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Pardi, Piazzi, Balata, Papi, Cinelli & Benedetti-Cecchi                                                          Posidonia oceanica epiphytes of Sicily

Table 2. Analysis of variance on mean percentage cover of epiphytes of Posidonia oceanica in Sicily.

                            encrusting red  encrusting brown filamentous    encrusting                     erect
                            algae                                          bryozoans                      bryozoans
                                            algae               algae                                                hydroids      Foraminifera

source of variation d.f. MS        F        MS F                MS F       MS F                           MS F       MS F          MS F

geographic area (¼A)        2 341,948 4.2 131,826 5.4* 772.9 22.6*** 23,973 4.9* 65.8 0.8                            66.5 3.2 7614 8.8**

habitat (¼H)                1 107,398 1.3   64,057 2.6          0.9 0.0           8792 1.8 68.2 0.8                  69.9 3.4 3242 3.8

A·H                         2 130,715 1.6   15,969 0.7          50.5 0.3 12,977 2.7 65.2 0.8 119.2 5.8* 1692 2.0

meadow (A · H) (¼M) 12 80,658 27.3*** 24,330 15.9*** 34.2 6.0*** 4857 17.2*** 86.4 22.3*** 20.6 6.4*** 865 14.6***

site (M) (¼S)         72 2957 2.7*** 1533 2.7*** 5.7 2.8*** 282 2.3*** 3.9 2.6*** 3.2 2.5*** 59 2.1***

plot (S) (¼P)         370 1088 5.5*** 563 2.2*** 2.0 2.8*** 120 2.0*** 1.5 2.4*** 1.3 2.4*** 28 2.3***

residuals             4050 200                           252 0.7                  59 0.6                             0.5 12

total 4499

Cochrane’s test (C)         0.011           0.014               0.011      0.019                          0.010      0.015         0.020

transformation              None            None                ln(x + 1)  None                           ln(x + 1)  ln(x + 1)     None

SNK test                                    W>N¼S               W>N¼S N¼W¼S                                          M: N ¼W¼ S W > N ¼ S

                                                                                                                     I: S > N ¼ W

                                                                                                                     N: I ¼ M

                                                                                                                     W: I ¼ M

                                                                                                                     S: I > M

*P < 0.05; **P < 0.01; ***P < 0.001.
N ¼ north; W ¼ west; S ¼ south; M ¼ Mainland; I ¼ Island.

Table 3. Variance components analyses conducted on untransformed data and expressed as percentage.

variance             encrusting    encrusting            filamentous        encrusting                     erect      hydroids      Foraminifera
components           red algae     brown algae           algae             bryozoans                      bryozoans
                                                                                                                      7.57         18.08
meadow               48.81         23.16                  6.56             21.06                          19.77       5.36          3.39
site                  5.87          4.93                  6.27              3.68                           4.22      12.62          9.04
plot                                7.90                 20.75              7.02                                     74.45
leaf                 13.95                               66.40                                            12.67                    69.49
                     31.36         64.01                                   68.24                          63.34

scale of observation has important implications, from the              islands and continental coasts. In the present paper, we
identification of the mechanisms that generate and main-                compared leaf epiphytes of Posidonia oceanica meadows
tain biological diversity (Underwood & Chapman 1996;                   between the two habitats represented by islands and
Willis & Whittaker 2002), to the prediction of how local               locations on the mainland coasts in different geograph-
and regional environmental changes will affect diversity at            ical areas around Sicily. This was performed by means
multiple levels of organization (Lockwood & McKinney                   of a multifactorial sampling design that enabled the
2001; Scott et al. 2002), to the analysis of relationships             quantitative assessment of variability at a hierarchy of
between species richness and ecological function (Waide                spatial scales, ranging from 10s of centimetres to 100s
et al. 1999). The study of spatial patterns of biodiversity            of kilometres. Significant differences were detected
in benthic organisms has received considerable attention,              among geographic areas for encrusting brown algae, fil-
and a synthesis of these studies suggests that small-scale             amentous algae, encrusting bryozoans and Foraminifera.
processes are at least as important as large-scale processes           A significant geographic area · habitat interaction was
in many assemblages (Fraschetti et al. 2005). Our results              detected only for hydroids, while spatial variability
show that this also applies to ESL, a system common to                 within each habitat was large and significant, partic-
many marine coastal areas.                                             ularly among shoots 10s of centimetres apart and
                                                                       meadows a few kilometres apart.
Conclusions
                                                                       Acknowledgements
Although a number of studies have investigated epi-
phytes on the leaves of seagrasses at different spatial                We dedicate the paper to Lucia Mazzella for her contri-
and/or temporal scales, few have focused on geograph-                  bution to the knowledge of Posidonia oceanica ecosystem,
ical patterns and on differences between the small                     and in particular to epiphytes. This research was supported

Marine Ecology 27 (2006) 397–403 ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd                                  401
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