Marine Biology

, Volume 148, Issue 5, pp 923–931 | Cite as

Sexual reproduction of the solitary sunset cup coral Leptopsammia pruvoti (Scleractinia, Dendrophylliidae) in the Mediterranean. 2. Quantitative aspects of the annual reproductive cycle

  • S. GoffredoEmail author
  • V. Airi
  • J. Radetić
  • F. Zaccanti
Research Article


There are few data on the reproductive biology of coral species living in temperate zones, particularly in the Mediterranean Sea. Leptopsammia pruvoti is a solitary coral that is commonly found in sea caves and under overhangs throughout the Mediterranean basin and along European coasts from Portugal to southern England. In this paper, we describe its annual reproductive cycle in the eastern Ligurian Sea near the city of Leghorn (Tuscany, Italy). Polyps were sexually mature at 3 mm in length (maximum diameter of the oral disc), were dioecious with a sex ratio of 1:1, and brooded their larvae. The maturation of spermaries took 12 months and oocytes 24 months. The rate of gonad development increased significantly from November to January, fertilization occurred from January to April and planulation during May and June. Seasonal variations in water temperature and photoperiod may have played an important role in regulating reproductive events. The amount of energy devoted to male gametogenesis (quantified by gonad index) was significantly higher in the sex separated species L. pruvoti than in the hermaphroditic dendrophylliid Balanophyllia europaea, whose reproduction has been studied in a previous work. We hypothesize that this difference is due to the contrasting sexuality or fertilization biology of these two species (cross-fertilization in the sex separated L. pruvoti versus possibly self-fertilization in the hermaphroditic B. europaea). Greater male sexual allocation in reproductive strategies characterized by dioecism or cross-fertilization when compared to those characterized by hermaphroditism or self-fertilization is common in plant mating systems. In relation to other solitary dendrophylliids, L. pruvoti presents an ‘r’-reproductive strategy.


Reproductive Cycle Reproductive Strategy Gonadal Development Mature Oocyte Oral Disc 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



We dedicate this study to Mario Pasquini, great friend, best diver, and our very important advisor and collaborator, who left us on 19 June, 2005. We also wish to thank E. Manzardo, S. Arnone, L. Tomesani, M. Longagnani, L. Podda, O. Panaro, for their valuable scuba assistance in collecting monthly samples. R. Falconi gave assistance in defining laboratory guidelines. G. Marconi provided significant help in preparing histological samples. J. P. Gilmour (Australian Institute of Marine Science) revised and significantly improved this paper. We also thank two anonymous reviewers for their valuable revision. The Marine Science Group ( provided scientific, technical, and logistical support. The Bologna Scuba Team gave assistance for the dives. Our research was supported by the Ministero dell’Istruzione, dell’Università e della Ricerca (MIUR), by the Associazione dei Tour Operator Italiani (ASTOI) and by Scuba Schools International, Italy (SSI). The experiments complied with the current laws of Italy.


  1. Acosta A, Zea S (1997) Sexual reproduction of the reef coral Montastrea cavernosa (Scleractinia: Faviidae) in the Santa Marta area, Caribbean coast of Colombia. Mar Biol 128:141–148CrossRefGoogle Scholar
  2. Avian M, Boero F, Mills C, Rossi L, Rottini-Sandrini L (1995) Cnidaria, Ctenophora. In: Minelli A, Ruffo S, La Posta S (eds) Checklist delle specie della fauna italiana. Edizioni Calderini, Bologna, pp 1–38Google Scholar
  3. Babcock RC, Bull G, Harrison PL, Heyward AJ, Oliver JK,Wallace CC, Willis BL (1986) Synchronous spawnings of 105 scleractinian coral species on the Great Barrier Reef. Mar Biol 90:379–394CrossRefGoogle Scholar
  4. Babcock RC, Willis BL, Simpson CJ (1994) Mass spawning of corals on high latitude coral reef. Coral Reefs 13:161–169CrossRefGoogle Scholar
  5. Bacci G (1975) Genetic and environmental controls of sex determination in marine animals. Pubbl Staz Zool Napoli 39:366–376Google Scholar
  6. Beauchamp KA (1993) Gametogenesis, brooding and planulation in laboratory populations of a temperate scleractinian coral Balanophyllia elegans maintained under contrasting photoperiod regimes. Invert Repr Dev 23:171–182CrossRefGoogle Scholar
  7. Beiring EA, Lasker HR (2000) Egg production by colonies of a gorgonian coral. Mar Ecol Prog Ser 196:169–177CrossRefGoogle Scholar
  8. Benayahu Y, Weil D, Kleinman M (1990) Radiation of broadcasting and brooding patterns in coral reef alcyonaceans. In: Hoshi M, Yamashita O (eds) Advances in invertebrate reproduction. Elsevier Science Publishers BV (Biomed Div), Amsterdam, pp 323–328Google Scholar
  9. Cairns SD (1999) Species richness of recent Scleractinia. Atoll Res Bull 459:1–12CrossRefGoogle Scholar
  10. Cairns SD, Hoeksema BW, Van Der Land J (1999) Appendix: List of extant stony corals. Atoll Res Bull 459:13–46CrossRefGoogle Scholar
  11. Charnov EL (1982) The theory of sex allocation. Princeton University Press, PrincetonGoogle Scholar
  12. Chomsky O, Kamenir Y, Hyams M, Dubinsky Z, Chadwick-Furman NE (2004) Effects of temperature on growth rate and body size in the Mediterranean Sea anemone Actinia equina. J Exp Mar Biol Ecol 313:63–73CrossRefGoogle Scholar
  13. Clayton PD, Collins DJ (1992) Reproduction and feeding ethology of a tropical, intertidal sand-dwelling anemone (Actinoporus elongatus, Carlgren, 1900). Hydrobiologia 237:31–38CrossRefGoogle Scholar
  14. Coma R, Ribes M, Cabala M, Gili JM (1995) Reproduction and cycle of gonadal development in the Mediterranean gorgonian Paramuricea clavata. Mar Ecol Prog Ser 117:173–183CrossRefGoogle Scholar
  15. Cruden RW (1977) Pollen–ovule ratios: a conservative indicator of breeding systems in flowering plants. Evolution 31:32–46CrossRefGoogle Scholar
  16. Dobson AP (1998) Conservation and biodiversity. Scientific American library, New YorkGoogle Scholar
  17. Fadlallah YH (1983a) Sexual reproduction, development and larval biology in scleractinian corals: a review. Coral Reefs 2:129–150CrossRefGoogle Scholar
  18. Fadlallah YH, Pearse JS (1982a) Sexual reproduction in solitary corals: overlapping oogenic and brooding cycles, and benthic planulas in Balanophyllia elegans. Mar Biol 71:223–231CrossRefGoogle Scholar
  19. Fadlallah YH, Pearse JS (1982b) Sexual reproduction in solitary corals: synchronous gametogenesis and broadcast spawning in Paracyathus stearnsii. Mar Biol 71:233–239CrossRefGoogle Scholar
  20. Fan TY, Dai CF (1995) Reproductive ecology of the scleractinian coral Echinopora lamellosa in northern and southern Taiwan. Mar Biol 123:565–572CrossRefGoogle Scholar
  21. Fan TY, Dai CF (1998) Sexual reproduction of the scleractinian coral Merulina ampliata in southern Taiwan. Bull Mar Sci 62:897–904Google Scholar
  22. Fautin DG (1992) Cnidaria. In: Adiyodi KJ, Adiyodi RG (eds) Reproductive biology of invertebrates. 5. Sexual differentiation and behaviour. Wiley, New York, pp 31–52Google Scholar
  23. Fonda-Umani S (1992) Successioni fitoplantoniche, micro e mesozooplanctoniche nell’Alto Adriatico. In: Marchetti R, Cotta-Ramusino M (eds) Atti del V congresso SITE. Società Italiana di Ecologia, Parma, pp 221–246Google Scholar
  24. Fujiwara M, Caswell (2001) Demography of endangered North Atlantic right whale. Nature 414:537–541CrossRefGoogle Scholar
  25. Giese AC, Pearse JS (1974) Introduction: general principles. In: Giese AC, Pearse JS (eds) Reproduction of marine invertebrates. I. Acoelomate and pseudocoelomate metazoans. Academic, New York, pp 1–49Google Scholar
  26. Glynn PW, Colley SB, Ting JH, Maté JL, Guzman HM (2000) Reef coral reproduction in the eastern Pacific: Costa Rica, Panama and Galapagos Islands (Ecuador). IV. Agariciidae, recruitment and recovery of Pavona varians and Pavona sp.a. Mar Biol 136:785–805CrossRefGoogle Scholar
  27. Goffredo S, Telò T (1998) Hermaphroditism and brooding in the solitary coral Balanophyllia europaea (Cnidaria, Anthozoa, Scleractinia). Ital J Zool 65:159–165CrossRefGoogle Scholar
  28. Goffredo S, Zaccanti F (2004) Laboratory observations of larval behavior and metamorphosis in the Mediterranean solitary coral Balanophyllia europaea (Scleractinia, Dendrophylliidae). Bull Mar Sci 74:449–458Google Scholar
  29. Goffredo S, Telò T, Scanabissi F (2000) Ultrastructural observations of the spermatogenesis of the hermaphroditic solitary coral Balanophyllia europaea (Anthozoa, Scleractinia). Zoomorphology 119:231–240CrossRefGoogle Scholar
  30. Goffredo S, Arnone S, Zaccanti F (2002) Sexual reproduction in the Mediterranean solitary coral Balanophyllia europaea (Scleractinia, Dendrophylliidae). Mar Ecol Prog Ser 229:83–94CrossRefGoogle Scholar
  31. Goffredo S, Mattioli G, Zaccanti F (2004a) Growth and population dynamics model of the Mediterranean solitary coral Balanophyllia europaea (Scleractinia, Dendrophylliidae). Coral Reefs 23:433–443CrossRefGoogle Scholar
  32. Goffredo S, Mezzomonaco L, Zaccanti F (2004b) Genetic differentiation among populations of the Mediterranean hermaphroditic brooding coral Balanophyllia europaea (Scleractinia, Dendrophylliidae). Mar Biol 145:1075–1083CrossRefGoogle Scholar
  33. Goffredo S, Radetić J, Airi V, Zaccanti F (2005) Sexual reproduction of the solitary sunset cup coral Leptopsammia pruvoti (Scleractinia, Dendrophylliidae) in the Mediterranean. 1. Morphological aspects of gametogenesis and ontogenesis. Mar Biol 147:485–495CrossRefGoogle Scholar
  34. Hall VR, Hughes TP (1996) Reproductive strategies of modular organisms: comparative studies of reef-building corals. Ecology 77:950–963CrossRefGoogle Scholar
  35. Harrison PL (1985) Sexual characteristics of scleractinian corals: systematic and evolutionary implications. In: Proceedings of the 5th international coral reef symposium, Tahiti, 4:337–342Google Scholar
  36. Harrison PL,Wallace CC (1990) Reproduction, dispersal and recruitment of scleractinian corals. In: Dubinsky Z (ed) Ecosystem of the world. 25. Coral reefs. Elsevier, Amsterdam, pp 133–207Google Scholar
  37. Harrison PL, Babcock RC, Bull GD, Oliver JK, Wallace CC,Willis BL (1984) Mass spawning in tropical reef corals. Science 223:1186–1189CrossRefGoogle Scholar
  38. Harvell CD, Grosberg RK (1988) The timing of sexual maturity in clonal animals. Ecology 69:1855–1864CrossRefGoogle Scholar
  39. Heltzel PS, Babcock RC (2002) Sexual reproduction, larval development and benthic planulae of the solitary coral Monomyces rubrum (Scleractinia: Anthozoa). Mar Biol 140:659–667CrossRefGoogle Scholar
  40. Jürgens A, Witt T, Gottsberger G (2002) Pollen grain numbers, ovule numbers and pollen–ovule ratios in Caryophylloideae: correlation with breeding system, pollination, life form, style number, and sexual system. Sex Plant Reprod 14:279–289CrossRefGoogle Scholar
  41. Kramarsky-Winter E, Loya Y (1998) Reproductive strategies of two fungiid corals from the northern Red Sea: environmental constraints? Mar Ecol Prog Ser 174:175–182CrossRefGoogle Scholar
  42. Kruger A, Schleyer MH (1998) Sexual reproduction in the coral Pocillopora verrucosa (Cnidaria: Scleractinia) in KwaZulu-Natal, South Africa. Mar Biol 132:703–710CrossRefGoogle Scholar
  43. Lacaze-Duthiers H (1873) Développment des coralliaires. Actinaires à Polypiers. Arch Zool Exp Gén 2:269–348Google Scholar
  44. Lacaze-Duthiers H (1897) Faune du Golfe du Lion. Coralliaires, Zooanthaires, Sclérodermés. Arch Zool Exp Gén 5:1–249Google Scholar
  45. Lam KKY (2000) Sexual reproduction of a low-temperature tolerant coral Oulastrea crispata (Scleractinia, Faviidae) in Hong Kong, China. Mar Ecol Prog Ser 205:101–111CrossRefGoogle Scholar
  46. Loya Y (1976) The red sea coral Stylophora pistillata is an r strategist. Nature 259:478–480CrossRefGoogle Scholar
  47. Maynard-Smith J (1978) The evolution of sex. Cambridge University Press, CambridgeGoogle Scholar
  48. Mione T, Anderson GJ (1992) Pollen–ovule ratios and breeding system evolution in Solanum section Basarthrum (Solanaceae). Am J Bot 79:279–287CrossRefGoogle Scholar
  49. Moran NA (1992) The evolutionary maintenance of alternative phenotypes. Am Nat 139:971–989CrossRefGoogle Scholar
  50. Neves EG, Pires DO (2002) Sexual reproduction of Brazilian coral Mussimilia hispida (Verril, 1902). Coral Reefs 21:161–168Google Scholar
  51. Oliver JK,Babcock RC, Harrison PL, Willis BL (1988) Geographic extent of mass coral spawning: clues to ultimate causal factors. Proceedings of the 6th international coral reef symposium, Townsville, 2:803–810Google Scholar
  52. Penland L, Kloulechad J, Idip D, van Woesik R (2004) Coral spawning in the western Pacific Ocean is related to solar insolation: evidence of multiple spawning events in Palau. Coral Reefs 23:133–140CrossRefGoogle Scholar
  53. Pianka ER (1970) On r- and K- selection. Am Nat 104:592–597CrossRefGoogle Scholar
  54. Richmond RH (1997) Reproduction and recruitment in corals: critical links in the persistence of reefs. In: Birkeland C (ed) Life and death of coral reefs. Chapman and Hall Inc., New York, pp 175–197CrossRefGoogle Scholar
  55. Richmond RH, Hunter CL (1990) Reproduction and recruitment of corals: comparisons among the Caribbean, the tropical Pacific, and the Red Sea. Mar Ecol Prog Ser 60:185–203CrossRefGoogle Scholar
  56. Rinkewich B, Loya Y (1979) The reproduction of the Red Sea coral Stylophora pistillata. I. Gonads and Planulae. Mar Ecol Prog Ser 1:133–144CrossRefGoogle Scholar
  57. Rodriguez J (1983) Estudio de una communidada planctonica neritica en el Mar de Alboran: II Ciclo del zooplankton. Bol Inst Esp Oceanogr 1:19–44Google Scholar
  58. Rossi L (1975) Sexual races in Cereus pedunculatus (Boad). Publ Staz Zool Napoli 39:462–470Google Scholar
  59. Shaw PW (1989) Seasonal patterns and possible long-term effectiveness of sexual reproduction in three species of sagartiid sea anemones. In: Ryland JS, Tyler PA (eds) Reproduction, genetics and distributions of marine organisms. Olsen and Olsen, Fredensborg, pp 189–199Google Scholar
  60. Siokou-Frangou I (1996) Zooplankton annual cycle in a Mediterranean coastal area. J Plankton Res 18:203–223CrossRefGoogle Scholar
  61. Soong K (1991) Sexual reproductive patterns of shallow-water reef corals in Panama. Bull Mar Sci 49:832–846Google Scholar
  62. Stearns SC (1992) The evolution of life histories. Oxford University Press, OxfordGoogle Scholar
  63. Steiner SCC (1995) Spawning in scleractinian corals from SW Puerto Rico (West Indies). Bull Mar Sci 56:899–902Google Scholar
  64. Stoddart JA, Black R (1985) Cycles of gametogenesis and planulation in the coral Pocillopora damicornis. Mar Ecol Prog Ser 23:153–164CrossRefGoogle Scholar
  65. Szmant AM (1986) Reproductive ecology of Caribbean reef corals. Coral Reefs 5:43–54CrossRefGoogle Scholar
  66. Szmant-Froelich A, Yevich P, Pilson MEQ (1980) Gametogenesis and early development of the temperate coral Astrangia danae (Anthozoa: Scleractinia). Biol Bull 158:257–269CrossRefGoogle Scholar
  67. Tanner JE (1996) Seasonality and lunar periodicity in the reproduction of pocilloporid corals. Coral Reefs 15:59–66CrossRefGoogle Scholar
  68. Tomascik T, Sander F (1987) Effects of eutrophication on reef-building corals. III. Reproduction of the reef-building coral Porites porites. Mar Biol 94:77–94CrossRefGoogle Scholar
  69. Tranter PRG, Nicholson DN, Kinchington D (1982) A description of the spawning and post-gastrula development of the cool temperate coral Caryophyllia smithii (Stokes and Broderip). J Mar Biol Assoc UK 62:845–854CrossRefGoogle Scholar
  70. Van Moorsel GWNM (1983) Reproductive strategies in two closely related stony corals (Agaricia, Scleractinia). Mar Ecol Prog Ser 13:273–283CrossRefGoogle Scholar
  71. Veron JEN (ed) (2000) Corals of the world. Australian Institute of Marine Science, TownsvilleGoogle Scholar
  72. Ward S (1992) Evidence for broadcast spawning as well as brooding in the scleractinian coral Pocillopora damicornis. Mar Biol 112:641–646CrossRefGoogle Scholar
  73. Willis BL, Babcock RC, Harrison PL, Oliver JK, Wallace CC (1985) Patterns in the mass spawning of corals on the great barrier reef from 1981 to 1984. Proc 5th Int Coral Reef Symp Tahiti 4:343–348Google Scholar
  74. Zibrowius H (1980) Les scléractiniaires de la Méditerranée et de l’Atlantique nord-oriental. Mem Inst Oceanogr (Monaco) 11:1–284Google Scholar

Copyright information

© Springer-Verlag 2005

Authors and Affiliations

  • S. Goffredo
    • 1
    Email author
  • V. Airi
    • 1
  • J. Radetić
    • 1
  • F. Zaccanti
    • 1
  1. 1.Department of Evolutionary and Experimental BiologyAlma Mater Studiorum,University of BolognaBolognaItaly

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