Bioresource Technology 58 (1996) 299-307
Copyright © 1997 Elsevier Science Limited
ELSEVIER PII:S0960-8524(96)O0120-4
DECLINE OF ULVA GROWTH IN THE LAGOON OF VENICE
Adriano Sfriso* & Antonio Marcomini
Department of Environmental Sciences, Universityof Venice, Calle Larga S.Marta 2137, 30123 Venice, Italy
(Received 11 July 1996; revised version received 23 August 1996; accepted 30 August 1996)
Abstract
Causes and effects of the remarkable regression of the
macroalga Ulva rigida C. Ag. in the central part of the
Venice lagoon since 1990 are reported. Climatic
changes triggered the progressive reduction of Ulva
coverage and production ( - 8 0 % in 1993; - 9 5 % in
1995) until its almost complete disappearance in 1996.
Grazers, especially Gammaridae, controlled 70% of
the current biomass production, on a yearly basis, and
largely overcame the biomass production in summer.
Concurrently, the sediment resuspension increased by a
factor of 6-10 in the lagoon areas where the biomass
disappeared, contributing significantly to control of the
growth of the residual biomass by light limitation,
especially by settling of a microlayer of sediment over
the laminar thalli. Finally, biomass harvesting, now
effective, and the heavy disturbance caused by fishing
boats equipped with hydraulic dredges to catch Tapes
philippinarum, a bivalve species recently introduced
into the lagoon and spread in the areas previously
covered by Ulva, currently play an important syner-
gistic role in Ulva control. Conversely, the overall
availability of nutrients did not appear to have any
significant influence on the observed macroalgal
decline. Copyright © 1997 Elsevier Science Ltd.
Key words: Venice lagoon, Ulva rigida, climatic
changes, particulate matter, grazing pressure, bio-
mass harvesting.
INTRODUCTION
Over the last century the lagoon of Venice has
undergone large modifications in its hydrodynamic,
pollution and eutrophication levels. In particular,
the uncontrolled discharge of nutrients by the indus-
trial area of Porto Marghera, which reached its
highest expansion during the second post-war
period, and the digging of wide and deep commer-
cial canals (Pavoni et al., 1992; Sfriso et al., 1992a;
Marcomini et al., 1995) has caused serious damage
to the environmental conditions of the central
lagoon located between the Malamocco-Marghera
*Author to whom correspondence should be addressed.
299
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0960-8524/96 $15.00
canal (Canale dei Petroli) and the salt marshes of
Burano and Torcelio. As a consequence, the physi-
cal and chemical characteristics of waters and
surface sediments promoted the occurrence of dys-
trophic conditions which, in turn, greatly affected
the lagoon phytobenthic associations. A dense and
monospecific population of Ulva rigida C. Ag.
replaced the rhizophyte meadows (Zostera marina
L., Zostera noltii Hornem. and Cymodocea nodosa
(Ucria) Asher.) previously covering the whole
lagoon. About half of the central lagoon (66 km 2)
was steadily covered by a deeply stratified macro-
algal biomass (3-20 kg fresh weight (fwt)m -z)
alternating conditions of luxury growth with others
of biomass collapse. Markedly high amounts of
nutrients were released by the decomposing biomass
and temporarily accumulated in the surface of the
sediments. In the 1948-1983 period the mean sedi-
ment concentrations of phosphorus and nitrogen
increased by approximately 30 and 2.3 times, respec-
tively (Perin, 1975; Perin et al., 1983), but decreased
slightly over the subsequent years (Sfriso et al.,
1995). The annual amount of nutrients involved in
macroalgal growth was by far higher (6.5 and 10.2
times for P and N, respectively; Sfriso & Marcomini,
1994) than the total nutrient inputs (P = 351 and
N = 2578 tonnes, Andreottola et al., 1990) entering
the lagoon by direct (industrial, agricultural, civil
and rainfall) and indirect (rivers and canals) sources.
Nutrients were not only recycled but also stored
temporarily both by the macroaigal standing crop (P:
--~119 tonnes, N: ~1355 tonnes) and the 5 cm top
layer of sediment (P: ~726 tonnes, N: ~4752
tonnes). In addition, the nitrogen loss occurring in
surface sediments and in the stratified biomass by
denitrifying processes was estimated to be
~10000-11000 tonne year ' (Sfriso & Marcomini,
1994; Silvestri et al., 1994), accounting for about half
of the total nitrogen recycled by the gross primary
production (GPP). The northern Adriatic sea was
firstly hypothesised, and then experimentally proved
(Sfriso et al., 1994), to act as an additional supplier
of nitrogen for the lagoon waters. Usually sea
coastal waters exhibit nutrient concentrations lower
than those in the lagoon, but over the March to
300 A. Sfriso, A. Marcomini
September period, during the luxury springtime algal
growth, the concentrations of nitrogen compounds
(especially nitrate) were found to be higher in the
sea than in the lagoon waters between sea inlets and
the areas populated by macroalgae (Sfriso et al.,
1994). Similarly, measures of nutrient concentrations
along the water column of the lagoon mouths during
flood and ebb tides showed that in the spring-sum-
mer nitrates were imported from the sea by tidal
exchange. The lagoon behaved as a denitrifying
reactor transferring land and sea nitrogen into the
atmosphere. Similar conclusions were inferred for
carbon (Sfriso & Marcomini, 1994), through a
marked lagoon import of bicarbonates from the sea
and a lagoon loss of CO2 into the air (Lvov et al.,
1996). No significant lagoon import of phosphorus
from the sea to the lagoon was found.
Starting from 1990, a progressive decrease of Ulva
density and distribution has been recorded over the
whole lagoon.
The objective of this work was the investigation of
factors responsible for the changes of biomass distri-
bution and production observed in the lagoon of
Venice over the last 5-year period. The available
data (physical parameters such as water tempera-
ture, nutrient concentration in water and surface
sediment, biomass production and sedimentation
fluxes) were routinely examined and compared with
the current measures of nutrient concentrations in
surface sediments and sedimentation fluxes. Also, a
field investigation was undertaken to assess the graz-
ing pressure by invertebrate herbivores and the
effect of biomass harvesting.
METHODS
Study area
The lagoon of Venice is a shallow water basin on
the west coast of the northern Adriatic sea. The
average mean depth referred to the mean tide level
is ~ 1 m. In more detail, ca. 75% of the total water
surface is within 2 m deep and only ~ 5 % of the
whole area is 5 m. The mean tide difference is
about +31 cm (Pirazzoli, 1974) and ca. 60% of the
total lagoon water is exchanged with the sea at any
12-h cycle. The water renewal is complete in a great
part of the lagoon, with the exception of some inner-
most areas where the water residence time may be
some days, depending on the tide regime: spring or
neap tides. The total lagoon area is ,,~549 km 2,
whereas the area exposed to the tides is -,~432 km 2.
The Venice lagoon is subdivided into three natural
areas, i.e. southern, central and northern basins,
which are separated by the Malamocco-Marghera
canal at the south and by the Burano-Torcello tidal
lands at the north. This study focused mainly on the
central part of the lagoon (~132 km 2 exposed to
tidal flows) where the main changes were observed.
Analyses
The temperature of the water column was measured
by a thermocouple electrode (precision 0.1°C) in a
mixed sample of 5-6 sub-samples collected by a
homemade bottle (i.d.: 4 cm, height: 150 cm)
plunged vertically into the water.
The photosynthetically active radiation
(PAR = 400-700 nm) and underwater transmission
was measured by a LI-193SA Spherical Underwater
Quantum Sensor (LI-Cor, Nebraska, USA).
The in situ biomass density was determined by an
aluminium, 1 square metre box (100 x 100 x 70 cm)
repeatedly (6-10 times) placed on the bottom
according to the procedure of Sfriso et al. (1991) for
shallow environments. The precision of biomass
measures varied from 1 to 3% in the presence of
standing crops of 5 kg fwt, to 6-7% when the bio-
mass was below this density.
The annual biomass production was estimated by
the biomass change method by summing the biomass
increases found significant (P<0.01) by ANOVA
(Sfriso et al., 1993) with a weekly (spring-summer)
or twice a week frequency (autumn-winter).
The grazing pressure of phytozoobenthos was
estimated by suspending samples of Ulva in cubic
wire net cages (25 x 25 x 25 cm). Homogeneous sub-
samples of Ulva biomass were dripped by a slow
centrifugation, the zoobenthic organisms removed
and aliquots of 150 g were suspended in the above
cages near the bottom within the natural biomass.
Three cages with 1 cm (allowing the grazing by the
more frequent herbivores) and three with 1 mm
(hampering any significant grazing) mesh sizes were
used. The reduction of light transmission inside the
1 mm mesh size cage was satisfactorily avoided by
building four of the six sides in plexiglas. The bio-
mass variation within the cages was recorded and
reset to 150 g fwt every 4-10 days, depending on
the season, by an electronic balance (precision:
0.1 g). The mean differences between the per cent
relative growth rate (%RGR) in the cages of dif-
ferent net sizes calculated by the formula
% RGR = ln(BJBo)/t x 100
(where B, = biomass at day t and Bo = the initial
biomass) showed the amount of macroalgae grazed
by the herbivores, which in the lagoon are repre-
sented mainly by small amphipods and isopods. The
growth rate estimations within the cages were
affected by an error of 5-10%. The particulate
matter (SPM) settling on the surface sediment
during time-integrated periods was collected by sedi-
ment traps (15 × 15 x 10 cm) designed especially for
shallow environments under strong tidal regimes
(Sfriso et al., 1992b). Sampling was performed with a
frequency depending on deposition rates
(15-30 days). Aliquots of ~250 ml of two mixed
SPM traps were stored in glass pots, frozen and
lyophilised for the determination of the dry weight
 Decline of Ulva growth in the lagoon of Venice
and estimating the SPM fluxes on the lagoon bot-
toms.
The 5 cm top layer of 5-6 surface sediment
cores, collected by a plexiglas corer (i.d.: 10 cm) and
mixed together, provided the sediment for nutrient
concentration analyses. Inorganic and total phos-
phorus were determined by shaking overnight in 1 N
HCI 0.2 g of finely homogenised sediment, before
and after 2 h combustion at 550°C, according to
Aspila et al. (1976). Phosphorus concentrations were
obtained spectrophotometrically by the phosphomo-
lybdenum blue method according to Strickland and
Parsons (1972). Organic phosphorus was calculated
by difference. Total nitrogen and organic carbon
were determined by a Carlo Erba CNS Elemental
Analyser model EA 1108, and organic carbon was
obtained after the removal of carbonate with 1 N
HCI.
RESULTS AND DISCUSSION
Observed changes of Ulva coverage and production
Starting from 1990 Ulva disappeared almost com-
pletely at the northern end of the translagoon
bridge, being replaced by frequent phytoplankton
blooms, and the presence of Ulva in the rest of the
lagoon was progressively reduced. The Ulva cover-
age, previously extending over ~66 km 2 of the
central lagoon, with a mean density of
3 kg fwt m 2 (Sfriso et al., 1993), decreased to
14 km 2 in 1993, to 2-3 km 2 in 1995 and currently
has almost completely disappeared. Concurrently,
the highest standing crop and the annual net pro-
duction of Ulva dropped from ~0.6 and ~1.6
million tonne fwt before 1990, to 0.1 and 0.4 million
tonne fwt in 1993, and to some thousands of tonnes
in 1996. Tubulous and ribbon-like genera, such as
Enteromorpha and Gracilaria, and other seasonal
species, i.e. Cladophora, Scytosiphon, Petalonia, Spyr-
idia, etc., spread in the bottoms previously covered
by Ulva, but without significant biomasses. Species
widespread in the past, such as Monostroma grevillei
Water Temp. (°C) - Biomass (kglm =, fwt)
ii ..................
Jan-89 May-89 Sep-89 Jan-90
-- Temperature ~
Fig. 1. Water column temperature and Ulva biomass trends during the 1989-1991 period. Vertical arrows show the same
data at successive years, the other ones the critical temperature periods.
301
(Thuret) Wittrock, Stictyosiphon adriaticus Kutz.,
Dictyopteris membranacea (Stackh.) Batt., Cystoseira
spp. , Cladophora prolifera (Roth) Kutz. and Laur-
encia papillosa (Forsk.) Grey., reappeared, while
new species never found before [Grateloupia dor-
yphora (Montagne) Howe, Sargassum muticum
(Yendo) Fensholt (Gargiulo et al., 1992; Curiel et
al., 1996a), Undaria pinnatifida (Harvey) Suringar
(Curiel et al., 1994), Antithamnion pectinatum (Mon-
tagne) Brauner in Athanasiadis of Tittley,
Radicilingua reptans (Kylin) Papenfuss, Radicilingua
thysanorhizans (Holiness) Papenfuss (Curiel et al.,
1996b) and Castagnea zosterae Thur.] were also
recorded. Although the phanerogams Zostera noltii
Hornem. and Zostera marina L. repopulated many
areas where Ulva had declined, this repopulation of
the bottom has been going on very slowly and up to
now it is confined to the canals and the shallowest
waters.
Changes of climatic conditions
The effect of unfavourable climatic conditions on
Ulva growth, accounting for 2-4 weeks of tempera-
ture fall in May-June during consecutive years
(1989-1991), may be regarded as an important
factor responsible for the initial decrease of Ulva
dominance. Figure 1 shows the temperature trend of
the water column together with the standing crop
changes recorded in this period in an area previously
heavily covered by Ulva. During the 1989 spring,
water temperature progressively increased from the
winter to the summer values without significant fluc-
tuations and the biomass quickly reached
~20 kg fwt m 2. After this, the biomass collapsed
during an anoxia. In May 1990, water temperature
increased up to 24°C then, for 3 consecutive weeks,
fell by 3°C. Biomass stopped growing at
10 kg fwt m 2 and rapidly collapsed. The year
after, 1991, water temperature reached 18°C and
successively decreased by ~-4
3 o (_ for ca. 1 month.
Biomass started to grow only in June and stopped at
10 kg fwt m 2 During this 3-year period
(1989-1991) the minimum water temperature (ca.
May-90 Sep-90 Jan-91 Ma' -91
Biomass /
]
302 A. Sfriso, A. Marcomini

2-3°C) shifted from January to March, thus pro-
longing the winter conditions for 2 months.
However, the direct effect of water temperature on
the growth rate of Ulva appeared to be far less
important than the influence of the cloudiness and
wind-induced water turbulence associated with the
adverse meteorological conditions. In fact, the
growth of Ulva in the temperature range 15-21°C
recorded in these adverse periods was not signifi-
cantly far from that of the optimal growth interval:
23-24°C (Duke et al., 1989; Sfriso, 1995; Riccardi &
Solidoro, 1996). Long periods of adverse climatic
conditions can, however, reduce the Ulva growth
quite effectively, especially by light inhibition due to
the settlement of resuspended sediment on the lam-
inar thalli (Sfriso & Pavoni, 1994). In addition,
during periods of cloudy weather, the availability of
nutrients resulting from the macroalgal decay and
the sediment release, due to the wind-induced water
turbulence, greatly favoured phytoplankton blooms.
Phytoplankton takes advantage of Ulva since its
growth is not hampered by particulate-matter settle-
ment, and water turbulence can expose it to more
favourable light conditions.
In 1994 and 1995 the climatic conditions were
similar to those found in 1990-1991, with the mini-
mum water temperature in March and long periods
rates occurring at Sacca Sessola over the 1989-1990
period, in the presence of high and wide biomass
coverage, and in 1992-1993, when the biomass was
spread in patches of only some 10 square metres.
On a yearly basis, in the presence of the reduced
biomass coverage, sedimentation fluxes increased by
more than four times, from ~ 65 to
,-~275 kg dwt m -2. The close relationship between
biomass coverage and the SPM fluxes, both at Sacca
Sessola and at San Giuliano, are displayed in Fig. 3.
During springtime, high biomass levels were coupled
with the lowest SPM fluxes (on average:
20-30 g dwt m -2 day-l), whereas negligible bio-
masses, such as that monitored over the rest of the
year, corresponded to SPM fluxes higher by more
than one order of magnitude (on average:
300-550 g dwt m -2 day-l). In 1992-1993, in the
presence of a reduced biomass coverage, the SPM
fluxes at Sacca Sessola were ~0.5-1.0 kg dwt m -2
A L B E R O N I (SPM = 41 k g l m z y)
1989-90
600
"0 400
200
O1
0

eoo
Jan Mar Ju! Sep Jan
of adverse conditions until May. The highest Ulva
S A C C A S E S S O L A (SPM = 65 kglm = y)
biomass was found in June, then Ulva was com-
pletely harvested (Sfriso & Marcomini, 1996).
,00
Biomass and SPM fluxes
A major effect of the adverse climatic conditions "--
O1 200
was the increased water turbidity and particulate- 0
matter settling. In order to quantify the Ulva growth Jan Feb Apr Jun Jul Sep Nov Jan
limitation due to sediment resuspension, the sedi- SAN G I U L I A N O (SPM = 140 k g / m 2 y)
ment fluxes of settling particulate matter (SPM)
were recorded in some areas covered by different ,1~ 900~............................
biomass densities and phytobenthic associations 600 . . . . . . . . . . . . . . . . . . . .
after the observed Ulva decline and the results were ~1 300
compared with those acquired before the decline. By 0
Jan Mar May Jul Oct Dec
means of suitable bottom traps that allowed collec-
tion and time integration of SPM during seasonal SACCA SESSOLA (SPM = 275 kglm" y)
and yearly intervals, it was found that the SPM 1992-93
fluxes were strictly dependent on the biomass cov-
•0 3000
erage. Figure 2 shows the histograms of
2000
sedimentation rates measured during two different
~I 1000
annual periods at stations located along the lagoon
0
mouth (Alberoni)-mainland (San Giuliano) tran- Apr Jun Aug Oct Dec Feb
sept. As expected, the annual amount of SPM
F U S I N A (SPM = 846 k g l m ' y)
increased progressively according to the landward
direction; i.e. from Alberoni near the sea and with a
sandy sediment (fraction <63 /~rn: <20%), in the "O 3000
direction of Sacca Sessola and San Giuliano. The
~. 1500
difference in SPM found between the latter two O)
stations, characterised by a fine sediment (fraction 0
Apr Jun Aug Oct Dec Feb
<63 /~m: from 80 to 100%, respectively) and high
Fig. 2. Fluxes of settled particulate matter (SPM) in some
biomass levels, appeared related mostly to the dif- areas of the central part of the Venice lagoon. The station
ferent biomass densities and biomass growth times. of Sacca Sessola was sampled in 1989-1990 and
The same Fig. 2 shows the changes of sedimentation 1992-1993, before and after the Ulva decline.
 Decline of Ulva growth in the lagoon of Venice 303

day -~, with peaks up to 3.3 kg dwt m - 2 day-'. factors limiting the algal growth because of the cov-
Before 1990 the waters of the lagoon were very erage effect on the macroalgal biomass. Light
clear, except over 1-2 months of massive biomass limitation occurs directly on the surface of the lam-
collapse, which triggered coloured waters (resuspen- inar thalli that are covered by a microlayer of SPM,
sion of colloidal sulphur and/or reduced compounds) so reducing algal growth (Sfriso & Pavoni, 1994).
and phytoplankton blooms, whereas nowadays the Furthermore, high sedimentation rates prevent any
central lagoon exhibits turbid waters throughout the spore or germling attack on shells and on any other
year. Clear waters, replaced by turbid ones at a dis- available hard substrate. The light transmission in
tance of only a few tens of meters from the biomass the water column was also highly reduced, although
edges, are occasionally found in spring only at the the irradiance measured close to macroalgal layers
Lido watershed near to the residual meadows of during the different seasonal photoperiods
Ulva that are currently growing close to the Lido (50-80 /~moi m - 2 s - J at minimum, in the presence
island. Then, between April and June, the biomass is of rain and cloud) was always sufficient to support
completely harvested by the reaping machines of the the growth of macroalgae. In fact, literature data on
Venice Municipality and the waters become turbid the compensation irradiance for Ulva report values
until a new significant biomass coverage occurs in ranging from 9.5 /~mol m 2 s - ' (Keith & Murray,
the successive year. 1980) down to 1.9 /tmol m -2 s ~ (Riccardi & Sol-
The inverse correlation between SPM fluxes and idoro, 1996).
the biomass level monitored at Sacca Sessola and
San Giuliano is plotted in Fig. 4 as a logarithmic Grazing pressure
relationship. The SPM fluxes were all below Under the conditions monitored after the Ulva
250 g dwt m -2 d a y - ' for biomasses exceeding decline, zoobenthic invertebrates, mainly amphipods
,-~3 kg fwt m -2. In contrast, in the presence of of the genus Gammarus, isopods of the genus Ido-
negligible biomass, SPM fluxes showed the highest tea, snails of the genus Monodonta and shrimps of
variability, depending on water turbulence. The the genus Pale•on, may play a key role in control-
fluxes of SPM appeared to be strictly dependent on ling the growth and abundance of macroalgae in the
the sediment coverage by the algal biomass but, at Venice lagoon. In the past, when Ulva exhibited bio-
the same time, SPM fluxes were one of the main mass of 10-20 kg fwt m - 2 o v e r tens of square

Sacca Sessola 1989-90

[ ]
i_ O, Biomass • SPM
J
20000
15000
10000
t " -~ iiiii
- .......................
i iiiiii iiiiii~,ii, ii i~iiiiiiiiii ~ilt ,500 '=.=:~
oOE2oo ...
01 5000
o In' • " ": ' : ~ : - -, -'-'" -- o
Mar Apr May Jun Jul Sep Oct Jan Mar

San Giuliano 1989-90

l © Biomas. • SPM ]
90001 . . . . . . . . . . . . . . . . I 1200 "I~

O)
3000 - ~ ............. ~ ........... ---.---- ,oo "O
ol" • -" =
Mar Apr May Jun Jul Sep Oct Jan

Sacca Sessola 1992-93

I ~ Biomass • SPM j
,0000
15000 ..................
t
=_ 3000 y
~1ooo0~t- , . . . . . . . . . . . . . . . . . . . . . _I2o0o ~
5000 l V . . . . . ~ _ _ _ ~ . . . . . . . . . ' _-= . . . . . . -J 1ooo
oi= = = "= " ~ " '-, o =
Apr May Jun Jul Aug Sep Oct Nov Feb
Fig. 3. Biomass and SPM relationship in some macroalgae highly populated areas of the Venice lagoon.
304
E
300(
200(
o
SPM-Biomass logarithmic relationship
y = 907-201*loglO(x)
i. . . . .
o The role of grazers in controlling the composition
Iooo
0 :
n . n found to be important in other shallow environ-
u) 0 ~00 0 0 ; O" u :
"I°?~;oo 20oo .oo .oo
Biomass (g fwt/m z d)
Fig. 4. Logarithmic relationship between macroalgal bio-
mass and SPM fluxes.
kilometres, the effect of the grazing pressure was
surely negligible in comparison with the biomass
production because zoobenthic populations were kil-
led by the frequent anoxic crises occurring in the
lagoon (Sfriso & Pavoni, 1994). Only outside the
anoxic areas was herbivore feeding significant, and
the surviving grazers (especially Gammaridae)
crowded together grazing macroalgae at rates up to
15% day-l (Sfriso & Pavoni, 1994). Nowadays, with
anoxias occurring only extraordinarily and for short
periods, herbivores are spread all around the lagoon
and grazing takes place without significant seasonal
variations. The amount of macroalgae consumed by
herbivores in 1994-1995 is shown by Fig. 5. The
1 mm mesh size Ulva cages, preventing the intro-
duction of grazers, displayed Ulva growth rates
between - 0 . 3 and 6.4% day -1 , with an annual
mean of 3.2% day -1. The cages with 1 cm mesh
size (including grazers) showed growth rates ranging
from - 2 . 8 to 5.5% day -I , with a mean of
1.1% day -1. Excluding a light limitation effect
within the 1 mm mesh size cages because of the
presence of plexiglas sides, the resulting grazing
pressure was between 0.1 and 6.4% day -1 , with an
annual mean value of 2.2% day--I. On an annual
basis, grazing reduced Ulva production by ca. 70%.
The role of invertebrate feeding was particularly
important when, between July and August, macro-
algal production was negative and decreasing by a
rate of over 2% day -1. If these results are extrapo-
lated to the natural biomass, the effect of the
grazing pressure, combined with the spring-summer
TOTAL CAGE RGR (mean: 3.2)
7 . . . . . . . .
~ 3
1 1994-1995, during the spring-summer growth explo-
m Fieldq-em°,'~RG-R,..... t~0) • %;-ra-z|ngpre~;uro(mun~"2.;)]
Fig. 5. Percent relative growth rate (%RGR) in cages with
mesh sizes of 1 mm (total area: black+grey) and 1 cm
(grey area). The grazing pressure (black area) is provided
by the difference.
A. Sfriso, A. Marcomini
.
] biomass harvesting and the growth limitation due to
higher sedimentation rates found in the absence of
biomass coverage, appears to comprehensively
explain the current Ulva distribution and accumula-
tion in the lagoon.
and density of macroalgal communities has been
u O ;
ments (Lubchenco & Gaines, 1981; Hawkins &
Hartnoll, 1983; Johnson & Mann, 1988; Hansen et
110oo 14ooo al., 1993; Sfriso & Pavoni, 1994; Jak et al., 1994;
Scholten, 1995). Hansen et al. (1993) found in a
eutrophic Danish estuary (Roshilde Fjord) that the
grazing pressure due to the genus Idotea controlled
the growth and the biomass accumulation in the
free-floating fronds of Ulva lactuca at rates up to
40% day-1. In this case the biomass coverage was
very low (5-10%) and the grazing activity could
completely remove the available biomass. The same
effect was observed in the Venice lagoon in 1992
when small biomass samples were suspended in
cages placed in areas where macroalgae were natu-
rally missing (Sfriso & Pavoni, 1994).
Nutrient concentrations
The nutrients flowing into the lagoon, especially of
industrial and urban origins, have been markedly
reduced over these last 20 years (Pavoni et al., 1992;
Sfriso et al., 1992a; Marcomini et al., 1995) because
of the remarkable decline of the industrial activity in
Porto Marghera, the coming into operation of mech-
anical-biological sewage treatment plants (i.e.
Campalto, Marghera, Lido, etc.) and the banning of
phosphorus (1989) in commercial detergents. The
overall effect of these changes in the central lagoon
is inferable from the concentrations of ammonium
over the 1962-1988 period. Near the industrial area
of Porto Marghera, this ion decreased from
1000-2500/~M to ca. 10 #M. In the surface sediment,
after an increasing trend observed up to the 1970s,
both total nitrogen and phosphorus began to
decrease (Sfriso et al., 1995). This pattern is better
visualised in Fig. 6, which presents phosphorus,
nitrogen and organic carbon trends in the top 5 cm
sediment of a lagoon station monitored in
1985-1986, in the presence of massive macroalgal
populations, and 9 years later (1994-1995, this
work) when macroalgae were negligible. The highest
sediment concentrations of these elements in
" --
1994-1995 were similar to, or even lower than, the
winter concentrations recorded in 1985-1986. Com-
paratively, the concentration peaks found in
sion and/or decay of the algal biomass were
negligible for phosphorus and very low for nitrogen
and organic carbon. Total phosphorus without the
contribution of the organic algal fraction was
steadily around 0.4 mg dwt g-1 throughout the
year, showing a decrease of ~50% between the
highest concentrations monitored in summer 1985
 Decline of Ulva growth in the lagoon of Venice 305

t985-1986 1994-1995

eoo
0') 4001

200'
Phosphorus
==soo
~ 400

200
Phosphorus

0 0
Mar Apr May Jun Jui Sep Oct Jan Apr Feb Apr May Jul Aug Sep Nov Jan

Total Nitrogen Total Nitrogen

~ 2 . . . . . . . . . . . . . . .
E'
1
Mar Apr May Jun Jul Sep Oct Jan Apr Feb Apr May Jul Aug Sep Nov Jan

Organic Carbon Organic Carbon

16 . . . . . . . . . . . . . .

~ ) 13
10
. . . . . . . . . . . . . . . . .
- --
~113 . . . . . . . . . . . . . . . . . .~ . .v . ~. . ~

7
Mar Apr May Jun Jul Sep Oct Jan Apr Feb Apr May Jul Aug Sep Nov Jan

Fig. 6. Total (black squares), inorganic (grey circles) and organic phosphorus (black triangles), total nitrogen and organic
carbon concentrations in the surface sediments (5 cm) at the Lido station in 1985-1986 and in 1094-1995. Arrows show
anoxic periods.

and 1994. Similarly, in 1994, the highest nitrogen
and organic carbon concentrations were, respec-
tively, --~40 and ~ 3 0 % lower than those in 1985.
The concentrations of organic carbon and total
phosphorus were, on average, even lower than the
winter concentrations found in 1985-1986. Although
the reduction of nutrient inputs to the lagoon over
the last decade was significant (Sfriso & Marcomini,
1995), the biomass decline monitored over the last
5 years appears to be the main factor explaining the
decreased amounts of nutrient accumulations in the
surface sediment corresponding to the growth-decay
cycles of Ulva (Sfriso & Marcomini, 1994; Sfriso et
al., 1994).
Biomass harvesting
Another important factor controlling the expansion
of Ulva is certainly the biomass harvesting by reap-
ing machines suitable for operation in shallow
waters. The collection of the nuisance biomass
carried out since the 1980s by the Venice Munici-
pality, before the natural Ulva decline, did not
produce the expected benefit, since only a very small
amount of the total available biomass was harvested
( ~ 100-200 m 3 day J of fresh material, amounting
to 30000-50000 m 3 year-~). The algal standing crop
before 1990 in the central lagoon was ~0.6 million
tonne year ~, amounting to net and gross produc-
tions of ~ l . 6 and ~11 million tonne year--J,
respectively (Sfriso & Marcomini, 1994). In that
period, the only effect of the algal harvesting was
cleaning up small portions of the lagoon within
green meadows extending over tens of square kilo-
metres, thus increasing the grow'th of the residual
biomass. It is well known that lower standing crops
display higher growth rates (Lapointe & Tenore,
1981; Sfriso et al., 1993). In addition, biomass har-
vesting in shallow waters produced a very high
sediment resuspension and release of nutrients
immediately available for uptake by the Uh,a or, in
the case of local complete biomass removal, for
extraordinary phytoplankton blooms.
Quite different results have been obtained by the
biomass harvesting which has been carried out after
the natural Ulva reduction, from 1991 up to now.
Nowadays, because of the decreased natural produc-
tion, the collection of nuisance macroalgae
performed by Consorzio Venezia Nuova with the
support of aerial images and field monitoring is
effective and reliable for hampering possible new
biomass spread over the lagoon (whenever favour-
able climatic conditions could again allow Ulva
growth), although the harvested biomass is similar
or even lower than that collected in the past: from
53000 m 3 year ' of fresh algae in 1990 to ca. 9000
m 3 year ~ in 1994 (Consorzio Venezia Nuova, 1994)
and in 1995 (Bernstein, personal communication).
Catch of Tapes
The most recent factor (1993-1996) of increasing
importance for hampering the re-colonisation by
306 A. Sfriso, A. Marcomini
Ulva, as well as by the rhizophytes, of a great part of
the central lagoon is the very intensive fishing of
Tapes philippinarum, a bivalve species recently intro-
duced in the lagoon but already spreading
abundantly over the central lagoon with biomasses
up to 0.7-1 kg m -2, fwt. The fishing carried out by
means of heavy hydraulic dredges (weight: 0.4-0.6
tonnes, mouth: 2-2.5 m, compressing water to
3.4-3.6 atm, ICRAM, 1992) is not permitted in the
lagoon since it creates furrows up to 30-40 cm deep
that persist for 6-8 months. However, this fishing is
illegally performed overnight, and actually a great
part of the lagoon sediment appears heavily dis-
turbed by this fishing practice.
CONCLUSIONS
Unfavourable weather conditions during consecutive
years, extending winter-like conditions up to May-
June when Ulva usually exhibits its maximum
growth, prompted this species to progressively
reduce its spatial expansion. Following this, the
effects of environmental factors, such as the
increased resuspension of particulate matter due to
the feedback effect of the reduced biomass coverage
(causing light inhibition) and the increased grazing
pressure (no longer hampered by anoxic crises),
together with an efficient harvesting of the residual
biomass and the disruptive action of Tapes fishing by
hydraulic dredges, contributed significantly to fur-
ther Ulva reduction up to its almost complete
disappearance. In particular, both the increasing
sediment disturbance by the Tapes catching and the
grazing effect on the growth of macroalgae and rhi-
zophytes deserve a deeper investigation to
understand the future development of the macro-
phytes in the lagoon of Venice.
Other factors, such as the abatement of nutrient
inputs from the mainland, appear to have played a
minor role, if any, in the trophic changes observed
over the last 5 years. The recent findings (Sfriso &
Marcomini, 1994; Sfriso et al., 1994; Sfriso & Marco-
mini, 1996) on nutrient trends in the surface
sediments have confirmed that macroalgae control
the distribution and residence time of nutrients in
the Venice lagoon, and not vice versa.
ACKNOWLEDGEMENTS
This work was financially supported mainly by the
European Community in the framework of the
EUMAC-project (Eutrophication and Macrophytes)
as well as by the project of the Italian National
Council of Research (CNR) 'The Venetian Ecosys-
tem. Research line 4.1'. The authors are also
grateful to Mrs Sonia Ceoldo for the valuable tech-
nical assistance.
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