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G. Di Maida et al. / Marine Environmental Research 87-88 (2013) 96e102 97
rhizomes growing on matte and sand was greater than that on rocky affecting meadows features and their health status e can also have
substratum. The opposite response was observed at meadow level, interesting policy and management implications.
with higher shoots density on rock than on sand and matte. Finally,
at the community level, epiphyte cover and composition did not 2. Materials and methods
differ among shoots growing on the three substrata.
2.1. Study area, sampling and laboratory analyses
Less attention has been paid instead to the effect of different
substratum typologies on growth performance. The vertical elon- The study was carried out in four locations were P. oceanica
gation rate of the rhizome is closely linked to sedimentation rate as meadows grow on three types of substratum: sand, rock and matte
shoots adapt their growth to different sediment deposition rates (Fig. 1). For substratum differentiation we choose plants growing on
(Boudouresque and Jeudy de Grissac, 1983; Boudouresque et al., sand, plants settled on visibly exposed rock and plants growing on
1984; Jeudy de Grissac and Boudouresque, 1985), through vertical visibly exposed and not buried matte. At each location sampling
growth plasticity (Manzanera et al., 2011; Marbà and Duarte, 1994). and measurements were carried out on meadows settled on each
Marbà and Duarte (1997) observed that where horizontal rhizomes kind of substratum. Shoot density was determined by using five
were buried deeper than 20 cm below the sediment surface the randomly placed quadrats (40 Â 40 cm), and fifteen short shoots
vertical growth rates were higher than those recorded over sub- (Hemminga and Duarte, 2000) connected to the horizontal axis
strates dominated by exposed P. oceanica matte. Furthermore, to were randomly collected for biometric analysis. Depth at each
counteract sediment deposition rate, in sheltered areas and on location was more or less constant, varying between 8 and 10 m. All
carbonate sediments rhizomes tend to grow vertically at a greater samplings and measurements were carried out in July 2008, except
rate, than those growing in exposed areas and on siliciclastic sed- for the Favignana location, which was examined in October 2008.
iments, which tend instead to grow with a lower rate and to expand
laterally because of the absence of sediment deposition (De Falco Rhizomes were analyzed by lepidochronology (Pergent, 1990),
et al., 2008). which provides reliable estimation of seagrass growth performance
(annual vertical growth and leaf production) (Pergent-Martini
Overall, these studies elucidated different aspects of P. oceanica et al., 2005). This retrodating technique allows us to isolate and
attributes developing on different substratum types, although to date rhizome segments corresponding to a 1-year period (lep-
date very few observations (if any) were simultaneously recorded on idochronological year). Each lepidochronological year is dated
meadows growing on the three prevalent types of substratum (i.e. starting from the rhizome apex (sampling year) downward and
rock, sand, and matte) in the Mediterranean Sea. Here, we aim at backdating the sequence of cycles with their corresponding
filling these gaps and at assessing the magnitude and variability of rhizome segment. This reiterative procedure is performed until the
the effect of substratum typologies on P. oceanica growth and rhizome segment connected to the horizontal axis is reached,
morphology attributes. More specifically, growth performance, leaf representing the year of shoot birth. Finally, the length of each
biometry and shoot density are examined and compared in four rhizome segment is measured and the shoot age determined so it
P. oceanica meadows growing on these prevalent substrata along the can be taken into account in the statistical analysis as a potential
north-western coasts of Sicily (Italy). We focused on these P. oceanica confounder (Tomasello et al., 2007).
attributes as these are now commonly used as seagrass quality ele-
ments to assess the ecological status of coastal water bodies under For each rhizome only the three most recent lepidochrono-
the European Union Water Framework Directive (WFD, 2000/60/EC; logical years were considered in the analysis since the number of
Lopez y Royo et al., 2010). Since substratum typologies are not preceding years per shoot varied considerably among the three
presently considered as a source of variation on seagrass character- substrata.
istics in this context, our findings e in addition to have interesting
methodological consequences on the causative variables potentially On leaf bundles biometric analyses according to Giraud (1977)
and Pergent-Martini et al. (2005) were carried out. These
Fig. 1. The four locations examined along the north-western coast of Sicily.