Journal of Experimental Marine Biology and Ecology,

215 (1997) 269288 L

Effects of sedimentation on subtidal macroalgal assemblages:
an experimental study from a mediterranean rocky shore
Laura Airoldi*, Francesco Cinelli
Dipartimento di Scienze dell Uouoe dell Ambiente, Universita` di Pisa, Via A. Volta 6, I-56126 Pisa, Italy
Received 24 April 1996; revised 16 October 1996; accepted 13 November 1996

Abstract

The effects of sedimentation on the development and structure of macroalgal assemblages were
investigated from June 1992 to December 1993 on a rocky subtidal shore South of Livorno
(Ligurian Sea). Experimental procedures were specifically designed to address: (1) which species
recruit in patches of bare rock under different sedimentation rates; (2) whether small-scale
variations of sediment deposition influence the local diversity of the assemblage; (3) whether the
effects of sedimentation are independent from the time of the year when succession begins.
Depositional environment was manipulated in the field by means of transparent Plexiglas panels
which on average reduced the amount of sediment sinking over the algal assemblage by 35%.
Possible effects on the penetration of light and flow microenvironment were assessed. Small-scale
variations of sedimentation rates significantly influenced the local structure and diversity of the
algal assemblage. Such effects were particularly evident during the recolonisation of patches of
bare rock and were dependent upon the time at which succession was initiated. The erect algae
were the ones more affected by depositional environment and showed different responses to the
experimental reduction of sediment supplies. All the plots were quantitatively dominated by a
filamentous turf mostly consisting of Polysiphonia setacea, whose growth was potentially
enhanced by reduced sedimentation rates. At the end of the experiments, the overall diversity and
evenness of the assemblage were lower in treatments with lessened sediment supplies. It is
suggested that small-scale spatial variability of the depositional environment may affect the
within-habitat diversity of algal assemblages either through direct effects on individual species or
on their propagules, and through indirect effects mediated by competitive outcomes. 1997
Elsevier Science B.V.

Keywords: Sedimentation; Diversity; Recruitment; Macroalgae; Turf; Subtidal; Rocky shore

*Corresponding author. Tel.: (139-50) 500943; fax: (139-50) 49694; e-mail: lairoldi@discat.unipi.it

0022-0981 / 97 / $17.00 1997 Elsevier Science B.V. All rights reserved

PII S0022-0981( 96 )02770-0
270 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

1. Introduction

The worldwide increase of water turbidity and sediment load in coastal areas as a
consequence of anthropogenic activities has stirred interest into the impact of sedi-
mentation upon rocky shore organisms. Such interest is reflected by an extensive
literature (Moore, 1977; Seapy and Littler, 1982; Taylor, 1983; Rogers, 1990; Engledow
and Bolton, 1994; Saiz Salinas and Isasi Urdangarin, 1994; Shaffer and Parks, 1994;
Airoldi et al., 1995). However, so far, few quantitative experiments have examined in
the field the effects of this important physical process on the composition and structure
of macroalgal communities (Neushul et al., 1976; Kendrick, 1991).
Sedimentation is believed to influence the distribution and diversity of seaweeds
(Devinny and Volse, 1978; Robles, 1982; Stewart, 1983; DAntonio, 1986; Shiel and
Foster, 1986; Santos, 1993). However, most studies have been limited to descriptions of
species typically associated with sediment influenced habitats (Lewis, 1964; Mathieson,
1967; Markham and Newroth, 1972; Mathieson and Fralick, 1972; Markham, 1973;
Daly and Mathieson, 1977), and the causal mechanisms that shape the local structure of
algal assemblages are poorly understood. The prevalent opinion is that high rates of
sediment deposition and movement are detrimental to the overall richness and diversity
of the community through exclusion of less tolerant species (Daly and Mathieson, 1977;
Little and Smith, 1980; Engledow and Bolton, 1994; Saiz Salinas and Isasi Urdangarin,
1994). On the other hand sediment disturbance may promote diversity through
preventing monopolisation of space by competitively dominant species and through
maintaining patchiness (Foster, 1975; Taylor and Littler, 1982; Littler et al., 1983;
McQuaid and Dower, 1990).
The purpose of the present study was to investigate the effects of sedimentation on the
development and structure of a macroalgal assemblage growing on a rocky subtidal
shore (south of Livorno, Western Mediterranean Sea). This tract of coast is characterized
by high sedimentation rates (Airoldi et al., 1996), and local patterns of sediment
deposition are variable on a scale of meters (Airoldi, unpubl. data), potentially
influencing the within-habitat diversity of the algal assemblage. Experimental pro-
cedures, including manipulations of the depositional environment, were specifically
designed to address (1) which species recruit on patches of bare rock under different
rates of sediment deposition, (2) whether small-scale variations of depositional environ-
ment influence the local diversity of the assemblage and (3) whether the effects of
sedimentation are independent from the time of the year when succession begins. The
quantitative characterisation and the temporal variations of the depositional environment
of the study area and possible influences on patterns of abundance of the algal species
are reported elsewhere (Airoldi et al., 1995, 1996).

2. Methods

2.1. Study site

The experiments were carried out from June 1992 to December 1993 on an exposed
 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288 271

rocky shore located about 10 km south of Livorno (Ligurian Sea). A detailed description
of the characteristics of the area can be found in Airoldi et al. (1995, 1996). The
experimental site was a gently sloping ( | 308, 1317 m in depth) field of sandstone
boulders colonised by an algal assemblage strongly dominated by a dense turf mostly
consisting of Polysiphonia setacea Hollenberg (Airoldi et al., 1995). This species, which
occurs on rocks and as epiphyte on other organisms, develops dense mats of upright
filaments up to 1 cm high from prostrate branches firmly attached to the substratum by
rhizoids (Verlaque, 1989). The calcareous alga Halimeda tuna (Ellis et Solander)
Lamouroux represented the only abundant erect species. The abundance of grazers was
low, and these were mainly represented by specimens of the herbivorous fish Sarpa
salpa (Linnaeus).
A total of 48 boulders, selected at random among 80 that had an horizontal top surface
larger than 35 cm 3 40 cm, were used in the experiments described below. The
minimum distance between boulders was 1.5 m.

2.2. Manipulation of depositional environment

In the following experiments sediment deposition over the algal assemblage was
reduced in treatment plots by means of 34 cm 3 39 cm transparent Plexiglas panels.
These were permanently anchored onto the top surface of the boulders, at a height of
approximately 15 cm from the algal assemblage, by means of four stainless steel bars
(diameter 4 mm) cemented with epoxy putty (Subcoat S, Veneziani) into holes produced
in the rock by hammer and chisel (Fig. 1). The panels were fixed at the upper threaded
extremities of the bars by means of nuts, in order to allow their removal and deployment
by divers during monitoring of the plots.
The effectiveness of the panels in reducing the amount of sediment sinking over the
algal assemblage was assessed from February to July 1993 by comparing sedimentation
rates over six treatment and six control plots established at random in the area. Small
cylindrical sediment traps (diameter 20 mm and aspect: ratio 2) were deployed vertically
stable on the horizontal top surface of the boulders by means of lead weights during 11
periods ranging from 5 to 28 days. Every time, however, some traps were lost, probably
both as a consequence of severe storms and vandals. Only eight periods, when at least
four traps could be retrieved from both treatment and control plots, were included in the
analysis. Traps were closed with plastic stoppers before retrieval. The sediment collected
in each trap was filtered through ignited and preweighed 25 mm Whatman GF / F glass
microfiber filters, rinsed with distilled water to remove salts and dried at 608C for 24 h
before weighing. The data were analysed using a two-way mixed model ANOVA with
panel (treatment vs. control) as fixed factor and time (eight levels) as random factor. The
number of replicates for each combination of panel and time was balanced by choosing
at random a constant smaller number of four traps from those retrieved (Underwood,
1981). In this and in the following experiments, before running the analysis the
assumption of homogeneity of variances was examined using Cochrans C or Hartleys
Fmax tests (Winer, 1971). Opportune transformations were applied when necessary
(Sokal and Rohlf, 1981). StudentNewmanKeuls tests were used for a posteriori
multiple comparisons of means.
272 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

Fig. 1. Panels used in treatment plots to reduce the amount of sediment sinking over the algal community on a
rocky subtidal shore South of Livorno (see text for further explanations).

To avoid shading of the algal assemblage as a consequence of the establishment of a

thin film of sediment and fouling over the transparent panels, these were carefully
cleaned in situ at least once a week throughout the experiments, except on six occasions
when bad weather conditions made diving unsafe for 14, 14, 17, 18, 20 and 22 days
respectively. The effectiveness of such cleaning was tested by comparing the irradiance
below the panels and in the control plots after a short ( , 10) or a long ( . 10) interval
of days had elapsed between two successive cleaning. Measurements were carried out by
means of a quantometer (Biospherical Instrument Irradiancemeter) and were repeated
throughout the research four times for each interval of days. Every time irradiance was
measured in six treatment and six control plots selected at random among those used for
the experiments described below. Data were analysed using a three-way mixed model
analysis of variance with panel (treatment vs. control) and interval ( , 10 days vs. . 10
days) as fixed factors and time (eight levels) as random variable nested in interval.
An additional set of measurements was taken in order to assess the flow mi-
croenvironment near the rockyseawater interface. The rate of dissolution of initially
dried (608C, 24 h) and weighed gypsum balls (diameter 2.3 mm, average weight
including steel bars and plastic labels 11 g) was measured on six occasions from July
1992 to June 1993 in the same six treatment and six control plots used to assess the
effectiveness of the panels in reducing rates of sediment deposition. Gypsum balls were
prepared following Gambi et al. (1989) and mounted on lead weights. Lead weights with
two to four balls were positioned in the field for periods variable from 24 to 48 h.
 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288 273

Additional balls closed in a transparent, non hermetic box were used each time to
calibrate the measurements (Bailey-Brock, 1979). After recovery, gypsum balls were
rinsed lightly in freshwater to remove salts, dried at 608C for 24 h and weighed. The
dissolution (weight loss) of each ball was converted to an estimate of flux (cm / sec)
according to the following function obtained by Bailey-Brock (1979)

Flux 5 3.65 (Me /Mc 2 1)

where Me is the weight loss of each gypsum ball during field exposure and Mc is the
weight loss of calibration balls. Fluxes estimated with balls mounted on the same lead
weight were pooled, and the mean was used as the dependent variable. Data were
analysed using a two-way mixed model ANOVA with panel (treatment vs. control) as
fixed factor and time (six levels) as random factor.

2.3. Effects of sedimentation on macroalgae

A first experiment was set up in June 1992 in order to determine the effects of
reduced sedimentation rates on the composition and the structure of the algal assem-
blage. Twelve 13 cm317 cm plots were established on the horizontal top surface of as
many boulders, permanently marked at their edges with epoxy putty and then assigned at
random to treatments (reduced sediment deposition by means of Plexiglas panels) and
controls (unmanipulated plots) with six replicates each. The abundance of algae was
monitored monthly by visual surveys and photographs taken with a 1:7 extension tube
on a Nikonos III underwater camera equipped with an electronic flash and Fujichrome
100 Asa films. In the laboratory the transparencies were projected onto a grid of 100
equally spaced dots, and the percent cover values of each species or group of species
was expressed as the number of hits after correction for those points which laid over
unidentifiable substratum. Species present in quadrats but not abundant enough to be
scored from intercepts with the grid were assigned an arbitrary cover value of 0.1%. The
photographic sampling did not allow quantification of the abundance of those specimens
covered by the algal turf. A major characterisation of the composition and the structure
of the assemblage was carried out at the end of the experiment, in June 1993, by
destructively sampling the plots with hammer and chisel. The samples were analysed in
the laboratory as described by Airoldi et al. (1995), and the abundance of species was
expressed in terms of biomass (dry weight) per unit area. For each sample the
ShannonWeaver diversity index (H9) and the species evenness (J) were calculated. The
data from the photographic sampling were analysed using Students t-tests on the
percent cover values of each species in each plot averaged over the sampling dates. The
data obtained from the destructive sampling (number of species, biomass, H9, J) were
analysed using Students t-tests.
A second 232 factorial experiment was conducted to examine the effects of
sedimentation on patterns of development of the algal assemblage following complete
removal of the biota at different times of the year. In June 1992, 12 experimental plots
(16 cm320 cm) were scraped clean of resident organisms by means of hammer and
chisel and then assigned at random to treatments (reduced sediment deposition by means
274 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

of Plexiglas panels) and controls (unmanipulated plots) with six replicates each. The
great anfractuosity of the substratum made cleaning operation rather difficult, and
prostrate axes of those specimens living in the deepest crevices were probably not
completely removed. A second series of cleared areas was produced in December 1992.
The time at which succession was initiated was not replicated within season: any
difference among the plots produced in different times may, therefore, reflect chance
temporal variation (in either patterns of recruitment and environmental factors) rather
than predictable seasonal trends. Patterns of colonisation of cleared areas exposed to
different sedimentation rates were sampled monthly for 1 yr by photographs and visual
observations as described above. The abundance of algae was estimated over a smaller
surface (13 cm317 cm) in order to avoid edge effects. The reproductive phenology of
the dominant alga Polysiphonia setacea was also studied throughout the research by
biweekly to monthly samples, in order to assess its reproductive status. Percent cover
data were analysed using a two-way ANOVA on the average values of each species over
time; both sedimentation (reduced vs. natural) and time of beginning (June vs.
December) were fixed factors. The plots produced in June were also sampled destruc-
tively after 1 yr, and the data analysed as described in the above experiment.

3. Results

3.1. Effectiveness of panels

On average, the panels used in the experiments significantly reduced the amount of
sediment sinking on the algal assemblage by about 35% (Fig. 2, Table 1). Although
sedimentation rates varied significantly among sampling periods, the differences
observed between treatment and control plots generally persisted over time, as indicated
by the lack of significant panel3time interaction (Table 1).
The weekly cleaning of the transparent panels prevented the establishment of a film of
sediment and fouling and the consequent shading of the organisms below. In fact, when
10 days at maximum had elapsed between cleaning, the irradiance measured below the
panels in treatment plots was not significantly different from that measured in the
controls (Fig. 3 Table 2). On the contrary, as expected, when bad weather conditions
prevented cleaning procedures for periods longer than 10 days, the irradiance below the
panels significantly decreased down to 37% of that measured in the control plots. The
magnitude of such differences, however, sharply lessened when environmental ir-
radiance was low (Fig. 3 Table 2). Moreover, it may be reasonably presumed that the
differences between treatment and control plots arose gradually, becoming relevant only
at the end of the intervals.
The presence of the panels over the algal assemblage did not affect the flow
microenvironment at the rockyseawater interface. The rates of dissolution of the
gypsum balls were, in fact, similar in treatment and control plots during all the
samplings (Fig. 4 Table 3).
 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288 275

Fig. 2. Effectiveness of panels in reducing the sedimentation rate on the algal assemblage. Sedimentation rates
(mg m 22 d 21 ) over the experimental plots (treatment panels vs controls) were measured by means of sediment
traps during eight periods from February to July 1993. Data are means (11 S.E.) over four replicates.

3.2. Structure of the algal assemblage

3.2.1. Photographic sampling

Throughout the first experiment, both treatment and control plots were quantitatively
dominated by a dense filamentous turf (mainly consisting of Polysiphonia setacea)
whose percent cover values were never lower than 78% (Fig. 5 Table 4). The erect
species were mostly represented by the articulated calcareous alga Halimeda tuna (Fig.
5). This species was generally less abundant in plots with reduced sedimentation rates,
but differences observed were not statistically significant (Table 4). Dictyota dichotoma
(Hudson) Lamouroux and Giffordia sp. were also present in spring and summer months
especially in plots with reduced sedimentation rates (Fig. 5). However, their distribution

Table 1
Analysis of the effectiveness of panels in reducing rates of sediment deposition on the algal assemblage
Source of variation df MS F P Cochrans C test(transformation)
a
Panel (5P) 1 4.296 11.18 0.012 C50.292 P.0.05
Time (5T) 7 3.841 9.775 ,0.001 a (logarithmic)
P3T 7 0.384 0.977 0.458
Error 48 0.393
Sedimentation rates in treatment (with panel) and control plots were compared at different times of the year.
Panel is a fixed factor, while time is random.
a
Significant P values (P,0.05).
276 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

Fig. 3. The irradiance (10 15 quanta cm 22 sec 21 ) below the panels and in the control plots after a short (,10 d)
or a long (.10 d) interval between two successive cleaning. Measurements were repeated four times for each
interval. Data are means (11 S.E.) over six replicates.

was scattered, and the differences observed between treatment and control plots were not
statistically significant (Table 4).

3.2.2. Destructive sampling

After 1 yr, at the end of the first experiment, 22 species showed either in treatment or
control plots mean biomass higher than 0.01 g m 22 (Table 5). Polysiphonia setacea
alone represented, on average, 55% of the total biomass of the assemblage in control
plots and 72% in treatment plots. Conversely, Halimeda tuna was more abundant in
controls than in treatments, where many dead specimens of this alga were found
entrapped within the dense matrix of filaments of P. setacea. Also the biomass of the

Table 2
Analysis of the effectiveness of panel cleaning
Source of variation df MS F P SNK tests
a
Panel (5P) 1 26.658 10.43 0.018 ,10 T 1 Control5Panel
Interval (5I) 1 52.574 0.527 0.494 ,10 T 2 Control5Panel
P3I 1 10.939 4.279 0.084 ,10 T 3 Control5Panel
Time (I) (5T) 6 99.63 248.6 ,0.001 a ,10 T 4 Control5Panel
P3T (I) 6 2.556 6.377 ,0.001 a .10 T 1 Control5Panel
Error 80 0.401 .10 T 2 Control.Panel
.10 T 3 Control.Panel
Cochrans C test C50.138 P.0.05 .10 T 4 Control.Panel
Transformation none
Irradiance in treatment (with panel) and control plots was compared after different intervals of days (,10 vs
.10) had elapsed between two successive cleaning. Measurements were repeated four times for each interval
of days. Panel and interval are fixed factors, while time is random and nested in interval.
a
Significant P values (P,0.05).
 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288 277

Fig. 4. Intensity of flow microenvironment (cm sec 21 ) below the panels and in the control plots estimated by
means of gypsum balls on six occasions from July 1992 to June 1993. Data are means (11 S.E.) over six
replicates (see text for further explanations).

prostrate species, which were commonly covered by the turf, was generally higher in
control plots. Overall, however, the variations of depositional environment had no
significant effects on the composition and structure of the assemblage, although a trend
toward a decrease of species diversity and evenness was observed in treatment plots
(Table 6).

3.3. Patterns of recruitment

3.3.1. Photographic sampling

In the recruitment experiment, encrusting coralline algae were among the first
colonists of the plots produced in both periods. The recruitment of these species
proceeded rapidly, and they reached mean percent covers up to 20% in both treatments
and controls within two months from the beginning of the experiment. After that time,

Table 3
Analysis of the effects of panels on water fluxes near the bottom surface
Source of variation df MS F P Cochrans C test(transformation)
Panel (5P) 1 0.203 6.483 0.052 C50.186 P.0.05
Time (5T) 5 1.895 60.19 ,0.001 a (logarithmic)
P3T 5 0.031 0.996 0.434
Error 36 0.031
The rate of dissolution of gypsum balls in treatment (with panel) and control plots was compared at different
times of the year. In each plot fluxes were estimated with 24 balls, and the mean was used as the dependent
variable. Panel is a fixed factor while time is random.
a
Significant P values (P,0.05) are shown in bold type.
278 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

Fig. 5. Effects of small-scale spatial variation of sediment deposition on patterns of abundance of macroalgae.
Data are mean percent covers (11 S.E.) on six replicates. Treatments, reduced sediment deposition; controls,
natural sediment deposition.

however, they were progressively overgrown by the filamentous turf, and it was no
longer possible to follow their development until the final destructive sampling.
Patterns of recruitment of the filamentous turf were significantly influenced by the
time of beginning of the experiment (Fig. 6 Table 7). The turf rapidly colonised both
treatment and control plots, reaching percent covers above 80% after 8 months in the
plots produced in June and after 6 months in those produced in December, but
restoration of a thick mat took longer (Airoldi, pers. obs.). In both periods cleared space
was filled by lateral vegetative encroachment of prostrate axes from the surrounding turf
and, to a lesser extent, by regrowth of outlived axes not completely removed from the
substratum (see Section 2). In spite of the frequent sampling, P. setacea was never seen
in a reproductive state, suggesting that recruitment from settled propagules did not
contribute to the recovery of this species.

Table 4
Analysis of the effects of sedimentation (reduced vs natural) on patterns of abundance of macroalgae
Algal species t df P Hartleys Fmax test Transformation
Filamentous turf 0.53 10 0.607 Fmax 52.53 P.0.05 none
Halimeda tuna 1.59 10 0.143 Fmax 51.07 P.0.05 none
Dictyota dichotoma 1.69 10 0.122 Fmax 514.7 0.01,P,0.05 none
Giffordia sp. 1.33 10 0.217 Fmax 5` P,0.01 none
Students t-tests were performed on the average percent cover values of each species trough time (see text for
further explanations).
 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288 279

Table 5
Mean biomass (g m 22 ) and standard deviations of the most abundant macroalgae in treatment (reduced
sedimentation rates) and control (natural sedimentation rates) plots at the end of the experiment to determine
the effects of sedimentation on the composition and structure of the assemblage
Species Treatments Controls
M S.D. M S.D.
Prostrate layer
Encrusting corallines 24.19 19.63 28.59 17.99
Peyssonnelia bornetii Boudouresque et Denizot 1 2.44 17.03 35.47
Peyssonnelia harveyana P.L. et H.M. Crouan ex J. Agardh 13.55 12.38 10.18 11.88
Peyssonnelia polymorpha (Zanardini) Schmitz 4.11 10.06
Peyssonnelia rubra (Greville) J. Agardh 0.68 0.61 1.25 1.14
Turf
Acrothamnion preissii (Sonder) Wollaston 0.08 0.19 0.74 1.66
Botryocladia boergesenii J. Feldmann 0.04 0.06 1
Chondria mairei G. Feldmann 0.07 0.14 1
Chordariacea unidentified 1 0.03 0.06
Dasyopsis plana (C. Agardh) Zanardini 0.04 0.1
Derbesia tenuissima (De Notaris) P.L. et H.M. Crouan 0.17 0.4
Feldmannophycus rayssiae (J. et G. Feldmann) Augier et
Boudouresque 1 0.47 0.79

Halopteris filicina (Grateloup) Kutzing 0.03 0.08
Jania rubens (Linnaeus) Lamouroux 0.01 0.07 0.07
Laurencia obtusa (Linnaeus) Lamouroux 0.07 0.11 0.45 0.71
Laurencia pelagosae (Shiffner) Ercegovic 0.19 0.46
Polysiphonia setacea (Hollenberg) 137.11 34.99 108.04 21.96
Erect layer
Dictyota dichotoma (Hudson) Lamouroux 0.22 0.19 0.34 0.34
Flabellia petiolata (Turra) Nizamuddin 1 1.22 2.92
Galaxaura oblongata (Ellis et Solander) Lamouroux 1 0.04 0.1
Giffordia sp. 1 0.08 0.19
Halimeda tuna (Ellis et Solander) Lamouroux 11.24 5.79 25.82 17.56
22
Only species that showed a mean biomass higher than 0.01 g m either in treatments or controls are reported.
1 indicates mean biomass ,0.01 g m 22 ; M, mean; S.D., standard deviation (n56).

Table 6
Mean values and standard deviations of the total number of species (N), total biomass (B), ShannonWeaver
diversity index (H9) and species evenness (J) in treatment (reduced sedimentation rates) and control (natural
sedimentation rates) plots at the end of the experiment to determine the effects of sedimentation on the
composition and structure of the algal assemblage
Variable Treatment (S.D.) Control (S.D.) t df P Hartleys Fmax test
N 13 (1.26) 13.17 (1.72) 0.191 10 0.852 Fmax 51.854 P.0.05
B 188.64 (26.42) 196.86 (50.17) 0.355 10 0.729 Fmax 53.605 P.0.05
H9 1.216 (0.4) 1.679 (0.36) 2.106 10 0.061 Fmax 51.228 P.0.05
J 0.33 (0.11) 0.45 (0.08) 2.115 10 0.06 Fmax 51.791 P.0.05
Untransformed data were analysed using Students t-tests.
280 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

Fig. 6. Effects of small-scale spatial variation of sediment deposition on patterns of recruitment of macroalgae
on patches of bare rock cleared in (A) June 1992 and (B) December 1992. Data are mean percent covers (11
S.E.) on six replicates. Symbols as in Fig. 5.
 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288 281

Table 7
Analysis of the effects of sedimentation (reduced vs natural) and time of beginning of the experiment (June
1992 vs December 1992) on patterns of recruitment of the algal assemblage
Source of variation df MS F P Cochrans C test (transformation)
Filamentous turf
Sedimentation (5S) 1 129.9 2.338 0.142 C50.496 P.0.05
Beginning (5B) 1 561.6 10.11 0.047 a (none)
S3B 1 40.81 0.734 0.402
Error 20 55.55
Halimeda tuna C50.687 P,0.01
Sedimentation (5S) 1 1.743 1.680 0.209 (none)
Beginning (5B) 1 3.657 3.526 0.075
S3B 1 2.249 1.037 0.156
Error 20 1.037
Dictyota dichotoma C50.503 P.0.05
Sedimentation (5S) 1 0.106 0.018 0.895 (none)
Beginning (5B) 1 11.84 1.996 0.173
S3B 1 11.31 1.907 0.182
Error 20 5.929
Padina pavonica C50.758 P,0.01
Sedimentation (5S) 1 0.371 0.465 0.503 (none)
Beginning (5B) 1 0.721 0.903 0.353
S3B 1 0.354 0.443 0.513
Error 20 0.798
Acetabularia acetabulum C50.567 P.0.05
Sedimentation (5S) 1 35.12 5.97 0.023 a (none)
Beginning (5B) 1 73.38 12.49 0.002 a
S3B 1 25.22 4.293 0.051
Error 20 5.875
Giffordia sp. C50.510 P.0.05
Sedimentation (5S) 1 81.04 18.82 ,0.001 a (none)
Beginning (5B) 1 30.58 7.104 0.014 a
S3B 1 28.52 6.627 0.018 a
Error 20 4.304
ANOVAs were performed on the average percent cover values of each species through time (see text for
further explanations).
a
Significant P values (P,0.05).

Recruitment of Halimeda tuna was very low both in treatment and control plots, and
during the experiment this species never reached mean percent covers higher than 3.4%
(Fig. 6). A tendency towards a major abundance of H. tuna in control plots produced in
June was observed, but such differences were not statistically significant (Table 7).
Dictyota dichotoma and Padina pavonica (Linnaeus) Lamouroux showed a marked
seasonal pattern, with a peak of abundance in late spring and summer (Fig. 6). These
species colonised both treatment and control plots, reaching percent covers sensibly
higher than in the adult surrounding assemblage, as also suggested by the comparison
282 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

with results of the first experiment (Fig. 5). No significant differences were observed
among treatments and controls (Table 7).
Also Acetabularia acetabulum (Linnaeus) Silva and Giffordia sp. showed marked
temporal patterns, colonising the cleared areas only during spring and summer months
(Fig. 6). However, patterns of recruitment of these species were influenced by both
variations of sedimentation rates and time of beginning of the experiment (Table 7). The
outcomes of such effects were different for the two species. A. acetabulum, in fact,
recruited especially in control plots produced in June, while Giffordia sp. exclusively
colonised treatment plots produced in both periods, reaching high percent cover only in
those produced in June (Fig. 6).

3.3.2. Destructive sampling

After 1 yr of succession, both treatment and control plots produced in June were
quantitatively dominated by the filamentous turf, which ranged from 85 to 94% of the
total biomass. Only ten other species showed mean biomass higher than 0.01 g m 22
(Table 8). Such a low number of species was mainly due to the negligible presence of
prostrate forms, which were almost exclusively represented by encrusting corallines with
low biomass. On the contrary, the abundance of erect species, especially Acetabularia
acetabulum, Dictyota dichotoma and Padina pavonica, was high if compared with that
of the natural assemblage in the study area (see controls of the first experiment and data
reported in Airoldi et al., 1995).
The influence of depositional environment on patterns of succession of the algal
assemblage was evident. The experimental reduction of sedimentation rates, in fact, not

Table 8
Mean biomass (g m 22 ) and standard deviations of the most abundant macroalgae in treatment (reduced
sedimentation rates) and control (natural sedimentation rates) plots at the end of the experiment to determine
the effects of sedimentation on the patterns of recruitment of the assemblage on cleared substrata
Species Treatments Controls
M S.D. M S.D.
Prostrate layer
Encrusting corallines 0.74 0.84 1.10 0.68
Turf
Acrothamnion preissii (Sonder) Wollaston 0.36 0.36 0.37 0.46
Chondria mairei G. Feldmann 0.04 0.08 0.03 0.05
Laurencia obtusa (Linnaeus) Lamouroux 0.04 0.1 0.44 0.56
Polysiphonia setacea (Hollenberg) 102.21 63.16 85.54 32.02
Sphacelaria cirrosa (Roth) C. Agardh 1 0.14 0.31
Erect layer
Acetabularia acetabulum (Linnaeus) Silva 0.27 0.25 1.10 0.94
Dictyota dichotoma (Hudson) Lamouroux 1.72 1.96 7.16 9.45
Giffordia sp. 1.84 3.81 0.40 0.88
Halimeda tuna (Ellis et Solander) Lamouroux 1.08 1.11 3.26 3.99
Padina pavonica (Linnaeus) Lamouroux 1 1.17 1.32
Only species that showed a mean biomass higher than 0.01 g m 22 either in treatments or controls are reported.
1 indicates mean biomasses ,0.01 g m 22 ; M, mean; S.D., standard deviation (n56).
 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288 283

Table 9
Mean values and standard deviations of the total number of species (N), total biomass (B), ShannonWeaver
diversity index (H9) and species evenness (J) in the experimental plots (treatment panels vs controls) at the
end of the experiment to determine the effects of sedimentation on patterns of recruitment of the algal
assemblage
Variable Treatment (S.D.) Control (S.D.) t df P Hartleys Fmax test
N 9.33 (1.21) 10.5 (0.84) 1.941 10 0.081 Fmax 52.095 P.0.05
B 109.12 (64.77) 100.69 (39.74) 0.272 10 0.791 Fmax 52.657 P.0.05
H9 0.44 (0.296) 0.82 (0.26) 2.342 10 0.041 a Fmax 51.291 P.0.05
J 0.14 (0.08) 0.24 (0.076) 2.307 10 0.043 a Fmax 51.221 P.0.05
Untransformed data were analysed using Students t-tests.
a
Significant P values (P,0.05).

only affected significantly the abundance of Acetabularia acetabulum, Dictyota dich-

otoma and Padina pavonica (whose total biomass was lower in treatment than in control
plots, Students t-test, t52.24, P,0.05), but also significantly decreased the overall
diversity and species evenness of the assemblage (Table 9). Such decrease of diversity
was partly determined by the increase of abundance of Polysiphonia setacea, whose
mean biomass was 19.5% higher in plots with reduced sedimentation rates. However, the
effects of sedimentation on the development of this species were not detected as
significant due to the high variability among replicate plots (Students t-test, t50.576,
P.0.05).

4. Discussion

4.1. Manipulation of depositional environment

Experimental manipulations of the subtidal depositional environment are still few

(Neushul et al., 1976; Rogers, 1983; Kendrick, 1991). The commonest approach used in
such experiments is the increase of sediment deposition through addition of known
amounts of sediment to the experimental plots. Reduction of sedimentation rates has
been attempted less frequently, through removal of sediment by water motion. In both
cases, the effectiveness of the manipulation is clearly dependent on the frequency of
application or removal of sediment as well as on environmental conditions (e.g., natural
sedimentation rates, water flow, composition of the biological community). However, no
attempt has ever been made to quantify the real effectiveness of the method employed.
Moreover, the duration and intensity of the treatment may greatly affect the outcome of
the experiments: prolonged sediment smothering, in fact, may have quite different
effects than short term deep burial by sediments (Marszalek, 1981; Rogers, 1990). In
this study transparent panels were used to reduce natural sediment deposition over small
portions of the algal assemblage, reducing the sedimentary flux by 35% on average. The
effectiveness of the panels was confirmed by the visual survey of the plots: after a few
months from the beginning of the experiments, in fact, the algal assemblage below the
panels looked clearer of sediment than in the control plots. This suggests that, although
284 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

the short-term reduction of sediment deposition below the panels was not quantitatively
very high, its persistence induced a long-term impact on the algal assemblage.
A major shortcoming of manipulative field experiments in which structures such as
cages, barriers or panels are used is the risk of introducing artifacts that may bias the
results (Connell, 1974; Underwood, 1986; Peterson and Black, 1994). This concern is
generally evaluated by integrating into the experimental design a procedural control
which should reproduce the method used in treatments but not affect the variable of
interest (but see Johnson, 1992). In the present study, however, we could not find any
adequate control to unconfound potential artifacts: partial panels, in fact, do not mimic
accurately the presence of the entire panels, while introducing a partial manipulation of
sedimentation rates. As an alternative, we directly quantified in the field the effects on
those variables (i.e., penetration of light and flow microenvironment) that could be most
affected by the presence of panels. Results have shown that the accurate and constant
maintenance of the panels kept the effects on irradiance down to a level which
presumably had no biological relevance. Effects on the flow microenvironment were not
detected either. Artifacts on grazing pressure were not assessed, but these may be
reasonably considered irrelevant due to the scarcity of grazers in the area. Possible
confounding effects on the inputs of propagules were not quantified either. However,
dispersal of propagules generally occurs through lateral transport by bottom currents
(Santelices, 1990; Amsler et al., 1992), which were not significantly affected by the
presence of panels.

4.2. Influence of depositional environment

Small-scale variation of the depositional environment significantly influenced the local

structure and diversity of the algal assemblage. Such effects were especially evident
during the colonisation of patches of bare rock and were dependent on the time of
beginning of the succession. The erect algae (i.e., those species developing above the
filamentous turf) were the most affected by variations of the depositional environment
and showed different responses to the experimental reduction of sediment deposition.
The filamentous turf quantitatively dominated all the plots, but its growth was potentially
enhanced by reduced sediment loads. At the end of the experiments the overall diversity
and evenness of the assemblage was lower in treatments subjected to reduced
sedimentation rates.
The above results provide support to the hypothesis that small-scale spatial variability
of depositional environments may affect the within-habitat diversity of algal assem-
blages. In the study area such influence was exerted particularly through control of
abundance of dominant species rather than control of the overall composition and
species richness of the assemblage. This suggests not only direct effects on individual
plants or their propagules, but also indirect effects through mediation of competitive
outcomes.
Direct effects of sediment on algae in rocky habitats include smothering and scouring.
The morphological and life-history attributes that might confer adaptive ability to algae
in environments with sediment deposition and movement have been discussed by several
authors (Daly and Mathieson, 1977; Stewart, 1983; Littler et al., 1983; Airoldi et al.,
 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288 285

1995); these include tough thalli, the regeneration of upright axes from basal thallus
parts that can physically resist or tolerate smothering and scouring, opportunistic cycles
of reproduction and growth and the capacity to reproduce vegetatively, which removes
dependence upon spore attachment to buried substrata and allows a rapid recovery from
damage. Adaptations of this kind can be found in all the major space occupiers of the
study area (Airoldi et al., 1995), and are particularly noticeable in the dominant species
Polysiphonia setacea. Our results, however, suggest that rates of growth of the turf were
negatively affected by sediment smothering. The abundance of the turf, in fact, tended to
increase under reduced sedimentation rates, although this pattern could be perceived
only from detailed measurements of the biomass at the end of the experiments. Higher
rates of growth of filamentous turfs with lower sediment supplies have been also
observed by Adey and Goertemiller (1987) in the Caribbean and by Kendrick (1991) in
the Galapagos. Sediment may be considered a structural constituent of algal turfs
(Scoffin, 1970; Stewart, 1983; Airoldi et al., 1995), and the characteristics of deposition-
al environment might directly affect their patterns of growth. Experimental inves-
tigations, including manipulations of the quality and size spectrum of sedimenting
material as well as estimates of the turf biomass, are actually in progress to verify this
hypothesis.
The results of clearing experiments suggest a possible role of competitive interactions
as a structuring factor for the algal assemblage, at least within the time scale considered.
The removal of the resident organisms, in fact, enhanced the recruitment of some erect
species which were sparse at the study site. Such effects were particularly evident in the
plots produced in June and exposed to natural sedimentation rates. A possible
explanation is that the algal turf could preclude the development of other species by
exploitation of space and / or interference interactions through overgrowing, and that
local stress and disturbance are necessary for the persistence of some components of the
assemblage. Variations of sedimentation rates might, therefore, influence indirectly the
distribution of some species and the local structure and diversity of the assemblage
through affecting patterns of growth of the algal turf.
Similarly to observations in other sediment-impacted areas (Robles, 1982; DAntonio,
1986), grazing pressure is low during most of the year at our study site (Airoldi, pers.
obs.), and disturbance is mainly related to physical factors (i.e., sediment scouring and
wave action) which vary greatly through time (Airoldi et al., 1996). The effects of
manipulation of the depositional environment varied among patches of cleared sub-
stratum produced at different times of the year, possibly reflecting differences in
propagule availability and post-settlement growth and mortality of organisms (Sousa,
1984; Breitburg, 1985; Menge, 1991). In particular, variations of sedimentation rates
affected recruitment of species especially in quadrats produced in June, while no effects
were observed in plots cleared in December. In the study area maximal sediment
deposition and movement generally occurs in the autumn, while in the summer the
assemblage is free from sediment burial and scour (Airoldi et al., 1996). Predictable
temporal patterns in the disturbance regime, together with the modes of recolonisation of
cleared areas could, therefore, be important determinants of the effects of sediment
deposition on the overall diversity of the assemblage.
In conclusion, the experimental data of the present study support the hypothesis that
286 L. Airoldi, F. Cinelli / J. Exp. Mar. Biol. Ecol. 215 (1997) 269 288

small-scale spatial variations in the depositional environment may affect the develop-
ment and local diversity of algal assemblages; the outcomes, however, are dependent on
a variety of factors, including disturbance regime, successional patterns and morphologi-
cal and life-history characteristics of the important space occupiers. It is suggested that
the causal mechanisms that shape algal communities in sediment impacted areas rely
either on direct effects on individual plants or their propagules and on indirect effects
mediated by competitive interactions. Multifactorial experiments, including manipulation
of both physical and biological factors, will help elucidate this hypothesis.

Acknowledgments

We wish to thank Marco Abbiati, Luigi Piazzi, Salvatore Portella, Stefano Acunto and
the Guardia di Finanza of Livorno for collaboration in the field work, Fabio Rindi for
help with the determination of algae, Lisandro Benedetti-Cecchi and Gary A. Kendrick
for help with the statistical analysis and valuable comments on the manuscript, and two
anonymous referees for helpful revision of the manuscript. This study was completed as
a part of the doctoral thesis research of Laura Airoldi at the University of Genova, Italy.

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