I TA L I A N H A B I TAT S

Lagoons, estuaries and deltas

23

Italian habitats Italian Ministry of the Environment and Territorial Protection / Ministero dellAmbiente e della Tutela del Territorio e del Mare Friuli Museum of Natural History / Museo Friulano di Storia Naturale Comune di Udine Scientific coordinators Alessandro Minelli Sandro Ruffo Fabio Stoch Editorial committee Aldo Cosentino Alessandro La Posta Carlo Morandini Giuseppe Muscio

I TA L I A N H A B I TAT S

Lagoons, estuaries and deltas Boundaries between the sea and rivers edited by Alessandro Minelli

Texts Annelore Bezzi Mauro Bon Francesco Bracco Daniele Curiel Francesca Delli Quadri Giorgio Fontolan Gilberto Gandolfi Alessandro Minelli Andrea Rismondo Francesco Scarton Margherita Solari Davide Tagliapietra Marco Ulliana Mariacristina Villani In collaboration with Maria Manuela Giovannelli Antonella Miola Marco Sigovini English translation Alison Garside Gabriel Walton Illustrations Roberto Zanella Graphic design Furio Colman Photographs Nevio Agostini 7, 15, 21, 82, 87, 88, 143, 145 Archive Museo Friulano di Storia Naturale 81, 84, 85, 90, 107/2, 148 Archive Naturmedia 86, 137 Mauro Bon 66, 132 Massimo Buccheri 80 Marco Cantonati 32 Compagnia Generale Ripreseaeree 12, 13 Ettore Contarini 101/2, 105 Giuseppe Corriero 58 Daniele Curiel 24, 25, 26, 27, 28, 29, 30, 31 Ulderica Da Pozzo 9, 11, 16, 19, 23, 135, 144 Vitantonio DellOrto 6, 20 ,41, 74, 75, 77, 112, 115, 116, 118, 119, 120, 121, 123, 124, 126, 127, 128, 129, 130, 131, 133, 134 Dario Ersetti 22, 39, 79, 147 Giorgio Fontolan 10, 18 Francesco Scarton 117, 122, 142 Paolo Glerean 138 Giuseppe Muscio 76 Nicola Novarini 113, 114 Arnaldo Piccinini 65, 68, 69, 70, 67 Andrea Rismondo 33, 34, 35, 36, 37 Lorenzo Seitz 40, 46, 47, 48, 56 Marco Sigovini 50, 51 Fabio Stoch 14, 59, 60, 61, 72, 140 Egidio Trainito 42, 44, 45, 52, 53, 54, 62, 63, 64, 71, 73 Marco Uliana 83, 91, 92, 93, 94, 95, 96,97, 98, 99, 100, 101/1, 102, 103, 104, 106, 107/1, 108, 109, 110, 111, 139 Roberto Valle 125 Augusto Vigna Taglianti 57 Francesco Zaramella 49, 141

Lagoons, estuaries and deltas

Boundaries between the sea and rivers

2009 Museo Friulano di Storia Naturale, Udine, Italy All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, without the prior permission in writing of the publishers. ISBN 88 88192 46 8 ISSN 1724-6539 Cover photo: Lagoon of Grado and Marano (Friuli, photo: U. Da Pozzo)

M I N I S T E R O D E L L A M B I E N T E E D E L L A T U T E L A D E L T E R R I T O R I O E D E L M A R E M U S E O F R I U L A N O D I S T O R I A N AT U R A L E C O M U N E D I U D I N E

Italian habitats

Contents
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Annelore Bezzi Francesca Delli Quadri Giorgio Fontolan Submerged vegetation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Daniele Curiel Andrea Rismondo

1 Caves and karstic phenomena

2 Springs and spring watercourses

3 Woodlands of the Po Plain

4 Sand dunes and beaches

5 Mountain streams

6 The Mediterranean maquis

Aquatic invertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Davide Tagliapietra Alessandro Minelli Fishes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 Gilberto Gandolfi Terrestrial vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Francesco Bracco Mariacristina Villani

7 Sea cliffs and rocky coastlines

8 Brackish coastal lakes

9 Mountain peat-bogs

10 Realms of snow and ice

11 Pools, ponds and marshland

12 Arid meadows

Terrestrial invertebrates. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 Marco Uliana Alessandro Minelli Terrestrial vertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 Mauro Bon Francesco Scarton Conservation and management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135 Annelore Bezzi Mauro Bon Francesco Bracco Francesca Delli Quadri Gilberto Gandolfi Mariacristina Villani Teaching suggestions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145 Margherita Solari Select bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151 Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 List of species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154

13 Rocky slopes and screes

14 High-altitude lakes

15 16 Beech forests The pelagic of the domain Apennines

17 Volcanic lakes

18 Mountain conifer forests

19 Seagrass meadows

20 21 Subterranean Rivers and waters riverine woodlands

22 23 Marine bioLagoons, constructions estuaries and deltas

24 Italian habitats

Introduction
ANNELORE BEZZI FRANCESCA DELLI QUADRI GIORGIO FONTOLAN

Foreword Italy is a country with more than 7500 km of various types of coastline. From the geomorphological viewpoint, one of the most interesting aspects of these coasts is the relationship between the sea and the rivers that discharge into it, modelling as they do the coastline by Mouth of the river Bevano (Emilia Romagna) distributing freshwater and sediments. The sea acts with tides, waves and long-term fluctuations in level (eustatism). This volume of Italian Habitats illustrates three environments which result from interactions between rivers and the sea: deltas, lagoons and estuaries. The vegetation and fauna of submerged environments are the subject of the first chapters, followed by descriptions of above-water environments. Deltas When a river flows into the sea, the speed of its current is drastically reduced, causing the sediment it carries to be deposited. If the energy of the waves is not sufficient to remove the deposited material completely, an accumulation forms, partly emerging and partly submerged, which is called a delta. This term derives from the triangular shape of many sedimentary bodies of this type, being similar to the capital letter of the same name in the Greek alphabet. In reality, the morphology and dimensions of a delta may vary widely, and in most cases differ significantly from the ideal model of the Nile Delta, which was the original inspiration for the name. Each delta is subdivided into a delta plain, delta front and pro-delta. The plain is a flat area, situated just above sea level, crossed by branching river channels (distributaries) edged by natural banks. Extremely heterogeneous morphological elements meet here that result in the development of diversified habitats. Fluvial processes are prevalent in the upper part; during

Po Delta (between Emilia Romagna and Veneto)

spates, large amounts of fine sediment escape from the channel beds and accumulate on the delta plain; the RIVERbanks of the channels rise, but are also DOMINATED DELTAS frequently breached, with the formation of breach fans and the consequent WAVETIDEdeposition of sand outside the channel. DOMINATED DOMINATED DELTAS DELTAS Processes due to marine dynamics act WAVES TIDES in the lower part of the delta plain and Classification diagram of different delta types the morphologies of coastal types meet, with frequent mixing of fresh and sea waters: lagoons, bays, sandbanks, tidal flats, abandoned strips of beach, and sand dunes. Sediments may be very heterogeneous in this zone, depending on the environment and how they were deposited. The delta front is the shallower part of the submerged delta, mainly composed of sandy sediment. This is the most active part of the deltaic structure, and the most interesting from the morphodynamic point of view. It is here that river water loses speed as it is discharged, and is consequently less capable of maintaining transported sediments in suspension. These then accumulate in structures called mouth bars (loss of competence). The mouth bars assume different shapes according to the hydrodynamic behaviour of the river at the point where it enters the receiving basin, i.e, the sea. The way in which river water expands into sea water, i.e., through turbulent mixing or as a floating body of water, depends not only on the speed of the current and slope of the seabed, but mainly on the density distribution of the water column. This distribution is generally influenced by the seasons, with alternating high and low regimes. The density distribution of the water column may be: homopycnal: when the waters are well mixed, with similar densities both at the surface and on the bottom; hypopycnal: when river water is less dense than that of the sea, because of a low sediment load. It tends to form a surface layer until some distance from the mouth, gradually losing its load of suspended sediments, which settle. This condition is common when there is little water in the river; hyperpycnal: typical of full spate regimes, when river water is denser than that of the sea and tends to move in contact with the bottom. The terminal part of the deltaic structure that develops towards the sea is called the pro-delta. This is the deepest part, originating from finer materials transported out to sea and deposited in layers on the seabed. In cross-section, this sediment accumulation is typically wedge-shaped and represents a
RIVER

significant historical and geological archive. The sediments that form it contain a record of the environmental and anthropic changes that have taken place in the river basin from which the sediments originated. According to a classical scheme, deltas are classified on the basis of shape, which is the result of the combined action of the sediment load carried by the river, the energy of wave motion, and the influence of tides. The Mississippi, in the Gulf of Mexico, is a typical example of a river-dominated delta, in which the rivers contribution dominates the physical modelling operated by the sea. This type of delta develops in conditions of shallow water, where currents, waves and tides are very weak, and therefore in a sheltered gulf or sea into which the river flows carrying large quantities of water and sediments. In these conditions, the delta tends to advance rapidly towards the sea (progradation). The result is a delta plain with a very uneven coastline, with tongues of emerging land with the typical elongated (digitate) shape jutting out into the sea, like a birds foot. These are produced by the rapid seaward advance of the distributaries, confined by natural banks, which erode the sandbars previously deposited at the mouth. In Italy, the most obvious examples of this type of delta are the mouths of the Po and Isonzo. Instead, the So Francisco, in Brazil, is the prototype of the wave-dominated delta, with its classic cuspidate shape. The morphology is characterised by a small number of distributary channels, from the mouth of which a very regular coastline departs, forming two lateral wings. The action of waves and currents

El o

Elo ng for ated m

ng

orm df ate

sp for idate m

Lo b for ate m

Es

ry tua

Cu

Mouth of the river Isonzo (Friuli Venezia Giulia)

10

along the coast prevail over the other components, carrying fine sediments out to sea and redistributing sand along the shore to form the typical beaches. The Italian examples are extremely varied, consisting of almost all the deltas on the Tyrrhenian coast (rivers Arno, Ombrone, Tiber and Volturno) and the Tagliamento on the Adriatic. In points where the discharges are very limited, the river mouth may become obstructed by accumulated sediments, which are only removed on the occasion of full spates. Examples are the mouths of the seasonal streams which discharge into the Ionian Sea in Basilicata (rivers Cavone, Bradano, Basento) and Calabria (Torbido, Precariti, Barruca, Novito), and, in Sicily, the Simeto in the Gulf of Catania. The growth phases of cuspidate deltas are marked by belts of sand or coastal dunes separated by depressions which sometimes contain marshes or pools. In some cases, these survive over long periods of time, and may then be clearly identified at the sides of the active delta as testimony to its evolutionary history. An example is the delta of the Tagliamento, which formed about 2000 years ago, in which eight belts of dunes can be identified, separated by depressions. The Arno and Ombrone share a similar evolutionary history: initially (between about 6000 and 25,000 years ago), they formed their own plains by filling wide lagoon inlets, into which they flowed with internal deltas. These lobes are still visible today on satellite images as drier areas. Once they had filled in their lagoons, they began to discharge directly into the sea only during Etruscan-

Roman times. Their growth was subsequently very rapid (around 7 km in 2500 years), due to sediments arriving from the drainage basins which were rapidly being deforested by man. On the delta of the Ombrone in particular, there are between 12 and 16 main sand belts, some of which are still separated by pools, known as chiari. The history of the evolution of the Po is extremely complex, as the river has been considerably influenced by human interventions aimed at regulation of its distributor channels. According to descriptions by Pliny and Polybius, there were three main distributaries during the 2nd-5th centuries A.D., forming the same number of cuspidate deltas. In more recent times, with a hydrographical network completely modified by man, the so-called modern delta has been formed, now extending for about 730 km2 and more similar in shape to that of digitate deltas. This evolution was caused by the increased sediment transport and raising of the banks of the distributor channels by the hand of man, which has caused their rapid progress towards the sea. The general reduction in river sediment loads during the latter half of the last century has recently led to general erosion of the Italian deltas and adjacent coasts. However, in this respect, it should be emphasised that the shoreline on both sides of a river mouth is, by its very nature, extremely ephemeral and subject to sudden variations both seawards and landwards, in response to high or low discharges of river water and high tides.

11

Mouth of the Allaro, a seasonal stream in Calabria

Mouth of the Piave (Veneto)

12

Types of deltas and estuaries

Isonzo Long sandbars edge the sea outlet of the only distributor channel of the river Isonzo, characterised by a weak regimen of waves and shallow bottoms (scale 1:100,000).

Annelore Bezzi Francesca Delli Quadri Giorgio Fontolan Tagliamento The strong currents along the shoreline have caused the cuspidate delta of the Tagliamento to become asymmetric. The dune cordons that reveal the formation of the nearby land over the last 2000 years, have been obliterated by farming (scale 1:200,000). Cavone The currents that transport sand along the shore partially close the mouth of this small river on the Ionian coast of Basilicata. The emerging sandbars are only eroded when the river is in full spate (scale 1:25,000).

13

Po The modern delta of the Po, which extends for approximately 730 km2, has a similar shape to that of digitate deltas, with various distributor channels flowing out to sea with their loads of sediment (scale 1:500,000).

Ombrone Owing to the lack of urbanisation in the area, the sandy belts of the cuspidate delta of the Ombrone are still clearly visible, equally aligned on either side of the river mouth as testimony to the different phases of growth (scale 1:100,000).

Tacina A typical image of the Ionian coast of Calabria: the meagre discharge of the Tacina seasonal stream is not sufficient to remove the sediment completely, which tends to obstruct the mouth (scale 1:25,000).

14

Estuaries When tides are dominant, river mouths assume a typical longitudinal funnelshaped configuration, often with banks elongated in the direction of the river. The sea may penetrate hundreds of kilometres upstream, and the main physical process is mixing of fresh and salt waters to create estuaries. In the oceanographic literature, the term estuary has a more general meaning. UNESCO uses Pritchards definition: a semi-closed coastal body of water that has free communication with the open sea and within which the seawater dilutes significantly with the freshwater arriving from the surface hydrographical network. In other words, it is the mixing of masses of fresh and salt water that gives rise to estuarine conditions within semi-closed basins. It is thus common to apply the name estuary to internal lagoon areas (such as the Lagoon of Venice and its estuary), where the arrival of a sufficient number of watercourses dilutes the marine characteristics of the basin and generates a marked salinity gradient between the edges of the lagoon and tidal inlets. Following this principle, the salt wedge may periodically intrude into the rivers discharging into the lagoon, extending the estuary environment to the final stretches of these watercourses. The above definition is too generic for any more precise distinction of the different geomorphological contexts, especially in the distinction between

lagoons and estuaries. An estuary might be more properly indicated as the lowest part of a river valley flooded by the sea, subject to tidal fluctuations and the meeting of fresh and salt waters. Seawater, therefore, may enter watercourses or correspond to an area of mixing, depending on how well tidal currents contrast river ones. According to this definition, given the limited tidal ranges of the Mediterranean, it is not easy to identify typical estuaries on a large scale in Italy. The Italian coasts are classified in the micro-tidal category (i.e., a tidal range of less than 2 m), ranges generally being less than 30 cm. This means that, compared with the Atlantic coasts of France and Northern Spain, which have tidal ranges of more than 4 m, the phenomena caused by tides are much less obvious along Italian coasts. There are no large rivers with estuaries in Italy. Smaller watercourses with low sediment loads may form a confined but significant estuary environment in semi-closed basins, examples being the lagoons of Grado-Marano, Caorle and Venice, and the smaller inlets of Scardovari, Canarin and Goro in the Po Delta. An example of a river mouth on the coast of the open sea with estuarine features is the river Magra, which flows into the Ligurian Sea: hydrographical studies of its last stretch have identified stratification, water with salinity comparable to that of seawater being found permanently on the bottom, which reaches as much as 5-7 km from the mouth when the river is low.

15

Mouth of the Mignone (Latium): when the river is low, the water stagnates and does not reach the sea

Area of the Goro inlet, Po Delta (Emilia Romagna)

16

Lagoons In Italy, the northern Adriatic is characterised by abnormal tidal excursions, with average spring tides of 1.1-1.2 m which, occurring as they do along a prevalently low-lying coast and affected by limited wave motion, gives rise to typical morphologies and habitats. These conditions, together with the significant sediment loads of the major Alpine rivers, led to the formation of the large lagoons of Grado-Marano and Venice. Those in the northern Adriatic are therefore the only Italian cases of true lagoons, meaning areas of semi-enclosed sea due to the presence of peninsulas or systems of barrier islands and regulated by the ebb and flow of tides through one or more channels of communication with the sea (tidal inlets). In fact, no stretch of water should be called a lagoon which is sited in environments with practically no tide, such as the Ligurian, Tyrrhenian and Ionian coasts. The northern Adriatic lagoons originated 5000-6000 years ago, when the sea flooded vast alluvial plains, and created large bays bordered by rapidly forming deltas. The sediments from the rivers Isonzo, Tagliamento, Piave and Adige were progressively transported along the shoreline, until they were distributed in the form of lidos, thus separating the large bays from the sea (then regulated by the tidal ebb and flow) and forming the lagoons. In greater detail, according to studies of deep sediments, the central part of the Lagoon of Venice began to form around 6000 years ago, whereas in the northern sector of the lagoon and the delta of the river Adige, the first lagoonal deposits appear to date back to slightly earlier times. The first dune belts of the coastal system of the Piave, further to the north, date back to just over
High tide level Sand bank Channel Shoulder Tidal level Mudflat Tidal creek Levels
High tide, maximum High tide, average Mean sea level Low tide, average Secondary channel Low tide, maximum

17

Thatched fishermens huts in the Lagoon of Grado-Marano

Cross-section of a lagoon

18

Tides

Annelore Bezzi Francesca Delli Quadri Giorgio Fontolan

Tides are caused by the force of attraction that the Moon and, to a lesser extent the Sun, exerts on the Earth. For this reason, the phases of the moon during a lunar month (29.5 days) give rise to significant differences in tidal ranges. During the phase of the new moon, let us imagine that the Moon lies on a line linking the Sun and the Earth and is thus not visible. At full moon, it is entirely visible, because it is situated on the same alignment but on the side opposite the Sun. During these two lunar phases, called syzygy, the attractions exerted by both Sun and Moon are combined, and so the differences between high and low tides are at their maximum (spring tides). Vice versa, when the Moon, Earth and Sun are at right angles to one another, the neap phase occurs (only half the Moon is visible), and the tides have a small range (neap tides).

The Lagoon of Grado-Marano (Friuli Venezia Giulia) during low (top) and high (bottom) tide.
Moon

SPRING TIDE

NEAP TIDE

Earth

lunar days
0 2 2 4

solar

combined

Sun

TIDAL DATA IN RELATION TO PHASES OF THE MOON 6 8 10 12 14 16 18 20

new

quarter

full

SPRING TIDE

NEAP TIDE

5000 years ago. Lagoon sediments found at depths of 6 metres are about 6000 years old. In the sector between the Piave and the Tagliamento, lagoon sediments deposited on the Pleistocene plain are dated to around 5000 B.C. Further east, the marine ingression creating the Lagoon of Marano goes back to Torcello in the Lagoon of Venice (Veneto) between 5500 and 4200 years ago. The Lagoon of Grado appears to have been formed much more recently, as testified by the presence of the Romans in the area, and can be traced to the process of easterly migration of the mouth of the Isonzo. The natural factors contributing towards the formation and maintenance of lagoons are, in the first place, those linked to the hydrodynamic system, which is closely regulated by the tidal action: the hydrographical network of a lagoon is composed of a dense criss-crossing of channels of different widths and depths, which allow the masses of water that enter through the tidal inlets to expand within the basin. Each individual inlet serves a lagoon basin, separated from the adjacent one by a watershed, an imaginary line representing the point of separation between the masses of water entering the lagoon from two neighbouring inlets. Between the watershed of two adjacent basins, the masses of water only mix in exceptional cases (e.g., strong winds), so the waters that flow in through an inlet as the tide rises, drain out through the same inlet as it falls. The network of lagoon channels can be subdivided into the larger main channels, which develop from the area of the tidal inlet towards the interior. They are often the relicts of the ancient fluvial system preceding the marine ingression, and are in some cases connected with the tributary watercourses of the lagoon itself. Secondary channels are smaller and shallower, they branch out from the main ones, and distribute and receive the waters from the surrounding land. During the period of spring tides, when the tidal range is at its peak, the water that flows through the lagoon inlets reaches its highest speeds, even faster than 1 m/s, and rapidly spreads through the main channels and then through the secondary ones, until it reaches the tidal flats. As the system of channels gradually multiplies, water speed falls to a few cm/sec. When the water reaches the tidal flats, the speed of the current again sharply diminishes, due to lateral

19

livello dellacqua (m)

20

Water exchange in a lagoon occurs through small channels (tidal creeks), mainly during high tides

expansion of water masses. This process is repeated in the opposite direction during the ebb tide: the main channel acts as a collector, all the secondary channels distribute on the tidal flats, and the lagoon waters flow out towards the sea, reaching their maximum speed at the tidal inlets. This is the hydrodynamic model which is responsible for the arrival, transport and deposition of sediments inside the lagoon, giving rise to the various morphologies found in the basin. The typical morphologies that characterise the northern Adriatic coastal lagoons are grouped in three morphological zones, according to Sandbanks may be stabilised by vegetation (Zangheri sandbank meadow, Emilia their position with respect to sea level: Romagna) the shoals, islands, beaches and internal coast of the lagoon are above the average level of high tides; the main channels and tidal inlets are below the average level of low tides; the intertidal areas, i.e., those lying between the average levels of low tides and high tides, which are periodically submerged. Typical examples are the tidal flats, i.e., the shallow lagoon beds and secondary channels, which emerge when low tides discharge water towards the sea. In lagons, islands are portions of land never submerged by high tides. They may constitute relicts of ancient beaches, or have originated from the accumulation of sediment transported by rivers. Shoals are the most characteristic landforms of lagoon environments. They are tabular raised areas, mainly composed of silty-clay sediments, which are only submerged by water in exceptional cases. Shoals have irregular topography and may contain saltpans, which receive an exchange of water through small, tortuous channels (tidal creeks), especially during high tides. Because of the complexity of the processes causing the formation of lagoons (distribution of freshwater and materials from rivers, marine ingressions, and later drops in water level), shoals are classified on the basis of their different origins: shoals of ancient river edges, or ones located at the sides of watercourses which still discharge into the lagoon, with a characteristic long, narrow shape;

21

22

shoals composed of the still emerging part of an ancient coastal plain invaded by brackish waters: they are located on the edge of the lagoon towards dry land, and their sediments contain a thick layer of deposits of continental origin, overlain by a thin layer of lagoonal type; residual shoals of a marshy environment, with layers of peat in the sediments: they were formed by river alluvial deposits in the lagoon basin which, following emersion, became covered by marsh vegetation. After river waters had been diverted they no longer received sufficient amounts of sediment to impede their rapid sinking, mainly due to compaction of the peat; channel shoals: these are in equilibrium, with sufficient sedimentation to compensate for slow sinking. Their shape is typically concave, with a lowerlying central area; the side bordering the channel is generally steep, whereas the other slopes gently towards a marsh or tidal flat. Channel shoals form at the points of convergence of lagoonal flows and in the zones of watershed where, in ideal conditions, the resulting water velocity is almost zero. These sandbanks grow in height because the waters that flow over them during high spring tides, slowed by vegetation, release part of their solid load, and expand due to the slowing down that sea currents of different intensity and direction exert reciprocally. For some authors, this last type constitutes a shoal in the strict sense - the result of the deposition of sediments arriving exclusively from the sea.

Mouth of the Agri (Basilicata)

Tidal flats are lagoon beds that only emerge at low tide, especially spring tides, and are not therefore suitable for colonisation by pioneer plants. They slope very gently (with gradients of only a few dozen centimetres per kilometre) and have a system of secondary channels discharging towards the main channel. These channels are meandering and a few metres wide; their minimum level coincides with the average of the low spring tides, so their maximum depth (slightly more than a metre) depends on the tidal range and coincides with the point of confluence with the main channel. Sandbanks, mudflats and channels in the Lagoon of Grado-Marano Lastly, some of the most typical environments in Italian lagoons are the fish-farm basins, exploited since early times for rearing and capturing fish. These often original basins are generally surrounded by embankments that separate them from the main lagoon, in order to protect them from the effects of tides and to regulate the entry of brackish and freshwaters from the hinterland. They currently occupy vast portions of the Lagoon of Venice and that of Grado-Marano and, to a lesser extent, those of Caorle and Caleri. As regards sediments, there is generally a progressive reduction in particle size from the tidal inlets towards more internal areas. These variations are usually associated with the energy level of the environment: lower where the basin is deeper, more sheltered and further from the sea; higher close to the tidal inlets, in the immediate vicinity of the channels, and in areas where the water is shallower and the bed is affected by the wind, waves and sea currents. Sandy sediments are deposited near the tidal inlets, often forming characteristic morphologies known as flow-tide delta (on the lagoon side) and ebb-tide delta (on the seaward side). Sand becomes progressively finer along the main channels and further in, especially on the tidal flats; fine sand is progressively replaced by mud, which becomes dominant towards the interior of the lagoon, to become exclusive, in its finest fraction, closest to the internal lagoon coast, as well as to the watershed.

23

Submerged vegetation
DANIELE CURIEL ANDREA RISMONDO

25

Foreword In optimal environmental conditions, the distribution of brackish water vegetation differentiates between that part of the body of water directly influenced by the sea and the more landward one, which is affected by confinement of waters or sometimes by freshwater. With distance Chaetomorpha from the points of exchange with the sea, submerged vegetation becomes simplified, because, as the type of sediments alters, from prevalently sandy to silty and becomes more subject to re-suspension, the morphological and physico-chemical characteristics of the water also change. Most of the phytobenthos of lagoons is typically of marine origin, with species that are capable of tolerating marked reductions in salinity. This dominance of marine species is also due to the fact that the typical organisms of internal waters are rarely capable of adapting to salinity higher than 5. When the arrival of freshwater is limited, salinity becomes less important for regulating the presence and abundance of submerged vegetation, whereas water transparency, sedimentation, hydrodynamics and the availability of nutrients become more so. As these environments are shallow (sometimes less than 40-50 cm), water turbulence due to weather - or human activities, such as mollusc-farming - may easily trigger events of re-suspension that reduce water transparency and limit compaction of sediments. When dystrophic conditions change with season, few variations occur in the structure of the communities, and the resilience of the plant communities allows typical conditions to be restored within a reasonable time. If dystrophic conditions persist for more than one year, submerged vegetation substantially modifies, with the gradual substitution of species with long life-cycles and high ecological value (such as seagrasses, brown algae like Cystoseira barbata and Fucus virsoides) by species with short life-cycles (such as the Ulvaceae or Cyanophyceae).

Sargassum in the Lagoon of Venice

26

The macroalgae: green algae Compared with the marine environment, where red algae prevail, green algae are the best-represented plants in the transition zone, especially in terms of abundance. These algae are generally euryecious and find optimal conditions in the shallows of internal and confined areas where water exchange is limited and nutrients are plentiful. In the transition environments of the northern Adriatic, the most abundant and best-known alga is Ulva laetevirens, with its widened leaf thallus, commonly known as sea lettuce. Other species of the genus Ulva (U. rotunda, U. curvata) or Gayralia oxysperma (morphologically similar but with a single layer of cells in the lamina) never reach the distribution and biomass levels of U. laetevirens. Another very widespread algal component is that of the Enteromorpha species, a group of green algae recently unified with the genus Ulva. These green algae, with tubular simple or branched thalli that may measure from a few centimetres to 50-60 cm, live in eutrophic environments. The most common species include U. intestinalis, U. clathrata and U. flexuosa. In summer months, the gases present in the cavities and products of photosynthesis raise the thalli from the bed, forming drifts floating on the surface of the water. During the summer, especially in the internal waters of fish-farm basins or in association with seagrasses, uniform carpets of the green filamentous algae

Ulva laetevirens

Chaetomorpha aerea and C. linum (perhaps conspecifics) are very frequent. In the summer months they are often mixed with green algae of the genus Cladophora (e.g., C. sericea, C. albida, C. rupestris, etc.) In the 1980s and early 1990s, all the lagoons in the northern Adriatic were afflicted by blooms of these green macroalgae, the The green alga Valonia aegagropila prevalent species in terms of extent and biomass being Ulva laetevirens. Recorded since the 1940s-50s in muddy areas of the Po Delta, in the Valli di Comacchio and Lagoon of Venice, but today much rarer and limited only to hollows, are the subspherical-shaped thalli, 3-8 cm long, of the green alga Valonia aegagropila. These macroalgae may form extensive, uniform, continuous coverings up to 30-40 cm thick, and with a wet biomass of more than 10 kg/m2. In fish-farm basins where salinity is maintained constant at values slightly above 10, partly in order to favour hunting, green algae belonging to the genera Chara and Lamprothamnion (L. papulosum) are found on the bottoms, mixed with the seagrasses Nanozostera noltii or Ruppia (R. maritima and R. spiralis) or with the green alga Valonia. These macroalgae are often not taken into consideration by marine algologists because they are only found in transition environments and adjacent flowing waters. Coherent substrates in these brackish environments host a few macroalgae which would have difficulty in living on loose substrates because of their need to anchor themselves to something solid. Both in vivified areas and in internal areas close to the transition to brackish waters, on the highest levels which are only wet during high tides (at the limit between the intertidal and supralittoral areas), are green algae of the genus Blidingia (B. minima, B. ramifera), which are resistant to emersion. The height of the tides and wave motion caused by shipping may raise the colonised level by up to 20-30 cm. Below this, starting from average sea level, in the intertidal zone, a filamentous distichous green alga (Bryopsis plumosa) may be observed, followed, but only in vivified environments like the Lagoon of Venice, by the green alga Codium fragile subsp. tomentosoides, with its cylindrical dichotomous branching thallus. This alga, a native of the Pacific Ocean and Asiatic seas, has been present in the northern Adriatic since the 1970s-80s. Although viewed as one of the most invasive algae, because of its ability to adapt to differing conditions of salinity and temperature, only isolated thalli are found, at least in the Lagoon of Venice.

27

28

Alien species
For more than a decade now, several species endemic to other seas have been found on both soft and firm substrates in transition environments in the Mediterranean. They have arrived thanks to the voluntary and involuntary actions of man. Until the Suez Canal was opened in 1869, the introduction of new species into the Mediterranean occurred accidentally through the Strait of Gibraltar. With the opening of the Suez Canal, the Red Sea and the Mediterranean, having remained separated for more than 10 million years, were put into communication again. For hydrological and ecological reasons, migrations are prevalently towards the Mediterranean. In the last 20-30 years, this slow but continual introduction of species has become insignificant compared with the much faster pace with which man has favoured the introduction of new species along the coasts of the Mediterranean. Transition environments, historically the sites of important economic activities (fishing, shipping and port structures, etc.), are consequently the main points at which new species arrive. If they adapt to the new environmental conditions, they can spread until they eventually reach an equilibrium with native species. The spread of others, which do not find optimal conditions in the new environment is limited, or they disappear. In the Mediterranean, the two most striking examples of transition environments affected by many alien species, are the Lagoon of Thau in Southern France and the Lagoon of Venice. In the latter, more than 20 species have been recorded on the substrates off Chioggia and Venice since the early 1990s, and, because they are found outside their potential distribution area, they are correctly defined as alien species

Daniele Curiel Andrea Rismondo or NIS - non-indigenous species. There have only been a few records for some of them (e.g., Sorocarpus sp., Prasiola sp.), probably because they have not adapted to the new and complex lagoon conditions. Instead, others have spread over the years and become integrated with native species (e.g., Antithamnion nipponicum, Grateloupia turuturu, Codium fragile subsp. tomentosoides). Yet others have spread rapidly and become dominant in the macroalgal community, with widespread diffusion and high abundance (Sargassum muticum, Undaria pinnatifida, Polysiphonia morrowii). A very unusual case, which now involves various lagoons of the Mediterranean, the Atlantic coasts of France and the northern seas, is that of the two brown algae, Sargassum and Undaria. Although having different lifecycles - Sargassum being perennial and Undaria annual - in the climate of the Northern Adriatic these two species have thalli observable from October and November, which disappear between June and July. Attempts have been made to eradicate them in various locations, so far with little success. Both species have extremely efficient dispersal systems for their millions of spores, which can be taken great distances by tides - and even fragments of thalli have high viability.

Red algae As in the marine environment, red algae are a numerically conspicuous part of the aquatic vegetation in transition environments, especially in those more highly vivified, like the lagoons of Venice and Grado-Marano. Being ecologically sensitive, especially to any reduction in salinity and increase in turbidity (natural or induced), red algae are limited in the fish-farm ponds, the inlets in the piallasse (marshy areas) between Romagna and the Veneto or towards the interior shores of the large lagoons of Venice and Grado-Marano. Only more tolerant species, which can Gracilaria adapt to waters rich in nutrients and sediments arriving from the rivers, can survive. On loose beds, the species typical of brackish water include those of the genus Gracilaria (G. bursapastoris, G. armata and G. gracilis). Gracilariopsis longissima (previously known as Gracilaria verrucosa) formed extensive macroalgae covers in the 1980s, alone or in association with Ulva. Today, the phenomenon of algal bloom appears to be more limited in the two large lagoons of the northern Adriatic, but reappears almost annually in the small to medium-sized transition environments in the Po Delta, although less intensely than in the past. With currents and winds, the thalli of Gracilaria / Gracilariopsis roll around on the bottom and accumulate in sheltered areas, where they may proliferate. In summer months, in the Lagoons of Venice and Grado-Marano, with their reduced hydrodynamics, more or less extensive covers of Spyridia filamentosa are to be found. Mainly in the spring and summer months, on loose bottoms without macroalgal cover, natural coherent substrates such as the valves of molluscs (Tapes, Cerastoderma, Cassostrea or serpulid tubes), or artificial substrates (stones, shingle) favour the presence of red algae with pulvini (cushion-like enlargements), 5-10 cm tall. Among the most eye-catching are the branched filamentous forms of Ceramium (the dark-red C. virgatum, or C. diaphanum, with alternating light

29

Undaria

30

and dark bands) and the Antithamnion (A. cruciatum, and, in the Lagoon of Venice, A. nipponicum, a native of Oriental seas). Among the branched laminar forms, Nitophyllum punctatum and Radicilingua thysanorhizans are especially striking. Conspicuous numbers of red algae are also to be found on the leaves of seagrasses, especially Cymodocea nodosa. On the oldest (external) leaves, a rich community of medium-small species develops, including the filamentous forms Ceramium, Antithamnion, Chondria capillaris or Chondria dasyphylla, and various species of the genus Polysiphonia. The red encrusting coralline algae of the genus Hydrolithon and Pneophyllum fragile may also be abundant, especially in the months of scarce or no growth of leaves. In the Lagoon of Grado-Marano, but to an even greater extent in that of Venice, there are many artificial coherent substrates (stones, shores of the islands, signalling devices for shipping, etc.), where species typical of these substrates may be observed. Above the average tidal level, Gelidium pusillum and Gigartina acicularis are frequent, with their branched filamentous thalli firmly anchored to the substrate and able to resist emersion for some hours. Below the average tidal level are species of the genus Polysiphonia (P. morrowii, a species native to the Oriental seas, P. harvey and P. denudata) and, lower down, Rhodymenia ardissonei, a sciophilous (shade-loving) alga, indicative of the lack of light in these environments caused by water turbidity.

Brown algae Brown algae do not find ideal conditions for growth in transition environments. On the coherent substrates of the two largest lagoons, species of ecological value can only be found in areas more affected by seawater distribution, with Fucus virsoides (a species of boreal affinity, present in the Mediterranean only in the northern Adriatic), Cystoseira barbata or Petalonia fascia. Dictyota dichotoma var. dichotoma and Scytosiphon lomentaria may also be observed in more internal areas. The microscopic species of the genera Myrionema, Leptonematella and the Fucus virsoides filamentous forms Ectocarpus and Hinksia are typically found on the leaves of the seagrasses. The latter are also the only brown macroalgae which, in the presence of nutrient availability and reduced hydrodynamics, may also flourish on loose substrates. In the Lagoon of Venice, this occurs mainly amongst the seagrasses when, following their summer proliferation, the filamentous thalli of Ectocarpus become entangled with the seagrass leaves, forming larger and larger clumps. Two brown algae, Undaria pinnatifida and Sargassum muticum, which are endemic to Far Eastern seas, have been found on artificial solid substrates in the Lagoon of Venice since the early 1990s. The large size of the thallus and the abundances these species are able to reach render them highly competitive with native species. Mention should also be made of the algae of the genus Vaucheria. In the deep beds of the lagoons of Venice and Grado-Marano, there has been a proliferation of these macroalgae in the past decade; unlike all the other species, they do not need a firm substrate to anchor themselves because they grow partly immersed in sandy-muddy sediments. Vaucheria dichotoma var. marina, today identified as V. submarina, is more common and has been recorded since 1800, but V. piloboloides has also recently been found in the Lagoon of Venice. These macroalgae play an important role in compacting the sediment layer and greatly reducing re-suspension of sediments during periods of water turbulence.

31

Polysiphonia morrowii

32

Phythoplankton
The highly variable environmental conditions of brackish-water environments, in terms of salinity level (10-40), temperature (sometimes exceeding 30 C), turbidity (Secchi disk values up to 10-30 cm) and dissolved oxygen (from anoxic to super-saturated conditions), mean that only phytoplankton with a wide ecological range and high quali-quantitative variability during the course of the year can survive. As well as abiotic factors, an important role is also played by biotic factors, such as the presence or absence of herbivorous predators. For clearer understanding of variability over the year, in the Lagoon of Venice in the early years of the 21st century there have been minima of 1x105 cells/litre in winter and peaks of 19x106 cells/litre in spring-summer. The most abundant taxa in all transition environments are the Bacillariophyceae and Pyrrophyta, more commonly known as diatoms and dinoflagellates. Diatoms have a silicified cell wall (frustule) formed of two interlocking valves. In the Lagoon of Venice, the most common and abundant diatoms belong

Daniele Curiel Andrea Rismondo to the genera Chaetoceros, Cocconeis, Nitzschia, Thalassiosira and Skeletonema and, among the forms which constitute mucilaginous tubules and are also epiphytes of seagrasses, Navicula and Melosira. Species of the genera Prorocentrum and Gyrodinium are frequent among the dinoflagellates. In the Lagoon of Grado-Marano, the diatoms of the genera Synedra, Navicula and Rhizosolenia are frequent and abundant. Among the dinoflagellates are Exuviella, Gymnodinium and Prorocentrum. In the brackish-water lagoons of Emilia Romagna, diatoms of the genera Nitzschia, Thalassiosira, Skeletonema and Melosira are common. Of the dinoflagellates, those of the genera Prorocentrum, Peridinium or Glenodinium have often produced large-scale algal blooms. The microalgae belonging to the chlorophytes (green algae) are less abundant, but no less important. They include species belonging to the Volvocales (Dunaniella being typical of hyper-saline basins) and Chlorococcales (with Chlorella).

Seagrasses Lagoon beds may be covered by varying densities of meadows of seagrass, a small but important group of flowering plants which are fully described in the Italian Habitats volume Seagrass Meadows. Posidonia oceanica, Zostera marina, Nanozostera noltii, Cymodocea nodosa and two species of the genus Ruppia are present in the Adriatic. The success of these macrophytes in the underwater environment of lagoons is due to the various morphological and physiological adaptations which they have had to develop to survive so well. Posidonia oceanica These include adaptation to the salty environment and to submersion (hydrophytic habitus), the capacity to resist the action of waves and tidal currents, and hydrophilic pollination and dissemination.

33

SPECIES Posidonia oceanica

GEOGRAPHICAL DISTRIBUTION Gulf of Trieste, including Grado coast Western Istrian coast Venetian coast (dead mattes) Western Istrian coast Gulf of Trieste Lagoons of Venice and Grado-Marano Lagoon areas of Po Delta Istrian bays and sheltered areas Western Istrian coast Gulf of Trieste Lagoons of Venice and Grado-Marano Lagoon areas of Po Delta

Zostera marina

Nanozostera noltii

Cymodocea nodosa

Istrian bays and sheltered areas Lagoons of Venice and Grado-Marano Scattered along lidos of Jesolo, Eraclea and Caorle Shoal and drainage areas of northern Adriatic lagoons in general

Ruppia sp.pl. Nitzschia (approx. 2000x) Cocconeis (approx. 2000x)

Seagrass distribution in the Adriatic Sea

34

Nanozostera noltii

The main factors regulating the presence and distribution patterns of seagrasses on the beds of coastal lagoons are, in order of importance: particle-size distribution of sediments, water turbidity, temperature patterns, local hydrodynamics and resulting water exchange, eutrophication and the consequent presence of macroalgae. The quality of waters and sediments and the hydro-morphological factors of basins tend to favour one or other of the existing species. In the Adriatic coastal lagoons, Ruppia is present in a variety of microenvironments. These cannot be precisely characterised in terms of Ruppia salinity, because the genus is to be found not only in the fish-farm ponds of Venice and Grado-Marano and in some habitats with very low salinity, but also in pools nearer the sea, which are subjected to high salt concentrations. In the northern Adriatic lagoons, Nanozostera noltii is a typical species of marshes and tidal flats, the more internal and confined areas subjected to cycles of emersion and with lower salinity than that of the sea. However, in the larger and morphologically more complicated basins, such as the Lagoon of Venice, the species may have more than one phenotype, sometimes colonising tidal flats with low, dense carpets and thus ensuring the morphological protection of these systems. At other times it forms extensive meadows on the deeper beds, where it grows much larger and with longer leaves. In the northern Adriatic, it is substantially homogeneous, being found both in the Lagoon of Venice and in that of Grado-Marano, on the same morphological systems and therefore on tidal flats and on the edges of deeper channels, on clearly silty beds. Very rare and extremely localised in the other, smaller, Adriatic basins, it has been reported close to the sluices or entry channels of waters such as those at Comacchio. Since the late 1990s, N. noltii has been undergoing a highly generalised process of retreat from the areas of sandbanks - particularly marked in the Lagoon of Venice. This has many causes - such as the general increase in

35

36

turbidity and prolonged periods of emersion during very hot sunny periods but the intensified erosion processes, especially in the lagoons suffering from severe human pressures, should not be underestimated. The larger Zostera marina, with leaves 6-7 mm wide and up to a metre long, lives almost always submerged in more turbulent and less confined waters, preferring bottoms with silty sand. Its distribution area covers truly lagoonal waters, as in the lagoons of Venice and Grado-Marano, where it forms extensive meadows. In the Po Delta and the brackish lagoons along the Romagna coast, its populations are rarer and more scattered, mainly in the channels linked with the sea. However, even truly marine waters cannot be excluded, because along the coast east of Grado, near the mouth of the river Primero, which has no artificial protection and is characterised by entirely natural beds, there is a whole alternation of sandbars and depressions which is widely colonised by Z. marina. Alternating with Cymodocea nodosa, it most probably takes advantage of the mixed sandy-silty sediments and the presence of discharging freshwater. Zostera marina is the most euryhaline of the species described here, tolerating salinity of between 5 and that of seawater; for this reason, paradoxically, it may spread towards the coastline, and in any case in the directions of flows of freshwater, more than Nanozostera noltii can.

In the lagoons of Venice and Grado-Marano, N. noltii forms large vegetated beds that extend from vivified sites to others which are partly confined. It avoids areas subject to water stagnation, as the upper limit of the favourable temperature range for this macrophyte is around 25-27C. Cymodocea nodosa, with a dense, robust root system, is found close to inlets where sediments are predominantly sandy and water exchange is medium to high. A stenohaline species, it cannot resist conditions of high turbidity and, although it grows at depths of up to 10 metres along the Istrian coast, it is not found deeper than 4 metres near the lagoon inlets, on occasional delta sandbars and in the other Adriatic coastal lakes like those in Apulia. C. nodosa is a species of sub-tropical origin and is highly seasonal, especially in the northern Adriatic lagoons, where the temperature may drop below zero in winter and rise above 30C in summer. In winter, the macrophyte uses a strategy of conservation of the root system and hypogeal (buried) stalks, where sugar is stored, and this may lead to the loss of all its leaves. Only with the marked rise in temperature, usually in May, is there rapid and intense re-growth which, in the space of a few days, gives rise to dense new meadows of brilliant green. At the inlets, on coastal sandbars, and in the brackish coastal lakes of the southern Adriatic, C. nodosa retains its leaf apparatus in the winter months, although it may be reduced, degraded and covered by epiphytes.

37

Foglie di Zostera marina

Cymodocea nodosa

Aquatic invertebrates
DAVIDE TAGLIAPIETRA ALESSANDRO MINELLI

39

Deltas, estuaries and lagoons are transitional coastal ecosystems characterised by gradients of many important environmental factors, such as water exchange, salinity, sediment structure, turbidity and nutrient load. The transition gradient is generally perpendicular to the coastline, from the sea landwards, with consequently progressively changing habitats and biological communities. Variations in biodiversity
Mouth of the Idume near Lecce (Apulia)

Gradually penetrating a transitional ecosystem, there is a progressive decline in the number of species. This phenomenon occurs in both directions - from the sea towards the land, and vice versa - and biodiversity reaches its minimum in the area where river and sea meet. The accumulation of organic matter in areas subject to slow exchange and hydrodynamics plays a key role in modulating the availability of oxygen, which becomes a strongly limiting factor that increases the vulnerability of biocoenoses. In Mediterranean lagoons, where the contribution of freshwater is lower and the seawater is less diluted, the reduction in the number of species along the sea-land axis is mainly attributed to the hydrology of the basins and sediment properties, and only secondarily to salinity. Tides greatly accelerate these exchanges, and their cyclical nature generates characteristic models of time-space distribution. The periods of emersion and submersion dictated by the tidal regime are other important factors affecting biological communities in transition environments. Most of the species which live in these areas are of marine origin, so that the degree of connection with the sea greatly influences the recruitment of species which require a larval stage involving marine dispersal, and this has

Cerastoderma shells

40

Detail of the shell of Acanthocardia spinosa, a bivalve

repercussions on the biodiversity and structure of the communities. It is therefore to be expected that, from the sea landwards, the progressive divergence from marine environmental conditions are tolerated by fewer and fewer species. The opposite situation occurs for species of river origin. Towards the river mouth, there is a progressive departure from typical river environments and the number of species tolerating the new environmental conditions gradually falls. This transition from a continental to a marine environment in deltas, estuaries and lagoons combines these two-way changes in biodiversity from the sea towards rivers and vice versa.

Classification zones and their fauna A salinity-based empirical method to classify coastal and transitional waters was proposed in 1958 after the IUCN Venice Symposium, an event at which the major international experts met to agree on a system of classification. The following zones were identified: hyperhaline, with salinity >40; euhaline 4030; mixohaline 30-0.5; and limnetic 5-0.5. As most transitional environments fall within the wide mixohaline zone, the latter was further subdivided into: mixo-euhaline with salinity >30 but lower than that of the adjacent sea; (mixo-)polyhaline, 30-18; (mixo-)mesohaline, 18-5; and (mixo-)oligohaline, 5-0.5. However, zoning based on salinity is extremely variable over time and is only based on one of the factors of the transition gradient. Instead, physiographical zoning is based on the most obvious morphological and structural characteristics of the environment, is less variable over time, and may be integrated, where needed, with the Venice System. The broad physiographical zoning of a typical transition environment such as a coastal lagoon is as follows. Moving from the sea towards the river, a lagoon basin is divided into three large sections. The first is the zone of the actual inlet in which the seas

energy is increased. Here, both inside and outside the inlet, the sediments, composed generally of sand or silty sand, fan out to form tidal flats. Channels then lead to a second zone, the central basin, which is generally rather shallow. This is where the energy from both sea and river is lowest and abundant deposits of fine sediments are found. As the central basin forms the most extensive part of the lagoon, it is not homogeneous and slopes gradually towards more internal shallows, with sandbanks along the edges of the channels. The complex interplay of tidal energy, river energy and hydrodynamics in general decides how close the sandbanks approach the sea, reducing areas of open water in the central basin. Marshland and sandbanks succeed one another towards the lagoon edge as far as the third large zone, composed of the mouth of the river. Here we find sandbanks, first mesohaline and then oligohaline, dominated by reed-beds, which are progressively replaced by less salty environments, until the true river environment is reached. It should be noted that drainage operations in almost all Italian lagoons have greatly reduced the zones of lagoon inlets and oligohaline marshes. In a delta dominated by tides, river energy is such that the central basin disappears and, with it, the capacity to trap fine sediments, which are thus deposited further inland, past the line of the mouth.

41

Lagoon of Venice, with the typical bricola, signalling deep-water channel for shipping

42

The crab
The common crab or green crab Carcinus aestuarii (= C. mediterraneus) is a decapod crustacean characteristic of the transition and coastal environments of the Mediterranean and the Black Sea. However, its status as a species is under discussion - according to some authors, it is merely a subspecies of Carcinus maenas, the affine and vicariant form (i.e., occupying the same ecological niche) which lives along the Atlantic shores of Europe. The species is well-adapted to the high spatial and temporal variability of environmental conditions in lagoons, estuaries and deltas. It is able to tolerate salinity values higher than those typical of the sea or ranging as low as an oligohaline state, and temperatures of between approximately 0C and 30C. The wide ecological range of the crab is confirmed by its omnivorous feeding habits, although predatory and detrivorous behaviour prevails. In conditions of high density, the common crab can regulate the abundances of its prey species (bivalves, polychaetes, and other invertebrates), significantly influencing the community structure. In turn, it constitutes an important food resource for limicolous birds and estuarine and sea fish such as eels, gobies and sea bass. Like all arthropods, the crab must moult several times during its lifespan, replacing the outer shell or exoskeleton, to allow for growth (see drawing). This process is divided into four stages: premoult; true moult, which only lasts for a few hours; post-moult, and inter-moult, the stage of resting and preparing for the next cycle. In general, adult males (m) moult in spring and autumn, whereas females (f) undergo only the first moult, later than the males. Mating takes place during this latter stage (a). The females

Marco Sigovini

43

II III IV

XII XI

f m

a
V VI VII VIII

IX

The life-cycle of the crab. On the left, a diagram showing moult phases - dashed line: periods of the moult; continuous line: inter-moult (for abbreviations, see text)

Carcinus aestuarii

lay their eggs between autumn and winter, depending on location. In many Mediterranean estuaries and lagoons, female crabs migrate towards the sea in winter, where the eggs then hatch. This is followed by a planktonic larval stage, during which stages called the protozoea, zoea (A) and megalops (B), follow one another. At the end of the last stage, the larvae, having returned to the estuarine or lagoon environment, settle down as benthic organisms, preferably in protected habitats like seagrass meadows, where they find shelter during their juvenile stage (C). From this moment onwards, they will moult several times a year, with lessening frequency, until reaching sexual maturity (D). Exploitation of crab meat as a human food resource is only of importance in the northern Adriatic, where a traditional type of fishing exists, principally at

Burano and Chioggia, of high ethnological and economic interest. Its targets are the moulting crabs, called moleche, or moeche, which are immediately removed from the water to stop re-calcification of the exoskeleton. Detailed knowledge of the crabs lifecycle is extremely important in the process of selection, and the molecanti, as these fishermen are called, have a whole series of names for the crabs during their various stages: gransio bon, an individual that still has to moult; gransio mato or duro, specimens which will not moult again during the fishing season; spintano, close to moulting; capelo, harvested at the moment when the shell detaches; and mastrzzo or struzzo, a few hours after the moult. Females with mature ovaries (masante) are also considered as delicacies and are gathered in autumn for the market.

44

Benthic fauna In analogy to the succession of different types of bed, which from sand become progressively silty sand, silt and then clay, populations of benthic macroinvertebrates evolve - from the communities classically attributed to the biocoenoses of fine sand in tidal Nassarius mutabilis deltas, to fine surface sand and muddy sand in environments sheltered from the more open areas of the central basin, until we reach the euryhaline and eurythermal lagoon biocoenoses of the more internal parts. We will therefore start from an imaginary inlet in a tidal delta, first on the seaward and then on the landward side. In the area facing the inlet, we find species typically connected with substrates with a high sand content and more turbulent waters. Some species of bivalves are well-known in fish markets, like the clam (Chamelea gallina) and the razor shell (Ensis minor), whose elongated shape allows it to bury itself rapidly in the case of danger. Tellins (Tellina pulchella, T. nitida, T. tabula, Donax semistriatus and D. trunculus) are also extremely common. Frequenters of beaches are certainly familiar with Acanthocardia tuberculata and A. echinata, bivalves with a lovely heart-shaped shell, the surface of which is characterised by radial longitudinal ribs with more or less pronounced tubercles, marked by deep inter-rib furrows. These species are accompanied by the miniscule Lentidium mediterraneum, millions of which often remain stranded at low tide, rapidly destined to enrich the fine shell detritus on the shore, the white Spisula subtruncata and the larger Mactra stultorum. It is also easy to identify some of the gastropods, such as the murex (Bolinus brandaris), together with its cousin Hexaplex trunculus, an active predator of bivalves whose spawn, like small white sponges, are often beached along the shore, and Nassarius mutabilis, a small ventricose snail which acts as a scavenger, feeding mostly on dead animals. Some families of molluscs arouse particular interest, in that species belonging to them succeed one another in a characteristic manner along the transition gradient. This is the case of venerid bivalves, mainly represented here by Chamelea gallina, and nassariid gastropods, represented by Nassarius mutabilis. Polychaetes include Owenia fusifromis, a tubicolous filtering worm, which lives in the sediment in a tube it creates by cementing grains of sand, and Arenicola

marina, which is responsible for the mounds of sand with the appearance of thick spaghetti that may be seen on sandy silty beds at low tide. This worm is extensively used as bait by fishermen and can tolerate sometimes considerable reductions in salinity. Another interesting polychaete is Sabellaria spinulosa, whose colonies have recently begun to cover hard substrates with a compact layer of tubes constructed with cemented sand, giving the overall appearance of a beehive. These polychaetes are important in that they create new substrates for other organisms. Among the crustaceans are some species of portunid crabs with moderate swimming ability belonging to the genus Liocarcinus, like the sand crab (Liocarcinus vernalis). Close to the shore lives the crab which might be defined as the king of the lagoon, Carcinus aestuarii (see box). Among the nooks and crannies of breakwaters and outer sea walls lurks what is probably the most robust of the Mediterranean crabs, the yellow or warty crab (Eriphia spinifrons), accompanied in the intertidal zone by the very fast-moving Mediterranean shore crab (Pachygrapsus marmoratus). The echinoderms include Echinocardium cordatum and Schizaster canaliferus, two heart-shaped sea urchins which live buried in sediment. Rocky substrates are home to Paracentrotus lividus, the classical sea urchin that causes so much pain to careless bathers descending the cement blocks often used to create breakwaters.

45

Sand crab (Liocarcinus vernalis)

46

Echinocardium cordatum

The isopod Tylos latreillei passes daytime hours buried in dry sand, usually a long way from the foreshore; at sunset it comes out into the open on to the wet beach, where it feeds on various organic materials: beached marine animals, algae, etc. Before dawn, it is once again resting in the dry sand. Regarding this species, worthy of mention here is the mass migration observed on Volano beach on the evening of 10 August 1968 by Antonio Mussels (Mytilus galloprovincialis) Giordani Soika, an attentive and untiring scholar of the natural history of the Veneto lagoons and the Po Delta. He saw millions of individuals, composing an almost compact column a dozen metres wide, marching on the damp sand along the shore, in a southerly direction. Many of these species enter the lagoon and settle in internal areas in tidal flats. Here the razor shell (Solen marginatus) replaces Ensis minor, while Chamelea gallina is less frequent, being flanked by other venerids like Dosinia lupinus and Paphia aurea. The Philippine clam (Tapes philippinarum) starts to put in an appearance here, but is much more common in the next zone. This species, introduced into the northern Adriatic in the last twenty years, has the typical characteristics of invasive species, such as a high reproduction rate and great tolerance to estuarine and lagoon environments. Unfortunately, very destructive systems are used to harvest it. As well as the Philippine clam, other exotic species introduced by man have become established in these environments in particular, the bivalve Scapharca inaequivalvis and the gastropod Rapana venosa. The spread of Scapharca is easily explained by the fact that it is resistant to anoxia, being able to rely on haemoglobin as a respiratory pigment. The large gastropod Rapana venosa, of Indo-Pacific origin, causes problems, since it is a predator which grazes on mussel and oyster beds. Its flesh is edible, but not greatly appreciated. The extremely common mussels (Mytilus galloprovincialis) find their ideal lagoon habitat in this zone. Indeed, this is where artificial mussel-beds are set up, prior to being moved some miles offshore, where the contribution of brackish water stimulates the growth of phytoplankton and where environmental conditions during the critical summer and winter periods are more stable, guaranteeing a longer growing period.

47

48

Nassarius mutabilis declines in abundance, being replaced by other nassariids such as Nassarius corniculus, N. nitidus and, to a lesser extent, Cyclope neritea. Nassariids pass their time buried in the sediment, leaving only their siphons protruding, and with these they monitor the water, waiting to detect the smell of the dead or injured animals on which they feed. Their sense of smell is particularly acute, as they have a chemical sensor inside their siphons enabling them to locate odours dozens of metres away with great precision. The zone adjacent to lagoon inlets often contains luxuriant seagrass meadows, with a prevalence of Cymodocea nodosa and Zostera marina. This is a speciesrich environment, where organisms which benefit from organic matter in the sediment that accumulates at the base of the plants are associated with organisms which choose the fronds as their preferred habitat. Among the former, both oligochaetes and sedentary polychaetes are to be found in abundance, such as the capitellid Notomastus latericeus, which is tubicolous as a juvenile but lives in tunnels in sediment as an adult. The invertebrates populating the seagrass fronds include isopods like Idotea baltica, which look like small fragments of leaves. They may sometimes be seen navigating, clinging to pieces of leaf that they use as boats, pushed by the beating of their paddle-shaped pleopods. Tanaidacea like Apseudes latreillei are also frequent, living in small tubes on the surface of the seagrasses, and easily distinguished from isopods by the pincers on their first pair of limbs. A

great many amphipods live among the seagrass fronds, like the gammarid Gammarus aequicauda, and graceful shrimps like Palaemon elegans, with their slender legs ringed in yellow and blue, and P. adspersus, which nibble at any particles they are able to grasp. Browsers on the microalgal film that grows on the leaf blades include various species of gastropods, among which Gibbula are particularly widespread, for example, G. adriatica, and the small cerithiid gastropods Bittium reticulatum and B. scabrum. In this environment rich in submerged vegetation, Asterina gibbosa also make their appearance - delicate starfish a couple of centimetres in diameter, and the small and fascinating anthozoan Anemonia viridis, which anyone who has ventured into underwater meadows without adequate protection knows, to their cost, because these anemones can sting with painful results. In the meadows closer to the inlets Pinna nobilis is also present. This is a pretty pinnate-shaped bivalve of large size, whose fan may rise about thirty centimetres from the bottom and whose byssus used to be spun and woven in some Italian localities (the island of SantAntioco off the south-western coast of Sardinia was once famous for this). P. nobilis is accompanied by the dark cockle (Chlamys varia) and the lucinid Loripes lacteus. The latter is an unusual bivalve, in that it plays host to bacterial symbionts in its gills. These fix carbon dioxide present in the water, using energy obtained from oxidation of sulphides, which are abundant in the sediments at the base of seagrasses. Some of the organic

49

Palaemon elegans

Pinna nobilis

50

Xilophagous organisms
Among the rigid substrates available to benthic communities, wood has the double function of acting as a support for encrusting populations, and food for other organisms, mainly bacteria, fungi and xylophagous invertebrates (wood-eaters). As the activity of bacteria and fungi is very slow, most of the macroscopic degradation of wood is the work of invertebrates. These include bivalve molluscs belonging to the family of the teredines, among which the most common are Bankia carinata, Lyrodus pedicellatus, Nototeredo norvegica and Teredo navalis, which excavate large tunnels as wide as a finger, and crustaceans the isopods Limnoria lignorum and Limnoria tripunctata and the amphipod Chelura terebrans, which are responsible for a denser network of galleries. The evolutionary history of these organisms has mostly taken place within environments like lagoons, estuaries and deltas, where there are large quantities of wood in contact with the water. In heavily anthropised environments, the majority are breakwaters or other wooden structures, placed there by man, and their degradation involves high replacement costs. The teredines have a wormlike body with a small shell at the front end. Evolutionary processes have transformed this from a protection for the animal into a digging implement, rendering it similar to the head of a drill bit, which these molluscs rotate with movements of their robust foot. These animals colonise a site when planktonic larvae arrive and subsequently metamorphose and grow inside the wood, building tunnels along the entire submerged section. The dispersal dynamics of individual species are influenced by the duration of their larval stage, which lasts for two or three weeks in T. navalis but only a few hours in L. pedicellatus. The tunnels, lined by a calcareous layer, have practically imperceptible dimensions in the first stretch, but increase markedly in diameter and length as the organism grows. The wood is pulverised and partly used as food by means of digestion assisted by symbiont bacteria. Salinity and temperature are the principal environmental factors determining spatial distribution: T. navalis, euryhaline (tolerating salinity of 10), is the species best adapted to estuary environments; Lyrodus

Marco Sigovini pedicellatus finds its optimum in less desalinated conditions, and other species require water more like that of the sea. Xylophagous crustaceans are also widely distributed in estuaries, deltas and lagoons. Although they lack a planktonic larval stage, their diffusion is guaranteed by currents which transport individuals to new sites. These organisms only attack wood within the mid-littoral belt. In synergy with the waves, the outer layers become detached, conferring the typical hour-glass shape on highly degraded wooden piles at the point of mean sea level. Although the system of tunnels is often highly unstable, it represents a new habitat that can be colonised by many other species of invertebrates.

51

The teredine Lyrodus pedicellatus

The hour-glass effect caused by xylophagous organisms eating away the wood

52

compounds obtained from carbon dioxide fixation are then transferred to the bivalve. The central basin of a lagoon may be divided into two large bands. The first, more external, facing towards the sea, has a succession of facies of classical biocoenoses of the muddy sands found in sheltered environments; the second, more internal, towards the river and dry land, contains some facies of euryhaline and eurythermal lagoon biocoenoses. In Upogebia pusilla the external band of the central basin, characteristic venerid bivalves are Paphia aurea and the carpet shell clam (Tapes decussatus). As already mentioned, Tapes philippinarum is very common, if not completely dominant. Other species of bivalves characteristic of this zone are Corbula gibba, Gastrana fragilis and Loripes lacteus. The rich phytoplankton and abundance of suspended organic matter explain the frequency of other efficient filterers, such as Mytilaster minimus and the oyster Crassostrea gigas. The latter species is extremely common, both here and in the more internal area. Oysters were introduced into Italian lagoons and estuaries around the middle of the last century and spread rapidly, creating true miniature reefs in some sites which form a habitat of hard substrate particularly important for the promotion of biodiversity in environments otherwise characterised by muddy substrates. The gastropods Nassarius reticulatus and Cyclope neritea abound on lagoon beds, together with high numbers of Cerithium vulgatum and C. alucaster, with its elongated shell which may reach eight centimetres in length. Bittium and Gibbula are also abundant on beds covered by macrophytes. Roving polychaetes include some predators of considerable size, sometimes a few dozen centimetres long, such as Nephthys hombergii, Marphysa and Glycera. In this band, the numbers of nereidid polychaetes like Alitta succinea (=Neanthes succinea) and the ragworms Perinereis cultrifera, P. rullieri and Hediste diversicolor begin to be significant, although they reach their peak in the next area. The mouth apparatuses of these polychaetes, even when due account is taken of scale, are extraordinary and formidable: among the predators, eunicids like Marphysa possess many pairs of mandibles of varying shape, whereas Glycera has a characteristic evertible pharynx with four hooked

mandibles. Although they cannot be defined purely as predators, the nereidids possess two powerful, falciform, serrated mandibles, with a set of conical teeth. Among the sedentary polychaetes, some are tubicolous like Amage adspersa and the maldanids, also known as bamboo worms, because of the unusual segmented structure of their body, which is similar to a bamboo cane. The cirratulids of the genus Cirriformia Hermit crab (Diogenes pugilator) are sedentary polychaetes of a reddishorange colour, with long filaments like tentacles, which give them the appearance of jellyfish. They live buried just beneath the surface of the sediment, from which their tentacular filaments emerge to capture the organic matter on which they feed. The high eutrophication of lagoons, estuaries and deltas, especially in the central basin, also explains the abundance, among the polychaetes, of species typical of environments with high organic loads, such as Capitella capitata, Heteromastus filiformis and Polydora spp. Polychaetes of the genus Polydora are not only found in areas rich in organic matter. When we eat an oyster or observe the inside of the valves, we may note blackish blisters containing mud and covered by a thin layer of motherof-pearl. In them live specimens of Polydora cornuta (=P. ligni) and P. ciliata. These polychaetes can perforate the shell of their host the oyster, aided by specialised bristles and, probably, acid secretions, until they penetrate the internal cavity. Sensing a foreign body, the oyster secretes a layer of motherof-pearl around the hole made by the polychaete, forming a chamber within which the worm then lives. Along the edge of channels, many holes about as wide as a finger may be seen: these are the ventilation shafts of the burrows of a crustacean prized as bait by fishermen: Upogebia pusilla. This thalassinid decapod, pale greenish-beige in colour, vaguely similar to scampi, reaches a length of around ten centimetres and lives in Y-shaped tunnels that reach down into the seabed to a depth of about fifty centimetres. The shells of gastropods, especially those of Nassarius nitidus, Cyclope neritea and Cerithium, are often inhabited by the hermit crab Diogenes pugilator, which is very abundant in this lagoon environment, especially

53

54

Cereus pedunculatus

along the edges of the largest tidal creeks between sandbanks and near seagrass meadows. A characteristic anthozoan of this belt is Cereus pedunculatus, which attaches itself by its basal disc to shells lying just below the surface of the sediment, from which its crown of tentacles protrudes. Some echinoderms are to be found In the most highly vivified areas. Examples are the holothurians Trachythyone tergestina and T. elongata, small sea cucumbers a few centimetres in length, and the starfish Asterina gibbosa in seagrass meadows. Many crustaceans can be found on hard substrates such as channel banks built in bricks, stone or cement, the poles and brcole (the typical triangular groups of thick oak pites defining deep-water channels), including the amphipods Corophium acherusicum, C. acutum, C. sextonae, Stenothoe tergestina and Jassa marmorata, and the cirripede Balanus amphitrite, easily recognised by its vertical purple stripes on a white background. We are now entering the innermost part of the central basin, an area of high sedimentation, distinguished by many sandbanks; the seabed is silty-clayey and the water is often desalinated. All the inhabitants of this area generally tolerate both seawater dilution and the resulting decrease in salinity, and relatively stagnant waters. The bivalves which characterise this zone are Cerastoderma glaucum, Abra segmentum (=A. ovata) and, in the more brackish areas also containing groundwater springs, Scrobicularia plana. Cerastoderma glaucum is very abundant in this band. Its rounded, slightly heart-shaped form prevents it from sinking very deep into the substrate, from which it often protrudes, becoming easy prey for crabs. Scrobicularia plana is a bivalve reaching more than five centimetres in length, whitish in colour and flattened in shape, which facilitates its burial in muddy sediments. Its long siphons brush the surface of the sediment, producing star-shaped traces around the holes from which they protrude. The flattened shape and long siphons of Abra segmentum are very similar to those of Scrobicularia, but are much smaller, being only about one centimetre long. This bivalve also alternates a detrivorous diet with food, which it obtains by filtering the water, and may be found in dense populations, sometimes subject to great fluctuations in numbers. It is in this belt that Nassarius nitidus progressively gives way to Cyclope neritea. More than a general tolerance to environmental factors, it is likely that their methods of reproduction determine the distribution of these two nassariids. N. nitidus lays rows of pointed capsules containing around a

55

56

hundred eggs, preferably on seagrass leaves; the resulting larvae, before settling, pass slightly less than a month as planktonic forms carried by currents. Conversely, Cyclope neritea does not go through a larval dispersal stage: it lays single eggs on hard substrates, such as shells, from which hatch miniature adults that can immediately cope with a less stable environment. A sedentary polychaete typical of this zone is the spionid Streblospio shrubsolii. This small annelid, about half a centimetre in length, loves environments close to estuaries. It has a very high growth rate and its colonies Nassarius sp. are very large, so it represents a good food source for the fry of many species of fish. Other polychaetes are associated with these species, such as Alitta succinea and the predator Nephthys hombergii. The behaviour of the nereidid A. succinea is particularly interesting, as it passes most of its existence on the seabed but, when it reaches sexual maturity, abandons it at night. The gametes of this worm are thus freed in the water close to the surface, which explains why, at certain times of the year, towards midnight, up to 4-5000 Alitta eggs per litre of water may be counted. The most common amphipods are Microdeutopus gryllotalpa, which feeds on fragments of algae, the detrivore Melita palmata, and the filter-feeder Ericthonius punctatus. The hard substrates of this belt are encrusted with barnacles (Balanus eburneus and B. improvisus), which also enter the next zone, often in large numbers. Our curiosity will undoubtedly be aroused by mounds of mud that rise from the bottom of pools between sandbanks: these mark the exit holes of the complex tunnels of another thalassinid crustacean, the ghost shrimp Callianassa tyrrhena, which may extend for more than half a metre beneath the surface of the sediment. The name of this species comes from its white translucent appearance, which does make it appear as if it were the ghost of a prawn or scampi. Another characteristic of the species is the larger size of one of the two chelipeds (pincers), which is used for digging the tunnels.

Two very common amphibious gastropods, Truncatella subcylindrica and Myosotella myosotis, are to be found on sandbanks. Lastly, this small world between water and land is home to the amphipods preferring sandy shores belonging to the genera Talitrus, Talorchestia and Orchestia. These are extremely lively animals of marine origin, which hop about swiftly on damp sand or amongst the beached detritus where they find their food. We are now nearing the estuary zone, where the entry of river water greatly reduces the salinity of lagoon waters and enriches sediments with organic matter. On deep bottoms, Cerastoderma makes way for Scrobicularia, while the tiny gastropods of the genus Hydrobia accompany the amphipod crustaceans Corophium orientale and C. insidiosum in pools between sandbanks. Corophium looks like a sort of mechanical excavator in miniature - only 1 or 2 centimetres in length, it has two very robust arm-like antennae about half the length of its body, with which it displaces sediments, raking up the particulate on which it feeds. This is the zone where the roving polychaete Hediste diversicolor, which fisherman often use as bait, finds its most favourable environment. Despite being a highly resistant species, it does need a certain amount of oxygen, and so is often found in the intertidal zone. This polychaete reproduces only once during its lifetime, which lasts at most a few years.

57

Amphipod of the genus Talitrus

58

Gametes are released more or less in synchrony in the entire population. Eggs and sperm leave the animal through an opening in the body wall, almost always close to the head. However, not all animals of the same age reproduce in the same year; some late developers mature during the following year, thus constituting a valuable reserve if any unforeseen events compromise the breeding success of their siblings. Victorellid bryozoans are to be found on hard substrates. The hydrozoan Cordylophora caspia and the tubicolous polychaete Ficopomatus enigmaticus are introduced species capable of creating reefs of great size. The amphipods Gammarus aequicauda, Leptocherirus pilosus and the tanaidacean Heterotanais oerstedi form large populations in the first stretches of this sector, but they are also found in other areas of the central basin, because of their tolerance to environmental factors. Shallows particularly rich in organic matter, in which there is very little available oxygen, are inhabited by capitellid polychaetes, including Capitella capitata and Heteromastus filiformis. Such is the affinity of Capitella for this type of sediment that its larvae appear to be attracted by the hydrogen sulphide that rises from it. The very abundant and tiny members of some species of tubifid oligochaetes are often to be found together with Capitella. Capitellids and oligochaetes are also abundant in environments rich in organic matter and not desalinated, as in the deposits forming at the base of seagrasses.

Planktonic fauna Mesozooplankton (planktonic animals 0.2-2 mm long) play a very important role as food source for the larvae of fish and other filter-feeders. Their seasonal variations in density follow those of phytoplankton with a slight delay, peaking between the end of spring and summer. The importance of plankton is recognised for all filtering organisms of the benthos, but less expert naturalists tend to under-estimate the role of plankton in the diet of vertebrates. Tangible proof of this comes from an examination of the stomach contents of the most common fish which live in The harpacticoid copepod Euterpina acutifrons these basins. The planktophagous dietary regime of the young of five species of fish of commercial interest is welldocumented, such as the sea bass (Dicentrarchus labrax), the gilthead seabream (Sparus auratus) and three species of mullet (Liza ramada, L. saliens and L. aurata). The young of all these species, until they reach a length of five centimetres, have a planktophagous diet, their preferred prey being calanids, polychaete larvae and cirripede nauplii. Whitebait (Atherina boyeri) are also active consumers of plankton. Small individuals feed mainly on the more minute fraction of zooplankton, composed of the earliest larval stages (nauplii) of copepod and cirripede crustaceans, as well as the larvae of polychaetes and molluscs. As the fish grow, their attention turns more to the larger components of the plankton, such as mysidacean crustaceans and decapod larvae, as well as benthic organisms, especially small polychaetes and amphipods. In proximity to the sea inlets, where fluctuations in salinity are more pronounced, the biocoenosis is rich in species, with cladocerans, cyclopoid copepods and neritic harpacticoids (for example, Oithona spp., Oncaea spp., Euterpina acutifrons) and tunicates (appendicularians). In lagoons, zooplankton is mainly marine (neritic: i.e., typical of the open sea) around the inlets and in the more external belt of the central basin. In internal parts it also includes a native component, composed of organisms that can pass their entire life-cycle in brackish waters, and also types of freshwater origin. An important component of meroplankton are the larvae of various

59

Tubicules of polychaetes Ficopomatus enigmaticus

60

species of annelid polychaetes, gastropod and bivalve molluscs, and cirripede and decapod crustaceans. Within the marine component of plankton, the important group of the copepods is represented by many species: the calanids, typical of neritic plankton, include Acartia clausi and Paracalanus parvus; the cyclopoids include Oithona nana, O. similis, O. plumifera, the pecilostomatoids include Oncaea sp., Corycaeus sp. and, lastly, the harpacticoids include Euterpina acutifrons and Microstella norvegica. Among the cladocerans, which are more abundant in the summer months, the most frequent species are Podon polyphemoides and Penilia avirostris. Other zoological groups, present at least seasonally, are the tunicates (appendicularians), hydromedusae, turbellarians, nematodes, ostracods, amphipods, mysids, chaetognaths (Sagitta sp.), and also the eggs and larvae of fish. A very curious although irregular presence is Noctiluca miliaris, a spherical unicellular organism almost a millimetre in diameter - an exceptional size for an organism composed of a single cell. It is capable of emitting light if stimulated by water movement - a phenomenon easily observed on the occasions between spring and summer when Noctiluca reproduces in infinite numbers in coastal waters. The components of the central basin, especially in the internal part, include the larvae of cirripede crustaceans, some rotifers (two species of Synchaeta and Brachionus plicatilis) and many copepods:

The copepod Canuella perplexa

Calanipeda aquaedulcis, Canuella perplexa, Halicyclops sp., Harpacticus sp., Microarthridium fallax, Acartia margalefi and A. tonsa. In the Lagoon of Venice, the seasonal abundance of zooplankton is at its minimum in January, with dominant species Acartia clausi, Paracalanus parvus, Centropages spp. and Oithona spp. There is a slight increase in April, and then a period of growth dominated by Acartia tonsa until the peak in July. The copepod Calanipeda aquaedulcis This is followed by a rapid, progressive decline in the following months. Absolute maxima and minima, in summer and winter respectively, are recorded in the most internal areas of the lagoon, mainly due to the thermophily (heatloving characteristics) of the dominant species. The larvae of polychaetes, molluscs, crustaceans, decapods and fish peak during late spring. Again in the Lagoon of Venice, in the last fifty years there have been marked changes in the specific composition of the lagoon zooplankton, especially within the genus Acartia. The most typically marine species, A. clausi, is still abundant near the inlets, while the brackish-water species A. latisetosa has become increasingly rare. A. margalefi typically occupies areas with intermediate characteristics, but since the 1980s it has progressively declined in favour of A. tonsa which, recorded for the first time in the Lagoon of Venice in 1992, has now become the dominant species of the community. In the oligohaline part, where a mass of denser saltier seawater penetrates upriver near the bottom beneath the river waters, there is a mixing of brackish and river water components of the plankton. Freshwater plankton, composed mainly of rotifers, is joined by marine copepods and a few larvae of another typically marine group, the polychaetes. True plankton, naturally, exists in the sea and in still internal waters, especially in lakes. Identification of true plankton in a river environment may be debatable. In the flowing waters of a river, as well as swimming organisms (essentially fish), organisms typical of lake beds may also be found, snatched from their natural environment and carried away downstream. However, it is difficult to imagine a true population of small river organisms, incapable of any particularly active movement, which can live perennially in the water without being inexorably carried off down towards the sea.

61

Fishes
GILBERTO GANDOLFI

63

The fish fauna in the brackish waters of lagoons, deltas and estuaries includes species that are best classified according to the characteristics of their biological cycles. Some are constantly present throughout the year, like anadromous species, the adults of which migrate to fresh water to spawn, and juveniles which follow the same but Whitebait (Atherina boyeri) inverse route to grow to maturity in the sea; conversely, the adults of catadromous species go to sea to breed, whereas juveniles ascend internal waters. Then there are species present in brackish-water environments for various periods during the year, with transfers to the sea to avoid unfavourable environmental conditions and also for breeding - because they have floating eggs which would not be able to develop in internal waters. Lastly, there are both marine and freshwater species which are found in brackish waters only occasionally. Resident species These are typically euryecious fishes, i.e., species able to tolerate wide and rapid variations in the chemical and physical parameters of the environment in which they live. Their capacities for adaptation allow them to pass their entire life-cycle in lagoon environments. Lagoons are populated by a very diversified resident fish fauna, due both to the different environmental conditions in individual areas, and to influences determined by the times and ways in which lagoons came into being in their particular geographical locations. The populations of fish species in internal waters have been affected by several events since the Messinian geological epoch, during a salinity crisis in the Mediterranean about 5 million years ago. In the northern Adriatic, fish were also influenced by preceding events in the middle Pliocene, when the Po Valley district

Black goby (Gobius niger jozo)

64

was affected by links then existing between the Adriatic and para-Tethys seas, and by more recent events, during the Pleistocene (approximately 2 million years ago), caused by phenomena in the Ice Ages. As regards the fish fauna of internal waters in general, the Po Valley district is richer in species and endemic forms than peninsular Italy and the Pipefish (Syngnathus acus) islands, for the reasons mentioned above and also because of the greater quantities of internal waters and the consequent greater abundance and variety of transition environments between fresh and sea waters. The characteristic common to the resident fish species in lagoon waters is that they are usually smaller than the marine or freshwater species belonging to the same genera or families. Living in a highly unstable environment, with frequent and rapid variations in temperature, availability of dissolved oxygen, salinity of the water and availability of food, these populations anticipate sexual maturity as much as possible, with the consequently reduced size of adults. In other words, they adopt a strategy of population growth, defined as type r, in which the adults produce a large number of descendants which quickly replace them. In their turn, the new generations exploit favourable environmental conditions and rapidly reach sexual maturity, managing to survive critical periods, although in fewer numbers. In lagoons, environmental crises generally occur in summer, because shallow waters heat up rapidly and there is a consequent reduction in oxygen, which dissolves in an amount inversely proportional to temperature. Conversely, autumn spates in rivers carry large quantities of nutrients into the lagoons which, after low winter temperatures, trigger a phase of high production in the food chain in spring. This section describes resident fish species which are common to all Italian lagoons. The whitebait (Atherina boyeri) is a gregarious species, feeding mainly on zooplankton and also adaptable to freshwater environments. The tiny South European toothcarp (Aphanius fasciatus) is also a gregarious species with a wide ecological range: it prefers lagoon shores rich in vegetation, but may also live in fresh water and pools with high salt concentrations. The pipefish (Syngnathus acus) and the black-striped pipefish (Syngnathus abaster), with their slim threadlike bodies, live in lagoons on sandy or muddy beds with abundant aquatic vegetation, sometimes migrating to coastal waters

to overwinter. They are gregarious species which feed on zooplankton. The females release fertilised eggs into the incubating pouch of the males, which feed the embryos through blood vessel connections until their development is complete. The peacock blenny (Salaria pavo) is a species with individual territorial behaviour, linked to the benthic environment and found on beds on which hard objects abound (pebbles, stones, submerged pieces of wood, etc.) suitable as shelters. The females lay their eggs in the nests of the males, which protect them until they hatch. This species has conspicuous sexual dimorphism. The black goby (Gobius niger jozo), a typical inhabitant of the Mediterranean basin, has similar biological characteristics to those of the peacock blenny: sexual dimorphism accentuated in the breeding season, parental care by the males, and a diet based on benthic invertebrates or small fish. Both the black goby and peacock blenny have poor adaptive capacity to highly desalinated waters. The three-spined stickleback (Gasterosteus aculeatus) may be found in lagoon areas scarcely influenced by salt waters and in channels adjacent to lagoons. In Italy, the species is absent in southern regions and in Sicily; elsewhere there are populations living in brackish waters and also freshwater ones, even far from the coast. In the breeding season, the male assumes a conspicuous livery and builds a nest using plant material. He then courts the female, inducing her to lay

65

Three-spined stickleback (Gasterosteus aculeatus)

66

her eggs in the nest and, after fertilising them, protects them by continually fanning them with his pectoral fins, to provide them with oxygen and keep them clean. In the lagoons of the northern Adriatic, as well as the black goby, there are also other species of the gobiid family. The grass goby (Zosterisessor Fish-farm ponds in the Lagoon of Venice ophiocephalus) is the largest of the gobiids which has adapted to lagoon environments, sometimes reaching more than 20 centimetres in length. It lives on muddy beds, in which the individuals of both sexes dig individual dens among the seagrass roots. In the breeding season, which begins at the end of February and lasts until May, the dens become more complex and are occupied by one adult male, a few immature males, and two to four females. Canestrinis goby (Pomatoschistus canestrinii) is a species endemic to the northern Adriatic, and frequents sandy beds. In the breeding season, the male occupies a territory around some small submerged object (a mollusc shell, submerged wood, seagrass rhizomes, etc.), beneath which the female lays her eggs. The male looks after the eggs, which hatch within a few days. The presence of Canestrinis goby in a small lake at the mouth of the river Sinni in the Gulf of Taranto is most probably due to accidental introduction into a fishfarm together with grey mullet fry. The lagoon goby (Knipowitschia panizzae) is a very small species that rarely exceeds four centimetres in length. It is practically endemic to Italian waters, extending only as far as the lagoons of Dalmatia. Records relating to the Lagoon of Lesina and lagoon environments of the Tyrrhenian coast very probably regard accidentally introduced populations that have recently become acclimatised. The lagoon goby lives on muddy beds, and the territorial behaviour of the male during the breeding season is similar to that of the previous species, but the hiding-place used as a nest is almost always a shell of some small bivalve of the genus Cerastoderma. Beneath this, after covering the half-shell with mud to conceal it, the male takes refuge to look after the eggs. Lastly, another representative of the gobiid family, Pomatoschistus tortonesei, is found in the Stagnone di Marsala in Sicily. This is another tiny species, even smaller than the previous one, which is typical of the lagoons on the African coasts of the central Mediterranean; its biology is practically unknown.

Catadromous species After feeding in internal waters, the adults of these species move into brackish waters prior to migrating to the sea in order to breed, while the juveniles make the reverse trip. Their presence in brackish waters is thus limited to specific periods of the year. Eel (Anguilla anguilla) Only one catadromous species, the eel (Anguilla anguilla), passes through the estuaries and deltas of Italian rivers in transit. It is known that the adults, having migrated from European waters for thousands of kilometres to reach the Sargasso Sea in the Atlantic Ocean, spawn there and die after releasing their gametes. The larvae (leptocephali) are transparent and have a characteristically flat shape. They are carried by the Gulf Stream towards the European coastline over a period varying from one to more than two years and, once they reach the continental shelf of Europe or enter the Mediterranean, by which time they are about 7 cm long, they metamorphose and assume a cylindrical shape. At this stage, they are called elvers, and begin to ascend rivers, especially at night when there is a rising tide. In Italian waters, this happens from October in the west until February in the Adriatic. The elvers distribute themselves over a wide range of environments: coastal brackish waters, canals, ponds, rivers and torrents, and lakes. They prefer soft bottoms and waters rich in vegetation, where they act as predators, feeding on invertebrates and small fish, prevalently at twilight and during the night. Their stay in fresh or brackish waters lasts for a few years - three to four for those which stay in brackish waters or on the plains, and up to seven or eight years for those living in colder waters. The former appear to be destined to become males, the latter females. This differentiation takes place after migration to the sea, as the adults which begin their breeding migration in autumn have not yet developed gonads. With maturity, their morphological characteristics modify, the colour on their backs varying from brownish-yellow to brownish-black, that of the belly from yellowish to silvery; the eyes become larger, the skin thickens, the pectoral fins become more pointed and the intestine begins to degenerate because the breeders do not feed during their long migration. They transform into what are known as silver eels, with lengths and weights varying from less than 45 cm and less than 200 g for those destined to differentiate as males, up to more than 1 m and 2 kg for the females.

67

68

Anadromous species The adults of these species, after feeding at sea, move seasonally into deltas, estuaries and lagoons, to ascend rivers until they reach suitable areas for spawning. After a stay in internal waters, the young individuals then take the inverse route, to pass their growing stages in marine waters until they reach sexual maturity. The fish species which make breeding migrations in internal Italian waters include the twaite shad and three different species of sturgeon, plus two species of lamprey the latter Beluga (Huso huso) belonging to a separate zoological class. The breeding specimens of twaite shad (Alosa fallax) swim up Italian rivers in spring, to spawn on gravel beds, sometimes very far from the sea. The young, after an initial growth stage in the river, descend to the sea in autumn, where they remain until they reach sexual maturity. Recently, differences in the number of branchiospines have shown that the populations of the Tyrrhenian are best considered as a separate sub-species (rhodanensis) from those of the Adriatic (nilotica). Of the three species of sturgeon that in the past ascended the major Italian rivers in spring, the largest, the beluga (Huso huso), which is typical of the Caspian Sea, Black Sea and the Adriatic, has only sporadically been recorded in the Po for some decades, and appears to be close to extinction. The common sturgeon (Acipenser sturio), which in the past had a wider distribution - ascending to spawn in all the major rivers of the northern Adriatic and also the Tiber and other rivers flowing into the Tyrrhenian Sea - seems to be following the same destiny. The third species, the Adriatic sturgeon (Acipenser naccarii), endemic to the Adriatic Sea and smaller than the others (rarely reaching more than 150 cm and 30 kg) appears to be able to pass its entire life-cycle in freshwater, as demonstrated by the persistence of a small population in the middle stretches of the Po and lower stretches of the Ticino, upstream from barriers

69

Adriatic sturgeon (Acipenser naccarii)

70

that would be insurmountable during migration. In the last year, operations to repopulate the Adriatic sturgeon have been under way, and specimens from fish-farms have been introduced into the Po and the rivers of the Veneto region. Among the four species of lamprey present in Italian waters, two make spawning migrations in rivers after passing their trophic stage in the sea as ectoparasites on fish or marine mammals. The sea lamprey (Petromyzon marinus) ascends rivers in spring or early summer, sometimes attached to the flanks of migrating twaite shad or sturgeon. It is found in all Italian rivers, whereas a second species, the river lamprey (Lampetra fluviatilis), only goes up the estuaries of rivers flowing into the Ligurian Sea and northern and central Tyrrhenian. Overall, the populations of species which make obligatory breeding migrations from the sea to internal waters, or vice versa, are those which have been most badly affected by human acts of environmental degradation. As well as the negative effects of water pollution on the development of eggs and initial growing stages of the species that make potamodromous (river) migrations, dams or other barriers built across rivers prevent adult individuals from reaching the beds suitable for reproduction. Similarly, it is now often impossible for eels to reach the stretches of rivers they inhabited in the past during their growing stage.

Seasonal species A few marine species feed temporarily in lagoons and estuaries, sometimes swimming for long stretches upriver, but they return to the sea both to spawn and to overwinter. Five species belonging to the mullet family are frequent in all the brackish waters of the Italian peninsula: flathead mullet (Mugil cephalus): fry enter internal waters from August to December, and adults migrate to the sea from summer to early autumn; thinlip grey mullet (Liza ramada): fry enter internal waters in autumn or at the end of winter, and adults migrate to Sea bass (Dicentrarchus labrax) the sea during the last months of the year; golden grey mullet (Liza aurata): fry enter internal waters in autumn or at the end of winter, and adults migrate to the sea from September to November; leaping grey mullet (Liza saliens): fry enter internal waters in summer, while adults migrate to the sea; thicklip grey mullet (Chelon labrosus): fry enter internal waters from April to June and adults migrate to the sea from February to April. These are all gregarious fishes with a diet based on plankton during the juvenile stages. Adults feed on algae and detritus rich in organic matter. These species are typical of lagoon and estuary environments with muddy and sandy beds; some swim upriver for varying stretches, being able to tolerate variations in salinity. In particular, the thinlip grey mullet has lengthened its ascent of the Po in the last few decades, now reaching distances of more than 200 km from the delta in summer, probably as a consequence of recent changes in the riverbed, which is much richer in organic matter than in the past. Other species which utilise brackish water for growing-on of juvenile stages are the sea bass (Dicentrarchus labrax) and the gilthead seabream (Sparus aurata). In late winter and in spring, the fry of these two species enter lagoons to feed on zooplankton. Being less tolerant of desalinated waters and cold temperatures than the mullets, they return to the sea during the winter months. Sea bass adults mostly

71

Sea lamprey (Petromyzon marinus)

72

prey on crustaceans and fish, and gilthead seabream mainly on bivalve molluscs. The five species of mullet, together with sea bass and gilthead seabream, are traditionally farmed in sometimes artificially enclosed brackish ponds. As these species cannot reproduce in brackish water, the fry are caught as they instinctively try to move upstream to internal waters, or else, they are bred by artificial techniques in which the males are induced to mature by treatment with gonadotrophic hormones. The plaice (Platichthys flesus luscus) lives in the lagoons and terminal stretches of the rivers of the northern Adriatic. This sub-species spawns in the sea in late autumn and winter. The larvae, which still have bilateral symmetry, enter the internal waters where they undergo metamorphosis when they are slightly more than 10 mm long: the left eye migrates to the right flank next to the right eye; the small fish acquires benthic habits, and its left flank gradually loses its pigmentation. The plaice, now flat, feeds on small invertebrates. Lastly, the sand goby (Pomatoschistus minutus elongatus) is found in lagoon and estuary waters along Italian coasts. It is slightly larger than its congeners which live permanently in lagoons. It spawns at sea and uses internal waters for its trophic stage, returning to the sea in winter or when the rivers are in full spate in early spring.

Occasional species In certain conditions - for instance, the autumn spates of rivers, or the summer penetration of the so-called salt wedge, which happens when waters are low some typically freshwater or marine species may occasionally be found in brackish-water environments, thus demonstrating their capacity to live with very different degrees of salinity to that to which they are normally exposed. It is not rare, for example, to observe some species of cyprinids in estuary waters, or the occasional pike (Esox lucius) in the most desalinated areas of the lagoons. Sporadically, in the hottest summers, the trout (Salmo trutta), a fish typical of mountain and hill waters, may even reach the sea in search of cooler waters. This sporadic behaviour mimics the regular anadromous migrations of the trout which populate the tributary rivers of the Black Sea, northern Atlantic Ocean and North Sea. The waters of the Mediterranean are too warm and salty for the needs of this species. Recordings of coastal marine species are more frequent - for example, the sardine (Sardina pilchardus), sprat (Sprattus sprattus), corb (Umbrina cirrosa), red mullet (Mullus barbatus) and sole (Solea solea). These are very probably only episodic visits of species which are not resistant to significant variations in salinity.

73

Fishing with large drop nets at the mouth of the Mignone (Latium)

Sole (Solea solea).

Terrestrial vegetation
FRANCESCO BRACCO MARIACRISTINA VILLANI

75

Estuaries, deltas and lagoons - those complex transition areas between fresh and salt waters, between land and sea are environments where the incessant interplay between the eroding action of winds and tides and the arrival of new material from rivers continually remodel the lines of the landscape and, following delicate equilibria, create a set of complex ecosystems in continual evolution. The deposition of sediments creates Vegetation on coastal dunes dunes, beaches and tongues of sand in the shape of sandbars which, over time, will be demolished by the waves, the impetus of high river spates, or subsidence. Vegetation plays a key role in stabilising and consolidating submerged and emerging sediments, which would otherwise be condemned to a state of perennial movement. Marked geomorphological variability corresponds to a variety of habitats, and consequently gives rise to plant communities which differ in their response to the salt gradient or particle size and texture of the substrate. Lagoon environments are differentiated into mudflats, sandbanks, tidal creeks, saltpans, marshes and inlets, which are all dominated by halophilous (salt-loving) vegetation, and the coasts and islands which host psammophilous (sand-loving) vegetation. Reed-beds Reed-beds are one of the most representative and widespread types of vegetation at river mouths and in coastal lagoons. In some situations - for example, at the terminal stretches of the major Italian rivers (Po, Adige, Tagliamento), favoured by the width of the riverbed and slow current - reed-

The Po Delta

76

beds are so extensive that they confer a characteristic physiognomy on the delta landscape. Reed-beds are easily identified by their typical appearance a dense and impenetrable grassland of gigantic herbaceous plants that may grow more than three metres above the level of the water. The undoubted main actor in this plant community is the reed (Phragmites australis), a tall grass which thrives in a variety of environments: from coastal areas to an altitude of 2000 metres, from fresh to brackish waters, from tranquil lakesides and ditches to riparian strips along streams, to lagoon sandbanks and marshes. Nor does the Reed (Phragmites australis) reed disdain environments without the constant or frequent availability of water, like flood plains or riverbanks, even in highly disturbed situations. Reeds have thin hollow stems, which may lengthen to more than three metres, and grow straight and very close together, forming dense stands of vegetation. They bear two rows of greyish linear-lanceolate leaves with sharp edges, accompanied by fine tufts of hairs where they join the stem. The long culms of reeds become more conspicuous in summer, when greyish-purple inflorescences appear at the top. Each panicle, 20-30 centimetres in length, is formed of a large number of feather-like spikelets, each bearing tiny flowers accompanied by soft and silky hairs. When the fruit have ripened, the entire spikelet breaks off the panicle, leaving only the basal elements (glumes) adhering. The hairs now become efficient organs of flight, to aid dissemination of the spikelet by air currents. The hypogeal (below-ground) apparatus of reeds is less obvious, but equally unusual. Robust creeping rhizomes, buried in muddy beds, form a dense subterranean tangle, a sort of net that constitutes the foundations of the reedbed. Each rhizome, covered in coriaceous scales, grows from year to year and may reach several dozen metres in length, producing both anchoring roots and new shoots at every node. Reeds are widely known for their traditional uses: at one time, the stems were cut during winter and used to insulate ceilings and roofs, to produce mats, stuff the seats of chairs or make brushes. In the Polesine (part of the Po Delta), the

arle, as reed mats are known locally, were placed vertically around the edges of vegetable gardens as protection against the wind. Over time, these uses and practices have progressively waned, but reeds are still exploited nowadays. They are successfully used in operations of natural engineering, thanks to the vigour of their vegetative reproduction. Portions of rhizomes or green cuttings are collected in areas where reeds grow abundantly and transplanted to wetland areas in need of environmental upgrading. Reeds are also one species very often used in the creation of phyto-purification systems, in which waste waters are channelled through a specially created route to prolong contact times with the plants. This permits efficient uptake of nitrogen, which is thus removed from the water and stored in plant tissues. The capacity of reeds to survive conditions of prolonged submersion make them ideal natural water purifiers. Owing to their wide ecological range, reed-beds may be found in different territorial ambits. In reality, the name reed-bed is used for different plant communities, which share physiognomic uniformity but which have substantial differences in composition, as the floral retinue changes according to ecological conditions. The common denominator to all communities is the clear quantitative dominance of reeds, a highly competitive species that creates a dense canopy, but which is not very conducive to the growth of other plants. As a consequence, few accompanying species survive.

77

Reed-bed in the Po flood plain

78

Plants and coastlines

Humans have exploited deltas and lagoons throughout history. Since the earliest times, man and the sea, just like man and rivers, have contended for space in repeated competition, first conquered by one, then by the other, but more often with no lasting results, because nature has often re-appropriated surface areas that man had previously succeeded in wresting from it. A significant example of this ongoing battle is the Lagoon of Venice, which has modified its geography innumerable times over the course of millennia. When, for various reasons, the coastline has receded, man has been able to exploit large areas of grassland for crops or as pasture for livestock, sometimes also building roads. An emblematic case of how plants can help us to understand the past
0 1 2 km

Mariacristina Villani Antonella Miola structure of the territory is that of the ancient road, the Via Annia which, traversing the plain between the rivers Sile and Piave (NE of Venice), linked the two large Roman settlements of Altino and Concordia Sagittaria. Vegetation gives clues as to why its route was moved further inland in Roman times (1st century AD): finds of fossilised plants, particularly the pollen of halophilous species, demonstrate the advance of the lagoon, which at that time invaded the area corresponding to the current village of Ca Tron (near Roncade, in the province of Treviso). In practice, the sea had won back what it had previously ceded flooding the original route of the road and making it necessary to build an alternative one, further from the coastline.
San Don
Me olo
Pia ve

They are mostly heliophytes, i.e., with a subterranean portion that roots them to the bed of the water-body, while the epigeal (above-ground) part emerges from the surface and lengthens in the subaerial environment. These are highly specialised plants. One of the main problems that plants in a reed-bed have to solve is lack of oxygen, which is poorly soluble and diffuses slowly in water. The roots, permanently buried in the asphyxial mud, with little oxygen, are particularly affected. A mechanism that counterbalances this lack is the translocation of oxygen from tissues that have plenty of it to others that do not have enough. The key to this process is the existence of a particular parenchymal tissue in the rhizome, called aerenchyma, which has many large intercellular spaces that are interconnected forming a system of conduits facilitating the movement of oxygen. In this way, oxygen can easily reach the root apparatus from the green epigeal parts, where it is produced by photosynthesis or taken from the atmosphere. Reed-bed species also have to tolerate another negative characteristic of the anoxic mud in which their roots are buried. The oxygen-poor environment stimulates the blooming of an anaerobic bacterial flora whose metabolism generates toxic products for the plants - although, by means of sophisticated physiological strategies, the latter manage not to suffer the consequences. Reed-beds are typical of wetlands and freshwater marshes, such as the small lakes behind the dunes, but those which flourish best are found in the riparian

79

Marteggia C Tron

Via

t limi on i um xim expans Ma on o f lag

An

nia

Caposile
Si

le

ia Ann Via

Portegrandi LAGOON OF VENICE

The Roman road Via Annia (south of Treviso) in relation to the maximum expansion of the Lagoon Marshy area at the mouth of the Agri (Basilicata)

80

Heterotopy
The proximity of the Eastern Alps to the coast and the existence of efficient transport systems towards the sea by rivers has led to the presence of what are more properly Alpine species of plants in coastal areas and, in particular, close to river mouths. The survival of their plant populations is probably closely linked to river currents regularly carrying seeds from higher land. This is an extreme case of heterotopy, a biogeographical term that describes the existence of populations of species dislocated outside their typical distribution area, due to transport of dispersed seeds. For example, the presence at the northern Adriatic river mouths of

Francesco Bracco Mariacristina Villani some species of shrubs which are usually components of holm-oak woodland is interpreted in this way: rock buckthorn (Rhamnus saxatilis ssp. saxatilis) and purple broom (Cytisus purpureus) are examples. One large tree also shares this peculiar phyto-geographical condition: the Austrian pine (Pinus nigra) which, although it grows spontaneously on the dunes at the mouth of the Tagliamento, is generally typical of the northeastern calcareous uplands. The presence of spring heath (Erica carnea) in this context has also been interpreted in this way, but it may also be a relict of the last Ice Age.

belt along the terminal stretches of rivers, in conditions of perennial or prolonged submersion. Here, other tall grasses are associated with reeds, such as reed canary grass (Phalaris arundinacea) and reed sweetgrass (Glyceria maxima), and sedges like the great pond sedge (Carex riparia) which, due to their height, manage to be competitive in capturing light. Also frequently found are other species of interest because they are rare and Marsh spurge (Euphorbia palustris) appear in the Red Lists of species at risk of extinction, such as the marsh spurge (Euphorbia palustris) or the fen ragwort (Senecio paludosus), with its yellow flower clusters that stand out among the reeds during the summer. In situations with shallow water and recurring, prolonged periods of drought, species typical of water meadows are associated with reeds, such as wood clubrush (Scirpus sylvaticus), with its characteristic triangular stem and bracts like large leaves that enfold the inflorescences; marsh woundwort (Stachys palustris) with its purplish-red corolla, purple loosestrife (Lythrum salicaria) and yellow loosestrife (Lysmachia vulgaris), with its bright yellow petals. Hedge bindweed (Calystegia sepium), with white funnel-shaped flowers, is almost always present, and succeeds in reaching the light by climbing up the reed stems. Another recurrent, but less frequent climbing species is bittersweet (Solanum dulcamara), which has clusters of dainty purple flowers with orange anthers. Halophilic reed-beds. Salinity does not represent a grave problem for reedbeds: they grow both in moderately brackish waters and in more salty conditions and, in response to the saline gradient, are accompanied by species more or less tolerant of salinity. In littoral zones, at river mouths and in creeks, where abundant freshwater mixed with seawater reduces the salt concentration, indicator species typical of areas with a moderate level of salinity appear next to reeds, such as the sea club-rush (Bolboschoenus maritimus) with its inflorescence composed of reddish spikelets; soft-stem bulrush (Schoenoplectus tabernaemontani), with its greyish-blue stems; spearleaved orache (Atriplex prostrata), the leaves of which are spear-like blades covered by a mealy bloom; and sea aster (Tripolium pannonicum ssp. tripolium), with its capitulum with tube-shaped flowers forming a central yellow

81

Austrian pine (Pinus nigra) at the mouth of the Tagliamento (Friuli Venezia Giulia)

82

disc surrounded by purple ligulate petals. In these situations, the saltmarsh aster (Symphyotrichum squamatum) is also frequent: an alien species present in disturbed environments and capable of living in soils with a modest salt content. It is a plant with flowering branches equipped with very thin linear leaflets bearing a large number of small heads of minute flowers which, at dissemination, develop a delicate pink pappus. Where the effects of seawater are more marked, salinity becomes a strongly conditioning factor that has two effects: on one hand, it limits the vigour of reeds, which grow shorter; on the other, it reduces floral richness, because only a few species, with particular adaptation strategies, can tolerate high concentrations of salt. In perennially haline environments, like sandbanks and salt-marshes, some species are able to join the floral retinue of reed-beds, although, in conditions of more accentuated salinity, they would form plant communities and become dominant. The most eye-catching of these include the sea-lavender (Limonium narbonense), belonging to the family of the Plumbaginaceae. This plant is insignificant in the vegetative stage, when it has only a basal rosette of spatula-shaped leaves, but becomes striking in late summer and autumn, when the floral stem lengthens, bearing a corymbed panicle dense with small pink-purplish flowers. These maintain their colour for such a long time, even when the corollas dry, that they are often gathered and sold in decorative bunches. The leaves and stem are sticky to the touch

because they are covered with cells that secrete the excess salts absorbed by the plant - a sophisticated physiological specialisation that allows them to survive high concentrations of toxic chlorides. The composition of halo-hygrophilic reed-beds also includes the annual seablite (Suaeda maritima), a plant species of the Chenopodiaceae that is called roscan in the hinterland of the Veneto lagoons. This term is also commonly used to designate other species belonging to the same family which, like the annual seablite, are well-known because they are edible. In halophilic reed-beds, another annual species plays a similar role. This is a glasswort (Salicornia veneta), an annual succulent species with fleshy stems and branches that appear to be leafless. The flowers, rudimentary and joined in two groups of three, are also only just observable with the naked eye. As the Latin name suggests, glasswort is a common and endemic plant of the Veneto coast, where it forms the most external strips of vegetation of the sandbanks on the lagoon edges, which are periodically submerged by seawater. Another species composing halophilic reed-beds is shrubby glasswort (Sarcocornia fruticosa), a perennial plant with woody stems at the base and fleshy above; more robust than the previous Chenopodiaceae, it may exceptionally reach one metre in height. With a similar morphology to that of Salicornia veneta, with the exception of its size, shrubby glasswort used to be eaten - although nowadays pollution in the lagoons makes this unadvisable.

83

Flowering of pink sea-lavender (Limonium narbonense)

Salicornia veneta

84

Other marsh vegetation The tall herbaceous plants that accompany reeds in reed-beds may take on a dominant role in favourable conditions and characterise the physiognomy of the vegetation, so that reeds are relegated to a subordinate role. An example are the tall grasslands dominated by bulrushes (Typha latifolia or T. angustifolia). Less extensive in comparison to reed-beds, these plant communities do not pass unobserved, especially during their flowering and fruiting seasons. The species of the Typha genus have an unusual inflorescence, in which the Typha latifolia male and female flowers form two separate clusters along the terminal axis. The male flowers, tiny and without petals, constitute the upper portion of the inflorescence. The female flowers, which are also tiny and without petals, form a kind of velvety sausage, composed of many thousands of microscopic flowers set very close together. Each female flower is composed of an ovary and supported by a peduncle, surrounded by a whorl of long bracts. Not only the position, but also the time the inflorescences remain on the axis are different: after the anthers have opened and the pollen is freed, the male flowers have finished their biological task, so they fall and leave a section of bare flowering axis above the sausageshaped spike. Instead, the female part turns brown after flowering, and conserves the different parts of which it is formed. The ovary, transformed into a small fruit, remains at the top of the peduncle. On this the bracts, which later play an important role, are also preserved. During the winter, the inflorescence dries and dissemination takes place: in response to hygroscopic stimuli, the bracts spread out and cause the floral peduncles to break; the sausage-shaped spike thus disintegrates and the seeds are transported by the wind in the form of cottony wisps. These communities are generally richer floristically than reed-beds, probably because bulrushes form less dense populations than reeds and create less severe shading conditions - and also because the long linear leaves perpendicular to the ground do not develop dense canopies.

In areas where the influence of brackish water is higher, halophilic reed-beds give way to Puccinellio palustrisScirpetum compacti, we find the typical formations of sea club-rush (Bolboschoenus maritimus). This robust member of the Cyperaceae presents many affinities with reeds: a robust hypogeal apparatus, dense stems, and a wide ecological range that allows it to live in both freshwater and oligohaline environments. Another species that can tolerate low rates of salinity is round-headed clubrush (Scirpoides holoschoenus), which forms communities in which it is dominant, often in damp areas behind Sea club-rush (Bolboschoenus maritimus) dunes. The bed of rushes is unmistakable when the inflorescences appear at the top of the stems: they have the appearance of small balls subtended by a bract that lengthens upwards almost as though it were an elongation of the stem. If salinity increases, beds of maritime rushes appear, with sea rush (Juncus maritimus), a species that is prickly to the touch, with its stems and rigid pointed leaves borne by robust horizontal rhizomes. Vegetation with sea rush flourishes best on sandbanks, where it usually grows in the more internal and higher areas, where the effect of tides is felt less. One of the interesting types of vegetation in wetter areas with weak saline distribution is Mariscetum serratae, the term used for the community dominated by sawgrass (Cladium mariscus), a vigorous species belonging to the family of the Cyperaceae. The stem, cylindrical at the base and triangular at the top, a few centimetres across, bears leaves with serrated, sharp margins folded along the midrib. The inflorescence, which develops during the summer, consists of an anthela composed of more overlapping and often interrupted brownish anthelas. This species is more common in freshwater environments, but its capacity to tolerate low levels of salinity means that it frequently appears in delta areas. Although sawgrass is considered a sub-cosmopolitan species, i.e., diffuse over most continents, the drainage operations that have taken place in coastal areas in the past have drastically reduced the presence of these communities throughout Italy.

85

86

Low salinity is also tolerated by cordgrass (Spartina versicolor), a robust grass that forms dense carpets on dunes in formations where it tends to substitute marram-grass, although it is also found on the edges of lagoons. Less halophilous than its congener small cordgrass (Spartina maritima), S. versicolor mainly reproduces vegetatively, through subterranean rhizomes that give rise to tussocks of leaves: because of this, its grasslands are very compact and species-poor. When they do appear, the inflorescences are composed of three or four sessile ears up to five centimetres in length. Lastly, one peculiar aspect of the vegetational landscape of the deltas should be mentioned - the belts of halo-nitrophilous vegetation. These communities are dominated by annual species with the distinctive feature that they can tolerate moderate salt contents and the physiological aridity that follows, but also high nitrate contents. Although often covering small areas, they play an important role as pioneer plants which anticipate the other more strictly halophilous species. The best-represented families are the Chenopodiaceae, with the oraches (Atriplex tartarica, A. prostrata) and the Polygonaceae, with golden dock (Rumex maritimus) and sea knotgrass (Polygonum maritimum). Their presence is often associated with disturbed environments. Natural plant debris is often accompanied by quantities of waste transported and deposited by water - a tacit but manifest accusation of modern mans uncivilised behaviour towards nature.

Consolidated dunes and damp hollows: a complex vegetational landscape As well as the more halophilous vegetation, the combined morphogenetic activity of the sea and rivers creates a complex landscape, which is also rich in communities far removed from psammophilous, marshy, salty or brackish ones. A complex environmental picture is created that flanks forestry aspects, xeric (heat-loving) vegetation, internal hollows containing freshwater, and coastal water-bodies of brackish or salt water. This is possible because of the detailed geomorphology of the landscape in both raised areas and low-lying ones. The higher areas generally correspond to dunes stabilised by vegetation, and may lie at some distance from the present-day coastline. Hollows are strongly influenced by the presence of salty or brackish water if they are in contact with the sea: conversely, if they are affected by the water table or surface water of land origin, they will have typically freshwater characteristics. In places where plants find it difficult to survive, as in the northern Adriatic littoral arc, the complex environmental situation also provides a little space for species of plants with different geographical distributions and very different ecological requirements. In this area, and as far south as the Po Delta, on the tops and flanks of consolidated dunes where the evolutionary process of the soil is more advanced, the landscape at river mouths hosts woodlands of holm

87

Sea knotgrass (Polygonum maritimum)

Mouth of the Reno (Emilia Romagna)

88

Mosaic of vegetation at the mouth of the Reno (Emilia Romagna)

oak (Quercus ilex), the evergreen oak more typical of the Italian Mediterranean landscape. This does not appear to be an exceptional case, as this woodland formation grows on the dune areas at the mouths of the Tagliamento and Adige and in the Po Delta. The low impact of this woodland in todays landscape is due to the relatively small extent of the dune and palaeo-dune systems, but also largely to the progressive elimination of this type of vegetation by mans exploitation of coastal areas. In contrast to the image of instability and strong dynamism of the coastal landscape, in these situations the soil - although essentially composed of sea sand, and thus dry and well-drained, appears to be well-structured and has a significant accumulation of organic matter in its shallower levels. The holm oak can form a closed, continuous tree canopy that may reach a height of 20 metres. Together with the holm oak, the most common tree species is the flowering ash (Fraxinus ornus), which is always co-present, but playing a decidedly subordinate role. Two generally Mediterranean tree species but with different phyto-geographical characteristics are therefore associated: the holm oak, typically distributed in the circum-Mediterranean areas with marked summer aridity (steno-Mediterranean), and the flowering ash, with a distribution area that covers not only the northern coasts of the Mediterranean, but also gravitates around the Black Sea (euro-Mediterranean-northern Pontic). The shrubs beneath the trees develop into a rather scattered cover, with a mixture of species with differing phyto-geographical characteristics. The tree vegetation of the dunes is associated in the landscape with shrub communities, which represent structurally simpler evolutionary stages of the vegetation cover, gradually preparing for colonisation by forest species. The first stage of colonisation by woody vegetation is that of low-growing shrubs, less than one metre high, defined as the precursor of the holm oak woodland, characterised by climbing shrubs with a small herbaceous contingent. The dominant species are wild asparagus, osyris and the heath Erica carnea, accompanied, especially in the most northerly part of the Adriatic, by purple broom (Cytisus purpureus) and the buckthorn Rhamnus saxatilis ssp. saxatilis, a small spiny shrub with reddish bark and small brown flowers with four petals. Very frequent, although not abundant, is the wall germander (Teucrium chamaedrys), a prostrate dwarf shrub, with small coriacious lobate leaves. The herbaceous species mainly derive from the neighbouring arid grasslands and, among these, tor-grass (Brachypodium rupestre) and cypress spurge (Euphorbia cyparissias) are frequent. It is within these ambits that the holm oak makes a timid appearance together with the flowering ash, although only at the seedling stage.

89

90

Where the sea is close enough to allow the salty air to reach the cordons of stabilised dunes, the vegetation is unable to form holm oak woodland, but reaches its climax as compact shrubland dominated by juniper (Juniperus communis), identifiable by its spiny-tipped leaves in whorls of three and its bluish-black berries. To this is associated, generally subordinately, the sea buckthorn (Hippophae fluviatilis), a densely Rosemary-leaved willow (Salix rosmarinifolia) branched shrub with silvery bark, linear leaves that are white beneath, and characteristic bright orange berries. This community is completed by many of the shrubs already mentioned for holm oak woodland and scattered herbaceous species including bladder campion (Silene vulgaris ssp. tenoreana) and the sedge Carex liparocarpos. River mouth environments sometimes contain systems of depressions that are often affected by salinity, but others - for instance, along the coast of the Veneto and Friuli plains - are influenced by the presence of fresh surface and groundwater, with low nutrient loads and high concentrations of calcium carbonate. In these cases, the depressions may contain accumulations of peaty sediments on which unusual plant communities settle and which, at sea level and at a short distance from the coast, sometimes include species that require cool environments. Typical examples are the peaty grasslands of purple moor-grass (Molinia caerulea ssp. caerulea) that are reminiscent of those in spring areas on the north-eastern plain, of which they reproduce a good part of the floral retinue. This includes black bogrush (Schoenus nigricans), narrowleaf plantain (Plantago altissima), fragrant leek (Allium suaveolens) and marsh gentian (Gentiana pneumonanthe). In slowmoving or still surface waters within these grasslands, rare hydrophyte communities of waters with low nutrient contents may also find refuge. An example is the aquatic vegetation dominated by fen pondweed (Potamogeton coloratus forma heterophyllus) with both floating and completely submerged elliptical leaves, both types being translucent, veined, and often reddish. The rosemary-leaved willow (Salix rosmarinifolia) also appears here as a woody colonising species. This shrub, rarely exceeding a height of two metres, has linear or linear-lanceolate leaves that are hairy and silvery beneath, and dark green and shiny above. It is a plant with Euro-Asiatic distribution, which in northern Italy is to

be found only rarely on the peaty soils of the high plain, montane and subalpine belts. The delicate environment in which it lives means that it is endangered throughout Europe, and its presence in coastal vegetation has also been drastically reduced, so that it now only survives in a few sites, such as those close to the mouth of the Tagliamento. More generally, where freshwater conditions converge towards the general mesotrophic or eutrophic state more Marsh marigold (Caltha palustris) typical of surface waters on the plains, the marsh and aquatic vegetation found in ditches and flooded hollows tends to coincide with that present in the waterbodies of the plains. In the slow-moving waters of the Po Delta, for example, aquatic communities have been recorded with fennel pondweed (Potamogeton pectinatus), a species which may also penetrate into brackish water. In the bodies of freshwater, the typical communities are dominated by several hydrophytes, all with large leaves growing on the surface of the water: the white water-lily (Nymphaea alba), yellow water-lily (Nuphar lutea), fringed water-lily (Nymphoides peltata) and water chestnut (Trapa natans). Only extremely rare groupings of marestail (Hippuris vulgaris) have been recorded. The hollows may also contain hygrophile or mesophile woodland vegetation, dominated by species of deciduous broadleafs that are in stark contrast to the evergreen woodlands on higher ground. This type of vegetation is even rarer than that of the holm oak woodland. A prime example is the Bosco della Mesola, which has been growing for the last thousand years on the complex of dunes and depressions created by the sedimentation of the Po di Volano and Po di Goro, within the Po Delta. The forest community that settles on the more ancient dune systems with flatter morphology is, on one hand, reminiscent of holm oak woodland and, on the other, is related to the mesophile forest vegetation typical of the plains. The soil is moderately damp and only sporadically water-logged, so a multilayered forest develops, with tall trees, many shrubs, and sparse herbaceous undergrowth. Lastly, the survival of marsh woodlands with alder (Alnus glutinosa) is very infrequent on plains, but becomes exceedingly rare at river mouths. For example, there are small areas of alder woods on soft black soil at the mouth of the Adige, at times with marsh marigold (Caltha palustris) in the herbaceous undergrowth.

91