm. Tube surface with a pattern of many planuloids alternately. varied less transverse rings, more or prominent, - The total diameter with longitudinal striations. Inner spines or cusps Medusa. is 5 mm. The diam- 4 the disk are arranged in distinct whorls of sixteen cusps; eter of central (from rhopalium to intermediate and is of the coronal furrow large (perradius), 4 (interradius) rhopalium) 3.69 mm; 1.9 8 small (adradius).