Experiments in nature and laboratory observations with Nausithoe aurea (Scyphozoa: Coronatae) support the concept of perennation by tissue saving and confirm dormancy

Silveira, Fábio Lang da; Jarms, Gerhard; Morandini, André Carrara

doi:10.1590/S1676-06032002000200009

Abstracts

Stephanocyphistomae of Nausithoe aurea from São Paulo State, Brazil (in subtropical western South Atlantic wa-ters), were relocated with their substrata in nature to study their survivorship under control and and experimental series - i.e. the polyps in the original orientation and inverted, and in each series exposed and buried polyps. We found that N. aurea survives over 13 months in nature, between 1/3 - 1/4 of 268 stephanoscyphistomae as normal feeding polyps, by segmentation produces planuloids and rejuvenates the polyps - an additional explanation for clustering of the solitary stephanocyphistomae. Dormant living tissues within the periderm of the tube were considered resting stages. The results support the concept that coronates in general have the capacity to save all living tissue and transform it to the energy saving sessile stage - the perennial polyp.

Cnidaria; Scyphozoa; Nausithoe aurea; perennation; resting stages; Brazil

Estefanocifístomas de Nausithoe aurea no Estado de São Paulo, Brasil (em águas subtropicais do Atlântico Sul ocidental), foram realocados com os seus substratos na natureza para estudar a sobrevivência em séries controle e experimental - isto é, os pólipos na posição original e invertidos e em cada série pólipos expostos e enterrados. Verificamos que N. aurea sobrevive por 13 meses na natureza, entre 1/3 - 1/4 de 268 estefanocifístomas como pólipos normais capazes de alimentação, por segmentação produz planulóides e rejuvenesce o pólipo - uma explicação adicional para a ocorrência agregada dos estefanocifístomas solitários. Tecidos vivos dormentes dentro da periderme do tubo foram considerados como estágios de quietação. Os resultados reforçam a idéia de que os coronados em geral têm a capacidade de conservar todo o tecido vivo e de transformá-lo durante o estágio séssil mais econômico energeticamente - o pólipo perene.

Cnidaria; Scyphozoa; Nausithoe aurea; perenização; estágios de quietação; Brasil

ARTICLES

Experiments in nature and laboratory observations with Nausithoe aurea (Scyphozoa: Coronatae) support the concept of perennation by tissue saving and confirm dormancy

Fábio Lang da Silveira^I(correspondence author; correspondência para); Gerhard Jarms^II; André Carrara Morandini^I

^IDepartamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11461, 05422-970, São Paulo, SP, Brasil. Tel.:+55-11-30917619 Fax: +55-11-30917513

^IIZoologisches Institut und Zoologisches Museum, Universität Hamburg, Martin-Luther-King Platz 3, 20146 Hamburg, Germany, Tel.: +49-40-428382086 Fax: +49-40-428382086 e-mails: fldsilve@usp.br, Gerhard.Jarms@zoologie.uni-hamburg.de, acmorand@usp.br

ABSTRACT

Stephanocyphistomae of Nausithoe aurea from São Paulo State, Brazil (in subtropical western South Atlantic wa-ters), were relocated with their substrata in nature to study their survivorship under control and and experimental series - i.e. the polyps in the original orientation and inverted, and in each series exposed and buried polyps. We found that N. aurea survives over 13 months in nature, between 1/3 - 1/4 of 268 stephanoscyphistomae as normal feeding polyps, by segmentation produces planuloids and rejuvenates the polyps - an additional explanation for clustering of the solitary stephanocyphistomae. Dormant living tissues within the periderm of the tube were considered resting stages. The results support the concept that coronates in general have the capacity to save all living tissue and transform it to the energy saving sessile stage - the perennial polyp.

Key-words: Cnidaria, Scyphozoa, Nausithoe aurea, perennation, resting stages, Brazil.

RESUMO

Estefanocifístomas de Nausithoe aurea no Estado de São Paulo, Brasil (em águas subtropicais do Atlântico Sul ocidental), foram realocados com os seus substratos na natureza para estudar a sobrevivência em séries controle e experimental - isto é, os pólipos na posição original e invertidos e em cada série pólipos expostos e enterrados. Verificamos que N. aurea sobrevive por 13 meses na natureza, entre 1/3 - 1/4 de 268 estefanocifístomas como pólipos normais capazes de alimentação, por segmentação produz planulóides e rejuvenesce o pólipo - uma explicação adicional para a ocorrência agregada dos estefanocifístomas solitários. Tecidos vivos dormentes dentro da periderme do tubo foram considerados como estágios de quietação. Os resultados reforçam a idéia de que os coronados em geral têm a capacidade de conservar todo o tecido vivo e de transformá-lo durante o estágio séssil mais econômico energeticamente - o pólipo perene.

Palavras-chave: Cnidaria, Scyphozoa, Nausithoe aurea, perenização, estágios de quietação, Brasil.

Full text available only in PDF format.

Texto completo disponível apenas em PDF.

Acknowledgements

This work was supported by Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP 99/12433-0) and CAPES/USP PROAP/2000. ACM received financial support from FAPESP (97/03325-3, 97/08137-0 and 99/05374-7). We thank an anonymous reviewer from FAPESP for sugges-tions to improve the experimental design at the start of the work and for later comments on the project. We are indebted to MSc Helena Krieg Boscolo for help in the fieldwork and to Dr. Márcio Custódio ( CEBIMar USP) for the identifica-tion of the sponge. We thank CEBIMar USP for providing the required facilities to obtain and examine the material throughout the work.

References

BUMANN, D. & G. JARMS, 1997. Localization of digestion activities in polyps of Nausithoe planulophora and Thecoscyphus zibrowii (Coronatae, Scyphozoa, Cnidaria). Helgoländer Meeresunter., 51: 477-485.

CÁCERES, C.E., 1997. Dormancy in invertebrates. Invert. Biol., 116(4): 371-383.

COLLINS, A.G., 2002. Phylogeny of the Medusozoa and the evolution of cnidarian life cycles. J. Evol. Biol., 15: 418-432.

HOLST, S. 2002. Können Polypen der Coronatae (Cnidaria, Scyphozoa) Fremdkörper aus ihren Röhren entfernen?. Diplomarbeit, Zoologisches Institut und Museum, Fachbereich Biologie der Universität Hamburg, Hamburg, 99p.

JARMS, G., 1990. Neubeschreibung dreier Arten der Gattung Nausithoe (Coronata, Scyphozoa) sowie Wiederbeschreibung der Art Nausithoe marginata Kölliker, 1853. Mitt. hamb. zool. Mus. Inst., 87: 7-39.

JARMS, G., 1997. The polyps of Coronatae (Scyphozoa), a review and some new results. In: den HARTOG, J.C. (ed.), Proceedings of the 6^th International Conference on Coelenterate Biology 1995. Nationaal Natuurhistorisch Museum, Leiden, p. 271-278.

JARMS, G., 2001. The life cycle of Nausithoe hagenbecki sp. nov. (Scyphozoa, Coronatae). Mitt. hamb. zool. Mus. Inst., 98: 13-22.

JARMS, G.; U. BÅMSTEDT ; H. TIEMANN; M.B. MARTINUSSEN & J.H. FOSSÅ, 1999. The holopelagic life cycle of the deep-sea medusa Periphylla periphyll a (Scyphozoa, Coronatae). Sarsia, 84: 55-65.

JARMS, G.; A.C. MORANDINI & F.L. da SILVEIRA, 2002. Polyps of the families Atorellidae and Nausithoidae (Scyphozoa: Coronatae). Biota Neotropica, 2(1): 11p.

KAWAGUTI, S. & A. MATSUNO, 1981. A new species of the Coronatae, Scyphozoa, from the Japan sea; Atorella japonica n. sp. Bull. Kawasaki Parmed. Coll., 1: 15-21.

KOMAI, T., 1936 On another form of Stephanoscyphus found in the waters of Japan. Mem. Coll. Sci. Kyoto (Ser. B), 11(3): 175-83.

MORANDINI, A.C., 1999. Gametogênese e desenvolvimento embrionário de Nausithoe aurea (Scyphozoa, Coronatae) do Canal de São Sebastião - SP. MSc Dissertation, Instituto de Biociências, Universidade de São Paulo. i-iv , 136 p.

MORANDINI, A.C. & F.L. da SILVEIRA, 2001a. New ob-servations and new record of Nausithoe aurea (Scyphozoa, Coronatae). Pap. Av. Zool., 41(27): 519-527.

MORANDINI, A.C. & F.L. da SILVEIRA, 2001b. Sexual re-production of Nausithoe aurea (Scyphozoa, Coronatae). Gametogenesis, egg release, embryonic development, and gastrulation. Sci. Mar., 65(2): 139-149.

ORTIZ-CORP'S, E.; C.E. CUTRESS & B.M. CUTRESS, 1987. Life history of the coronate scyphozoan Linuche unguiculata (

KOMAI, T., 1936 On another form of Stephanoscyphus found in the waters of Japan. Mem. Coll. Sci. Kyoto (Ser. B), 11(3): 175-83.

MORANDINI, A.C., 1999. Gametogênese e desenvolvimento embrionário de Nausithoe aurea (Scyphozoa, Coronatae) do Canal de São Sebastião - SP. MSc Dissertation, Instituto de Biociências, Universidade de São Paulo. i-iv , 136 p.

MORANDINI, A.C. & F.L. da SILVEIRA, 2001a. New ob-servations and new record of Nausithoe aurea (Scyphozoa, Coronatae). Pap. Av. Zool., 41(27): 519-527.

MORANDINI, A.C. & F.L. da SILVEIRA, 2001b. Sexual re-production of Nausithoe aurea (Scyphozoa, Coronatae). Gametogenesis, egg release, embryonic development, and gastrulation. Sci. Mar., 65(2): 139-149.

ORTIZ-CORP'S, E.; C.E. CUTRESS & B.M. CUTRESS, 1987. Life history of the coronate scyphozoan Linuche unguiculata (Swartz, 1788). Carib. Jour. Sci., 23: 432-443.

SILVEIRA, F.L. da. & A.C. MORANDINI, 1997. Nausithoe aurea n. sp. (Scyphozoa: Coronatae: Nausithoidae), a species with two pathways of reproduction after strobi-lation: sexual and asexual. Cont. Zool., 66(4): 235-246.

SILVEIRA, F.L. da. & A.C. MORANDINI, 1998a. Asexual reproduction in Linuche unguiculata (Swartz, 1788) (Scyphozoa: Coronatae) by planuloid formation through strobilation and segmentation. Proc. biol. Soc. Wash., 111(4): 781-794.

SILVEIRA, F.L. da. & A.C. MORANDINI, 1998b. New ob-servations on dormancy mechanisms in Linuche unguiculata (Swartz, 1788) (Scyphozoa: Coronatae). Bol. Mus. Nac., N.S., Zool., 393: 1-7.

SÖTJE, I. & G. JARMS, 1999. Description of Thecoscyphus zibrowii Werner, 1984 (Scyphozoa, Coronatae) with re-marks on the ontogeny. Mitt. Hamb. Zool. Mus. Inst., 96:5-13.

WERNER, B. 1966. Stephanoscyphus (Scyphozoa, Coronatae) und seine direkte Abstammung von den fossilen Conulata. Helgoländer wiss. Meeresunters., 13: 317-347.

WERNER, B., 1971. Stephanoscyphus planulophorus n.spec., eine neuer Scyphopolyp mit einem neun Entwichlungsmodus. Helgoländer wiss. Meeresunters., 22: 120-140.

WERNER, B., 1973. New investigations on systematics and evolution of the class Scyphozoa and the Phylum Cnidaria. Publs Seto mar. biol. Lab., 20: 35-61.

WERNER, B. 1974. Stephanoscyphus eumedusoides n. sp. (Syphozoa, Coronatae), ein Höhlenpolyp mit einem neuen Entwicklungsmodus. Helgoländer wiss.Meeresunters., 26: 434-463.

WERNER, B. 1979. Coloniality in the Scyphozoa: Cnidaria. In: LARWOOD, G. & B.R. ROSEN (eds.), Biology and systematics of colonial organisms. Academic Press, Lon-don, p. 81-103.

Date Received 07/19/2002

Revised 09/23/2002

Accepted 10/02/2002

BUMANN, D. & G. JARMS, 1997. Localization of digestion activities in polyps of Nausithoe planulophora and Thecoscyphus zibrowii (Coronatae, Scyphozoa, Cnidaria). Helgoländer Meeresunter., 51: 477-485.
CÁCERES, C.E., 1997. Dormancy in invertebrates. Invert. Biol., 116(4): 371-383.
COLLINS, A.G., 2002. Phylogeny of the Medusozoa and the evolution of cnidarian life cycles. J. Evol. Biol., 15: 418-432.
HOLST, S. 2002. Können Polypen der Coronatae (Cnidaria, Scyphozoa) Fremdkörper aus ihren Röhren entfernen?. Diplomarbeit, Zoologisches Institut und Museum, Fachbereich Biologie der Universität Hamburg, Hamburg, 99p.
JARMS, G., 1990. Neubeschreibung dreier Arten der Gattung Nausithoe (Coronata, Scyphozoa) sowie Wiederbeschreibung der Art Nausithoe marginata Kölliker, 1853. Mitt. hamb. zool. Mus. Inst., 87: 7-39.
JARMS, G., 1997. The polyps of Coronatae (Scyphozoa), a review and some new results. In: den HARTOG, J.C. (ed.), Proceedings of the 6^th International Conference on Coelenterate Biology 1995. Nationaal Natuurhistorisch Museum, Leiden, p. 271-278.
JARMS, G., 2001. The life cycle of Nausithoe hagenbecki sp. nov. (Scyphozoa, Coronatae). Mitt. hamb. zool. Mus. Inst., 98: 13-22.
JARMS, G.; U. BÅMSTEDT ; H. TIEMANN; M.B. MARTINUSSEN & J.H. FOSSÅ, 1999. The holopelagic life cycle of the deep-sea medusa Periphylla periphyll a (Scyphozoa, Coronatae). Sarsia, 84: 55-65.
JARMS, G.; A.C. MORANDINI & F.L. da SILVEIRA, 2002. Polyps of the families Atorellidae and Nausithoidae (Scyphozoa: Coronatae). Biota Neotropica, 2(1): 11p.
KAWAGUTI, S. & A. MATSUNO, 1981. A new species of the Coronatae, Scyphozoa, from the Japan sea; Atorella japonica n. sp. Bull. Kawasaki Parmed. Coll., 1: 15-21.
KOMAI, T., 1936 On another form of Stephanoscyphus found in the waters of Japan. Mem. Coll. Sci. Kyoto (Ser. B), 11(3): 175-83.
MORANDINI, A.C., 1999. Gametogênese e desenvolvimento embrionário de Nausithoe aurea (Scyphozoa, Coronatae) do Canal de São Sebastião - SP. MSc Dissertation, Instituto de Biociências, Universidade de São Paulo. i-iv , 136 p.
MORANDINI, A.C. & F.L. da SILVEIRA, 2001a. New ob-servations and new record of Nausithoe aurea (Scyphozoa, Coronatae). Pap. Av. Zool., 41(27): 519-527.
MORANDINI, A.C. & F.L. da SILVEIRA, 2001b. Sexual re-production of Nausithoe aurea (Scyphozoa, Coronatae). Gametogenesis, egg release, embryonic development, and gastrulation. Sci. Mar., 65(2): 139-149.
ORTIZ-CORP'S, E.; C.E. CUTRESS & B.M. CUTRESS, 1987. Life history of the coronate scyphozoan Linuche unguiculata (
KOMAI, T., 1936 On another form of Stephanoscyphus found in the waters of Japan. Mem. Coll. Sci. Kyoto (Ser. B), 11(3): 175-83.
MORANDINI, A.C., 1999. Gametogênese e desenvolvimento embrionário de Nausithoe aurea (Scyphozoa, Coronatae) do Canal de São Sebastião - SP. MSc Dissertation, Instituto de Biociências, Universidade de São Paulo. i-iv , 136 p.
MORANDINI, A.C. & F.L. da SILVEIRA, 2001a. New ob-servations and new record of Nausithoe aurea (Scyphozoa, Coronatae). Pap. Av. Zool., 41(27): 519-527.
MORANDINI, A.C. & F.L. da SILVEIRA, 2001b. Sexual re-production of Nausithoe aurea (Scyphozoa, Coronatae). Gametogenesis, egg release, embryonic development, and gastrulation. Sci. Mar., 65(2): 139-149.
ORTIZ-CORP'S, E.; C.E. CUTRESS & B.M. CUTRESS, 1987. Life history of the coronate scyphozoan Linuche unguiculata (Swartz, 1788). Carib. Jour. Sci., 23: 432-443.
SILVEIRA, F.L. da. & A.C. MORANDINI, 1997. Nausithoe aurea n. sp. (Scyphozoa: Coronatae: Nausithoidae), a species with two pathways of reproduction after strobi-lation: sexual and asexual. Cont. Zool., 66(4): 235-246.
SILVEIRA, F.L. da. & A.C. MORANDINI, 1998a. Asexual reproduction in Linuche unguiculata (Swartz, 1788) (Scyphozoa: Coronatae) by planuloid formation through strobilation and segmentation. Proc. biol. Soc. Wash., 111(4): 781-794.
SILVEIRA, F.L. da. & A.C. MORANDINI, 1998b. New ob-servations on dormancy mechanisms in Linuche unguiculata (Swartz, 1788) (Scyphozoa: Coronatae). Bol. Mus. Nac., N.S., Zool., 393: 1-7.
SÖTJE, I. & G. JARMS, 1999. Description of Thecoscyphus zibrowii Werner, 1984 (Scyphozoa, Coronatae) with re-marks on the ontogeny. Mitt. Hamb. Zool. Mus. Inst., 96:5-13.
WERNER, B. 1966. Stephanoscyphus (Scyphozoa, Coronatae) und seine direkte Abstammung von den fossilen Conulata. Helgoländer wiss. Meeresunters., 13: 317-347.
WERNER, B., 1971. Stephanoscyphus planulophorus n.spec., eine neuer Scyphopolyp mit einem neun Entwichlungsmodus. Helgoländer wiss. Meeresunters., 22: 120-140.
WERNER, B., 1973. New investigations on systematics and evolution of the class Scyphozoa and the Phylum Cnidaria. Publs Seto mar. biol. Lab., 20: 35-61.
WERNER, B. 1974. Stephanoscyphus eumedusoides n. sp. (Syphozoa, Coronatae), ein Höhlenpolyp mit einem neuen Entwicklungsmodus. Helgoländer wiss.Meeresunters., 26: 434-463.
WERNER, B. 1979. Coloniality in the Scyphozoa: Cnidaria. In: LARWOOD, G. & B.R. ROSEN (eds.), Biology and systematics of colonial organisms. Academic Press, Lon-don, p. 81-103.

Publication Dates

Publication in this collection
11 June 2013
Date of issue
2002

History

Received
19 July 2002
Reviewed
23 Sept 2002
Accepted
02 Oct 2002

This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License.

[1] BUMANN, D. & G. JARMS, 1997. Localization of digestion activities in polyps of Nausithoe planulophora and Thecoscyphus zibrowii (Coronatae, Scyphozoa, Cnidaria). Helgoländer Meeresunter., 51: 477-485.

[2] CÁCERES, C.E., 1997. Dormancy in invertebrates. Invert. Biol., 116(4): 371-383.

[3] COLLINS, A.G., 2002. Phylogeny of the Medusozoa and the evolution of cnidarian life cycles. J. Evol. Biol., 15: 418-432.

[4] HOLST, S. 2002. Können Polypen der Coronatae (Cnidaria, Scyphozoa) Fremdkörper aus ihren Röhren entfernen?. Diplomarbeit, Zoologisches Institut und Museum, Fachbereich Biologie der Universität Hamburg, Hamburg, 99p.

[5] JARMS, G., 1990. Neubeschreibung dreier Arten der Gattung Nausithoe (Coronata, Scyphozoa) sowie Wiederbeschreibung der Art Nausithoe marginata Kölliker, 1853. Mitt. hamb. zool. Mus. Inst., 87: 7-39.

[6] JARMS, G., 1997. The polyps of Coronatae (Scyphozoa), a review and some new results. In: den HARTOG, J.C. (ed.), Proceedings of the 6^th International Conference on Coelenterate Biology 1995. Nationaal Natuurhistorisch Museum, Leiden, p. 271-278.

[7] JARMS, G., 2001. The life cycle of Nausithoe hagenbecki sp. nov. (Scyphozoa, Coronatae). Mitt. hamb. zool. Mus. Inst., 98: 13-22.

[8] JARMS, G.; U. BÅMSTEDT ; H. TIEMANN; M.B. MARTINUSSEN & J.H. FOSSÅ, 1999. The holopelagic life cycle of the deep-sea medusa Periphylla periphyll a (Scyphozoa, Coronatae). Sarsia, 84: 55-65.

[9] JARMS, G.; A.C. MORANDINI & F.L. da SILVEIRA, 2002. Polyps of the families Atorellidae and Nausithoidae (Scyphozoa: Coronatae). Biota Neotropica, 2(1): 11p.

[10] KAWAGUTI, S. & A. MATSUNO, 1981. A new species of the Coronatae, Scyphozoa, from the Japan sea; Atorella japonica n. sp. Bull. Kawasaki Parmed. Coll., 1: 15-21.

[11] KOMAI, T., 1936 On another form of Stephanoscyphus found in the waters of Japan. Mem. Coll. Sci. Kyoto (Ser. B), 11(3): 175-83.

[12] MORANDINI, A.C., 1999. Gametogênese e desenvolvimento embrionário de Nausithoe aurea (Scyphozoa, Coronatae) do Canal de São Sebastião - SP. MSc Dissertation, Instituto de Biociências, Universidade de São Paulo. i-iv , 136 p.

[13] MORANDINI, A.C. & F.L. da SILVEIRA, 2001a. New ob-servations and new record of Nausithoe aurea (Scyphozoa, Coronatae). Pap. Av. Zool., 41(27): 519-527.

[14] MORANDINI, A.C. & F.L. da SILVEIRA, 2001b. Sexual re-production of Nausithoe aurea (Scyphozoa, Coronatae). Gametogenesis, egg release, embryonic development, and gastrulation. Sci. Mar., 65(2): 139-149.

[15] ORTIZ-CORP'S, E.; C.E. CUTRESS & B.M. CUTRESS, 1987. Life history of the coronate scyphozoan Linuche unguiculata (

[16] KOMAI, T., 1936 On another form of Stephanoscyphus found in the waters of Japan. Mem. Coll. Sci. Kyoto (Ser. B), 11(3): 175-83.

[17] MORANDINI, A.C., 1999. Gametogênese e desenvolvimento embrionário de Nausithoe aurea (Scyphozoa, Coronatae) do Canal de São Sebastião - SP. MSc Dissertation, Instituto de Biociências, Universidade de São Paulo. i-iv , 136 p.

[18] MORANDINI, A.C. & F.L. da SILVEIRA, 2001a. New ob-servations and new record of Nausithoe aurea (Scyphozoa, Coronatae). Pap. Av. Zool., 41(27): 519-527.

[19] MORANDINI, A.C. & F.L. da SILVEIRA, 2001b. Sexual re-production of Nausithoe aurea (Scyphozoa, Coronatae). Gametogenesis, egg release, embryonic development, and gastrulation. Sci. Mar., 65(2): 139-149.

[20] ORTIZ-CORP'S, E.; C.E. CUTRESS & B.M. CUTRESS, 1987. Life history of the coronate scyphozoan Linuche unguiculata (Swartz, 1788). Carib. Jour. Sci., 23: 432-443.

[21] SILVEIRA, F.L. da. & A.C. MORANDINI, 1997. Nausithoe aurea n. sp. (Scyphozoa: Coronatae: Nausithoidae), a species with two pathways of reproduction after strobi-lation: sexual and asexual. Cont. Zool., 66(4): 235-246.

[22] SILVEIRA, F.L. da. & A.C. MORANDINI, 1998a. Asexual reproduction in Linuche unguiculata (Swartz, 1788) (Scyphozoa: Coronatae) by planuloid formation through strobilation and segmentation. Proc. biol. Soc. Wash., 111(4): 781-794.

[23] SILVEIRA, F.L. da. & A.C. MORANDINI, 1998b. New ob-servations on dormancy mechanisms in Linuche unguiculata (Swartz, 1788) (Scyphozoa: Coronatae). Bol. Mus. Nac., N.S., Zool., 393: 1-7.

[24] SÖTJE, I. & G. JARMS, 1999. Description of Thecoscyphus zibrowii Werner, 1984 (Scyphozoa, Coronatae) with re-marks on the ontogeny. Mitt. Hamb. Zool. Mus. Inst., 96:5-13.

[25] WERNER, B. 1966. Stephanoscyphus (Scyphozoa, Coronatae) und seine direkte Abstammung von den fossilen Conulata. Helgoländer wiss. Meeresunters., 13: 317-347.

[26] WERNER, B., 1971. Stephanoscyphus planulophorus n.spec., eine neuer Scyphopolyp mit einem neun Entwichlungsmodus. Helgoländer wiss. Meeresunters., 22: 120-140.

[27] WERNER, B., 1973. New investigations on systematics and evolution of the class Scyphozoa and the Phylum Cnidaria. Publs Seto mar. biol. Lab., 20: 35-61.

[28] WERNER, B. 1974. Stephanoscyphus eumedusoides n. sp. (Syphozoa, Coronatae), ein Höhlenpolyp mit einem neuen Entwicklungsmodus. Helgoländer wiss.Meeresunters., 26: 434-463.

[29] WERNER, B. 1979. Coloniality in the Scyphozoa: Cnidaria. In: LARWOOD, G. & B.R. ROSEN (eds.), Biology and systematics of colonial organisms. Academic Press, Lon-don, p. 81-103.