Marine Biology

, Volume 153, Issue 4, pp 589–598 | Cite as

Animals on marine flowers: does the presence of flowering shoots affect mobile epifaunal assemblage in an eelgrass meadow?

  • Masahiro NakaokaEmail author
  • Masatoshi Matsumasa
  • Tetsuhiko Toyohara
  • Susan L. Williams
Research Article


Eelgrass, Zostera marina, produces two types of shoots: morphologically simple vegetative shoots and highly branched flowering (reproductive) shoots, the latter found only in summer months. We examined whether the abundance and diversity of mobile epifaunal assemblage are affected by the presence of flowering shoots in an eelgrass meadow of Otsuchi Bay, northeastern Japan. Comparisons of epifauna in natural vegetation revealed that density and species richness did not differ significantly between sites consisting of both flowering and vegetative shoots, and those only of vegetative shoots. A transplant experiment, conducted to examine the colonization rates of epifauna to defaunated eelgrass planted with different combination of vegetative and flowering shoots, showed no obvious variation in abundance and species richness. At species level, the density of some species such as a tanaid Zeuxo sp. and a polychaete Platynereis sp. was higher at sites and/or treatments with flowering shoots, whereas that of some gastropods, such as Lirularia iridescens and Siphonacmea oblongata was higher at sites without flowering shoots. The species-specific response led to dissimilarity of epifaunal assemblage between sites and among treatments with different densities of vegetative and flowering shoots. Similar patterns observed for natural vegetation and the transplant experiment suggest that the variation in assemblage structure is caused by habitat selection of each species, for example, the utilization of flowering shoots as feeding ground and nursery by Zeuxo sp.


Macrophyte Assemblage Structure Transplant Experiment Shoot Density Vegetative Shoot 
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 wish to thank the staff of the Otsuchi Marine Research Center, Ocean Research Institute, University of Tokyo, for facilitation of our research. We are specially indebted to B. Nyden, K. Morita, K. Hirano, K. Sado, M. Kurosawa and Y. Iwama for their help in the fieldwork. We are grateful to Y. Hashiguchi and Y. Okuzaki for laboratory assistance, and to M. Shimanaga for taxonomic identification of copepods. The research is partially supported by grants from the Ministry of Science, Education and Culture, Japan to M. N. (nos. 08740954 and 08680594), and by the research fellowship program by Japan Society for the Promotion of Science to S. L. W. (no. S-97122).


  1. Aioi K (1980) Seasonal change in the standing crop of eelgrass (Zostera marina L.) in Odawa Bay, central Japan. Aquat Bot 8:343–354CrossRefGoogle Scholar
  2. Aioi K, Nakaoka M (2003) Seagrasses of Japan. In: Green EP, Short FT (eds) World atlas of seagrasses. University of California Press, Berkeley, pp 185–192Google Scholar
  3. Almany GR (2004) Does increased habitat complexity reduce predation and competition in coral reef fish assemblages? Oikos 106:275–284CrossRefGoogle Scholar
  4. Attrill MJ, Strong JA, Rowden AA (2000) Are macroinvertebrate communities influenced by seagrass structural complexity? Ecography 23:114–121CrossRefGoogle Scholar
  5. Bell SS, McCoy ED, Mushinsky HR (1991) Habitat structure: the physical arrangement of objects in space. Chapman & Hall, LondonCrossRefGoogle Scholar
  6. Bell SS, Fonseca MS, Stafford NB (2006) Seagrass ecology: new contributions from a landscape perspective. In: Larkum AWD, Orth RJ, Duarte CM (eds) Seagrasses: biology, ecology and conservation. Springer, Dordrecht, pp 625–645Google Scholar
  7. Bologna PAX, Heck KL (1999) Macrofaunal associations with seagrass epiphytes—relative importance of trophic and structural characteristics. J Exp Mar Biol Ecol 242:21–23CrossRefGoogle Scholar
  8. Chemello R, Milazzo M (2002) Effect of algal architecture on associated fauna: some evidence from phytal molluscs. Mar Biol 140:981–990CrossRefGoogle Scholar
  9. Clarke KR, Warwick RM (2001) Change in marine communities: an approach to statistical analysis and interpretation, 2nd edn. PRIMER-E, PlymouthGoogle Scholar
  10. Day RW, Quinn GP (1989) Comparisons of treatments after an analysis of variance in ecology. Ecol Monogr 59:433–463CrossRefGoogle Scholar
  11. Dean RL, Connell JH (1987a) Marine invertebrates in an algal succession. II. Tests of hypotheses to explain changes in diversity with succession. J Exp Mar Biol Ecol 109:217–247CrossRefGoogle Scholar
  12. Dean RL, Connell JH (1987b) Marine invertebrates in an algal succession. III. Mechanisms linking habitat complexity with diversity. J Exp Mar Biol Ecol 109:249–273CrossRefGoogle Scholar
  13. De Troch M, Vandepitte L, Raes M, Suàrez-Morales E, Vincx M (2005) A field colonization experiment with meiofauna and seagrass mimics: effect of time, distance and leaf surface area. Mar Biol 148:73–86CrossRefGoogle Scholar
  14. Den Hartog C (1970) The sea grass of the world. North Holland Publishing Company, AmsterdamGoogle Scholar
  15. Duffy JE (2006) Biodiversity and the functioning of seagrass ecosystems. Mar Ecol Prog Ser 311:233–250CrossRefGoogle Scholar
  16. Duffy JE, Canuel EA, Richardson JP (2003) Grazer diversity and ecosystem functioning in seagrass beds. Ecol Lett 6:1–9CrossRefGoogle Scholar
  17. Duffy JE, Richardson JP, France KE (2005) Ecosystem consequences of diversity depend on food chain length in estuarine vegetation. Ecol Lett 8:301–309CrossRefGoogle Scholar
  18. Edgar GJ (1990) The influence of plant structure on the species richness, biomass and secondary production of macrofauna assemblages associated with Western Australian seagrass beds. J Exp Mar Biol Ecol 137:215–240CrossRefGoogle Scholar
  19. Edgar GJ, Robertson AI (1992) The influence of seagrass structure on the distribution and abundance of mobile epifauna: pattern and process in a Western Australian Amphibolis bed. J Exp Mar Biol Ecol 160:13–31CrossRefGoogle Scholar
  20. Grabowski JH (2004) Habitat complexity disrupts predator–prey interactions but not the trophic cascade on oyster reefs. Ecology 85:995–1004CrossRefGoogle Scholar
  21. Heck KL, Orth RJ (1980) Structural components of eelgrass Zostera meadows in the lower Chesapeake Bay—decapod crustacea. Estuaries 3:289–295CrossRefGoogle Scholar
  22. Heck KL, Orth RJ (2006) Predation in seagrass beds. In: Larkum AWD, Orth RJ, Duarte CM (eds) Seagrasses: biology, ecology and conservation. Springer, Dordrecht, pp 537–550CrossRefGoogle Scholar
  23. Hellwig-Armonies M (1988) Mobile epifauna on Zostera marina, and infauna of its inflorescences. Helgoländer Meeresunters 42:329–337CrossRefGoogle Scholar
  24. Hovel KA, Fonseca MS, Myer DL, Kenworthy WJ, Whitfield PE (2002) Effects of seagrass landscape structure, structural complexity and hydrodynamic regime on macrofaunal densities in North Carolina seagrass beds. Mar Ecol Prog Ser 243:11–24CrossRefGoogle Scholar
  25. Jernakoff P, Brearley A, Nielsen J (1996) Factors affecting grazer–epiphyte interactions in temperate seagrass meadows. Oceanogr Mar Biol Ann Rev 34:109–162Google Scholar
  26. Jenkins G, Walker-Smith G, Hamer P (2002) Elements of habitat complexity that influence harpacticoid copepods associated with seagrass beds in a temperate bay. Oecologia 131:598–605CrossRefGoogle Scholar
  27. Kelaher BP (2005) Does colonization contribute to spatial patterns of common invertebrates in coralline algal turf? Aust Ecol 30:40–48CrossRefGoogle Scholar
  28. Lewis FG (1987) The crustacean epifauna of seagrass and macroalgae in Apalachee Bay, Florida, USA. Mar Biol 94:219–229CrossRefGoogle Scholar
  29. Mukai H, Suzuki T, Nojima S (1999) Morphological implications of seagrass substratum for epifauna community in Rottnest, Western Australia. In: Walker DI, Wells FE (eds) The seagrass flora and fauna of Rottnest Island, Western Australia. Western Australia Museum, Perth, pp 255–274Google Scholar
  30. Nakaoka M (2002) Predation on seeds of seagrasses Zostera marina and Zostera caulescens by a tanaid crustacean Zeuxo sp. Aquat Bot 72:99–106CrossRefGoogle Scholar
  31. Nakaoka M (2005) Plant–animal interactions in seagrass beds: ongoing and future challenges for understanding population and community dynamics. Popul Ecol 47:167–177CrossRefGoogle Scholar
  32. Nakaoka M, Aioi K (2001) Ecology of seagrasses Zostera spp. (Zosteraceae) in Japanese waters: a review. Otsuchi Mar Sci 26:7–22Google Scholar
  33. Nakaoka M, Toyohara T, Matsumasa M (2001) Seasonal and between-substrate variation in mobile epifaunal community in a multispecific seagrass bed of Otsuchi Bay, Japan. PSZN Mar Ecol 22:379–395CrossRefGoogle Scholar
  34. Nakaoka M, Kouchi N, Aioi K (2003) Seasonal dynamics of Zostera caulescens: relative importance of flowering shoots to net production. Aquat Bot 77:277–293CrossRefGoogle Scholar
  35. Nelson WG (1979) An analysis of structural pattern in an eelgrass (Zostera marina L.) amphipod community. J Exp Mar Biol Ecol 39:231–264CrossRefGoogle Scholar
  36. Nelson WG, Bonsdorff E (1990) Fish predation and habitat complexity: are complexity threshold real? J Exp Mar Biol Ecol 141:183–194CrossRefGoogle Scholar
  37. Orth RJ (1992) A perspective on plant–animal interactions in seagrasses: physical and biological determinants influencing plant and animal abundance. In: Price JH, John DM, Hawkins SJ (eds) Plant–animal interactions in the marine benthos. Clarendon Press, Oxford, pp 147–164Google Scholar
  38. Orth RJ, Heck Jr KL, van Montfrans J (1984) Seagrass faunal communities: a review of the influence of plant structure and prey characteristics on predator–prey relationships. Estuaries 7:339–350CrossRefGoogle Scholar
  39. Parker JD, Duffy JE, Orth RJ (2001) Experimental tests of plant diversity effects on epifaunal diversity and production in a temperate seagrass bed. Mar Ecol Prog Ser 224:55–67CrossRefGoogle Scholar
  40. Russo AR (1990) The role of seaweed complexity in structuring Hawaiian epiphytal amphipod communities. Hydrobiologia 194:1–12CrossRefGoogle Scholar
  41. Stoner AW, Lewis FG (1985) The influence of quantitative and qualitative aspects of habitat complexity in tropical seagrass meadows. J Exp Mar Biol Ecol 94:19–40CrossRefGoogle Scholar
  42. Takeuchi I, Hirano R (1995) Clinging behavior of the epifaunal caprellids (amphipoda) inhabiting the Sargassum zone on the Pacific coast of Japan, with its evolutionary implications. J Crustacean Biol 15:481–492CrossRefGoogle Scholar
  43. Tatsukawa K, Tanaka S (1982) Studies on the fish community in coastal waters of Otsuchi Bay. Otsuchi Mar Res Center Rep 8:49–68Google Scholar
  44. Toyohara T, Nakaoka M, Aioi K (1999) Population dynamics and reproductive traits of phytal gastropods in seagrass bed in Otsuchi Bay, northeastern Japan. PSZN Mar Ecol 20:273–289CrossRefGoogle Scholar
  45. Toyohara T, Nakaoka M, Tsuchida E (2001) Population dynamics and life history traits of Siphonacmea oblongata Yokohama on seagrass leaf in Otsuchi Bay (Siphonariidae, Pulamonata). Venus (Jap J Malaco) 60:27–36Google Scholar
  46. Virnstein RW, Howard RK (1987) Motile epifauna of marine macrophytes in the Indian River Lagoon, Florida. 1. Comparisons among three species of seagrasses from adjacent beds. Bull Mar Sci 41:1–12Google Scholar
  47. Worthington DG, Ferrell DJ, McNeill SE, Bell JD (1992) Effects of the shoot density of seagrass on fish and decapods: are correlation evident over larger spatial scales? Mar Biol 112:139–146CrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2007

Authors and Affiliations

  • Masahiro Nakaoka
    • 1
    Email author
  • Masatoshi Matsumasa
    • 2
  • Tetsuhiko Toyohara
    • 3
  • Susan L. Williams
    • 4
  1. 1.Graduate School of ScienceChiba UniversityChibaJapan
  2. 2.Department of Biology, School of Liberal Arts and SciencesIwate Medical UniversityMoriokaJapan
  3. 3.Marine Biological Research Institute of Japan Co., LTDOsakaJapan
  4. 4.Bodega Marine LaboratoryUniversity of California at DavisBodega BayUSA

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