OIKOS 91: 233 – 244. Copenhagen 2000
Eelgrass, Zostera marina, growth along depth gradients: upper
boundaries of the variation as a powerful predictive tool
Dorte Krause-Jensen, Anne Lise Middelboe, Kaj Sand-Jensen and Peter Bondo Christensen
Krause-Jensen, D., Middelboe, A. L., Sand-Jensen, K. and Christensen, P. B. 2000.
Eelgrass, Zostera marina, growth along depth gradients: upper boundaries of the
variation as a powerful predictive tool. – Oikos 91: 233–244.
1200 measurements of eelgrass (Zostera marina ) biomass, shoot density and cover
along 19 depth gradients in Øresund, located between Denmark and Sweden, were
analysed to characterise growth of eelgrass in relation to depth. The large data set
allowed analyses of boundaries of distribution as well as of average trends. Natural
variability is large in shallow water where populations are disturbed by wave action
and other physical parameters. Models based on average values, therefore, did not
adequately describe growth regulation by resources, and only explained a minor part
(up to 30%) of the overall variation in data. In contrast, boundary functions, which
describe the upper bounds of distributions, focus on the variation produced by the
ultimately growth-regulating resource, and therefore provide models with high pre-
dictive power. An exponential model explained up to 90% of the variation in upper
bounds of eelgrass shoot density as a function of depth and indicated that shoot
density was ultimately regulated by light availability. The boundary functions demon-
strated that eelgrass shoot density, biomass and cover followed markedly different
patterns as functions of depth and were affected differently by the factors governing
their distribution. In addition, boundary functions revealed informative spatial
structures in data and illustrated whether a given general trend was caused by
changes in maximum values, minimum values or both. For example, upper and lower
boundaries of biomass-shoot density relations changed markedly with depth, demon-
strating depth-related changes in intraspecific succession and competition patterns.
Boundary functions are therefore suggested as a promising tool for analysing
ultimate regulating factors of distribution and growth of organisms when large data
sets are available.
D. Krause -Jensen, A. L. Middelboe, and P. B. Christensen, Dept of Lake and Estuarine
Ecology, National Enironmental Research Inst., Vejlsøej 25, DK-8600 Silkeborg,
Denmark (dkj@dmu.dk).– K. Sand -Jensen (and present address of ALM), Freshwater
Biological Laboratory, Uni. of Copenhagen, Helsingørsgade 51, DK-3400 Hillerød,
Denmark.
Seagrasses are important structural components and
primary producers of shallow, soft-bottom coastal wa-
ters throughout the world (Den Hartog 1970, Mann
1982). The lower depth limit of seagrass meadows
delimits the coastal area potentially covered by sea-
grasses, and several studies have documented that vari-
ations in seagrass depth limits among species and sites
are mainly regulated by light attenuation in the water
column (e.g. Ostenfeld 1908, Backman and Barilotti
1976, Duarte 1991, Sand-Jensen et al. 1994). In contrast
to seagrass depth limits, seagrass abundance along
depth gradients has not been intensively analysed. Pre-
vious studies indicate that seagrasses display either a
bell-shaped distribution along depth gradients, with
maximum abundance at intermediate depths and lower
abundance in shallow and deep water (Bay 1984, Sand-
Jensen et al. 1997), or a general decline in seagrass
abundance with increasing depth (Buesa 1975, Hulings
1979, West 1990). Exposure, desiccation and ice-scour
may reduce seagrass abundance in shallow water
Accepted 10 May 2000
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ISSN 0030-1299
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OIKOS 91:2 (2000) 233