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I TA L I A N H A B I TAT S Volcanic lakes 17 Italian habitats Italian Ministry of the Environment and Territorial Protection / Ministero dell’Ambiente e della Tutela del Territorio e del Mare Friuli Museum of Natural History / Museo Friulano di Storia Naturale - Comune di Udine I TA L I A N H A B I TAT S Scientific coordinators Alessandro Minelli · Sandro Ruffo · Fabio Stoch Editorial committee Aldo Cosentino · Alessandro La Posta · Carlo Morandini · Giuseppe Muscio “Volcanic lakes · Fire, water and life” edited by Fabio Stoch Texts Silvia Arisci · Marcello Bazzanti · Arnaldo Angelo De Benedetti · Renato Funiciello · Mauro Iberite · Laura Lepore · Fiorenza Gabriella Margaritora · Luciana Mastrantuono · Giuseppe Morabito · Michela Rogora · Marco Seminara · Fabio Stoch · Daria Vagaggini In collaboration with Raffaella Berera · Vezio Cottarelli English translation Elena Calandruccio · Gabriel Walton Illustrations Roberto Zanella Volcanic lakes Fire, water and life Graphic design Furio Colman Photographs Nicola Angeli 47, 48 · Archive Museo Friulano di Storia Naturale 58, 60, 61, 62, 63, 64/3, 106 · Andrea Balestri 64/5 · Raffaella Berera e Vezio Cottarelli 90 · Compagnia Generale Ripreseaeree 10, 116 · Vitantonio Dell’Orto 98, 107, 109, 111, 112, 114, 115, 134, 135, 137 · Giuseppe Di Lieto 113 · Dario Ersetti 59 · Paolo Fabbro 6, 7, 15/1, 15/2, 18, 21, 38, 40, 139 · Renato Funiciello 11, 22, 132, 145 · Mauro Iberite 55, 64/4 · Giuseppe Ippolito 86 · Luca Lapini 108, 136 · Giuseppe Morabito 45, 51, 52, 53 · Giuseppe Muscio 13, 26, 41, 46 · Naturmedia 99, 102 · Roberto Nistri 64/2, 103, 105, 130, 138 · Fabio Stoch 9, 16, 17, 20, 23, 27, 28, 29, 33, 34, 37, 42, 43, 44, 57, 64/1, 64/3a, 65, 66, 67, 68, 69, 70, 71, 72, 73, 74, 75, 80, 84, 85, 87, 89, 92, 93, 95, 96, 97, 104, 110,121, 122, 123, 125, 126, 129, 131, 142, 143, 144 · Damiano Vagaggini 24, 25, 35, 54, 78, 79, 88, 100, 101, 117, 118, 119, 120, 124, 128 ©2007 Museo Friulano di Storia Naturale, Udine, Italy All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or trasmitted in any form or by any means, without the prior permission of the publishers. ISBN 88 88192 34 4 ISSN 1724-6539 Cover photo: Lakes Nemi and Albano, Latium (photo Compagnia Generale Ripreseaeree) 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 Contents Italian habitats Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Fabio Stoch · Daria Vagaggini Geological aspects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Arnaldo Angelo De Benedetti · Renato Funiciello 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 Hydrochemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Silvia Arisci · Laura Lepore · Michela Rogora Phytoplankton. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Giuseppe Morabito Macrophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 Mauro Iberite 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 Zooplankton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Fiorenza Gabriella Margaritora · Daria Vagaggini Zoobenthos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79 Marcello Bazzanti · Luciana Mastrantuono Vertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 Marco Seminara 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 Conservation and management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 Fabio Stoch · Daria Vagaggini Suggestions for teaching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 Marco Seminara Select bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147 19 Seagrass meadows 20 21 Subterranean Rivers and waters riverine woodlands 22 23 Marine bioLagoons, constructions estuaries and deltas 24 Italian habitats Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149 List of species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151 Introduction FABIO STOCH · DARIA VAGAGGINI Even the most inattentive passenger in an aircraft flying over the Italian peninsula cannot fail to notice round, sky-blue bodies of water dotting the landscape below. Although these lakes, large and small, apparently calm and surrounded by steep shores, are spotted at a glance, only those who know them really well are aware of their true nature and complexity. The protagonists of this pubblication are volcanic lakes, unique Italian environments which formed in ancient times, at the dawn of civilisation. They have therefore witnessed man’s social and cultural development in a strikingly beautiful natural context. Findings of Lake Nemi (Latium) Neolithic human settlements along the shores or below the surface of volcanic lakes show how closely associated mankind and lakes have been since prehistoric times. Later, the Etruscans and Romans were the first populations to use these environments for water supplies. An example is Trajan’s aqueduct, later called dell’Acqua Paola which, even today, after a thousand years, conveys the water of Lake Bracciano directly into the Janiculum Fountain in Rome. Volcanic lakes were not only used to supply water, but also aroused curiosity and fear, as well as poetical inspiration. For instance, in the 6th book of the Aeneid, Virgil tells us that the Cumaean sibyl took Aeneas to the gates of the Underworld near the Lake Averno (facilis descensus Averno: noctes atque dies patet atri ianua Ditis - the descent into the Averno is easy: the gate to dark Hades is open night and day). Researchers are tracing the invisible line that connects the past to the present, aiming at improving knowledge of the mechanisms that regulate these delicate ecosystems and, as lovers of nature, at discovering and learning to respect volcanic lakes. Lake Albano (Latium): the crater-shaped rim of the lake 7 8 Early research dates back to the late 19th century, when enthusiastic scientists with simple instruments walked long hours to perform their geological surveys, or sailed across these lakes to draw bathymetric maps and to collect samples for chemico-physical and biological analyses. Although modern scientists have better instruments and more knowledge, they share the same eagerness to learn and understand these habitats. Prolonged research has highlighted several characteristics that make volcanic lakes extremely valuable environments - first, their origin due to volcanic action in Quaternary times, which was very powerful in Italy. It created the craters and calderas (wide, shallow craters due to collapse of the central part of a volcano) that today contain these lakes. The particular morphology of these bodies of water, which are generally very deep for their total area, is due to their particular formation. The apparent tranquillity of their waters actually conceals deep changes in the Earth’s crust caused by still unremitting volcanic action - there are still phenomena of secondary volcanism, such as the release of gases - the same that terrified people in ancient times. In addition to their peculiar geological characteristics, volcanic lakes host several animal and plant communities which either live in their waters or use them for their nutrient content. The plant successions circling their shores and submerged aquatic macrophytes are the “lungs” of these ecosystems, together with various kinds of micro-algae, the fundamental link in the food-chain. These lakes are also inhabited by free-floating animals and those closely associated with lake bottoms. These links in a foodchain dominated by fish and aquatic birds all live and reproduce in these environments, invaluable troves of biodiversity. Precisely because of the importance of their fauna, all these lakes are included in protected areas, Sites of EU Interest (Habitats Directive) or in Special Conservation Areas (Birds Directive). We may be led by these words to think that the relationship between man and nature is compatible with the welfare of both. Unfortunately, this is not so. Several threats to their integrity and equilibrium loom over volcanic lakes as broken links of the invisible line connecting past and present. The nonsustainability of some human activities - such as intensive agriculture, excessive animal grazing and tourism near these standing waters, the introduction of alien species into them and changes in shore profiles - are only some of the factors which in recent years have caused gradual deterioration of the quality of the waters, and have jeopardised their flora and fauna. This new volume of the Habitat Series has two objectives. One is to describe volcanic lakes from several viewpoints, from their origins to their chemicophysical characteristics and biological communities. The other is to emphasise their value for the conservation of nature by focusing on the main problems related to their management. We hope this volume may contribute towards protecting these precious environments; which have supported man for millennia and which now risk irreversible deterioration. The area around Naples in a map of 1817-19, before the lakes in the Astroni crater were reclaimed Lush vegetation along the banks of Lake Martignano (Latium) 9 Geological aspects ARNALDO ANGELO DE BENEDETTI · RENATO FUNICIELLO Over the past two million years, Italy is the European area that has most been affected by Quaternary volcanism. This phenomenon is associated with the faulting of the Tyrrhenian edge of the Apennines, along Tuscany and Campania, an event which took place over the last 10 million years. The peri-Tyrrhenian volcanic arc (Pleistocene-Holocene epochs) runs for 420 km NW-SE from Val d’Era in the north (Orciatico and Montecatini, Val di Cecina), to Vesuvius in the south, with only a few points inland (San Venanzo, Cupaello and Pollino) or near the Apennine faults (Monte Vulture). This volcanic arc developed along the Outcropping eruptive rock forms the island of Martana in the middle of Lake Bolsena western margin of the Tyrrhenian backarc basin. A thinning of the Earth’s crust is visible in the geophysical structure of the lithosphere, together with extensive, progressive thermic flow within the Tyrrhenian basin which, in its southern part, gives rise to one of the most active geodynamic elements in the Italian area. Thinning allowed the subcrustal intrusion of a type of syenitic rock at Larderello (Tuscany) to the area around Rome (Sabatino District), together with mantle upwelling in the southern Tyrrhenian basin. Distensive tectonics began in the Middle-Upper Miocene through the formation of normal faults running NW-SE and subsidence to the NE, and perpendicular transcurrent NE-SW-running faults that dismembered the Apennine chain. The result was the formation of sedimentary basins filled with sand and clay to depths sometimes reaching 1000 m, and also deposits from environments transitional to shallower sea basins before external volcanism started. Aerial view of the Sabatino Volcanic District. Left: eastern shore of Lake Bracciano; right: Lake Martignano and (top) crater that once hosted Lake Stracciacappa. 11 12 deep basins in African - Adriatic domain newly formed oceanic crust Miocene mountain chain slip vectors of orogenetic transport strike - slip faults inactive front of Europa-verging mountain chain active thrust front active normal faults Simplified sketch showing the formation of the Tyrrhenian Sea, and front migration of the Apennine chain Volcanoes are closely associated with this faulting network that produced tecto-volcanic structures like the calderas of the Alban Hills and the Phlegraean Fields or large dyke networks like the Somma-Vesuvius structure. Integrated geological and geophysical data show that, under the volcanic arc and post-orogenic clastic deposits, the Mesozoic-Cenozoic (the Mesozoic epoch dates back 248-65 million years ago; the Cenozoic followed) carbonatic succession thinned, overthrust and laterally extended along a décollement surface on the crystalline basement at a depth of Volcanic deposits in the Vulture mountains (Basilicata) about 7-8 km. According to spacetime distribution and petrographic criteria, the Apennine chain is divided into three petrographic areas called magmatic provinces: Tuscan, Roman, and the recently identified Lucan. In each of these provinces, igneous rocks with differing petro-chemical affinities outpoured inside the main magmatic associations. From the morphological viewpoint, volcanism gave rise to: caldera edifices like those of Bolsena-Latera, Bracciano-Sacrofano, Vulcano Laziale (Alban Hills) and the Phlegraean Fields; stratovolcanoes like Vico, Faete, Roccamonfina, Ventotene, Ischia, Procida, Monte Somma-Vesuvius and Vulture; domed shield volcanoes like Monte Amiata, the centres of Tolfa, Ceriti-Manziate, Monti Cimini and the Ponza islands; or monogenic volcanoes like those of the Umbrian District, Monti Ernici, and Ponza. Many volcanic structures such as cones, craters, calderas, lava flows and lava fields are still well-preserved. Calderas and craters are the best places for rain and hydrothermal water to collect. This is due to the elevated rims of both structures and to the not very permeable nature of volcanic products, which hinders water drainage. The formation of a lake is therefore the result of a series of factors, the most essential of which is the substrate and its relationship with the surface watertable, as craters and calderas are like large natural wells fed by these waters. According to their origin, volcanic lakes are divided into primary and secondary. 13 Primary lakes are standing waters contained in volcanic edifices like calderas, tecto-volcanic depressions and craters, and are described in this volume. Secondary lakes are all basins whose origin, at least partially, was due to volcanism, but they formed in rocks which are not of volcanic origin (like sinkhole lakes, particular types of basins described on p. 16). Lastly, there are pseudo-volcanic lakes forming from the outflows of hot springs and mineral waters. In Italy, there are only a few lakes of this type, examples being Bagno dell’Acqua on the island of Pantelleria, and the lake at Arquà, in the Euganean Hills. 14 TN AO VE TS MI TO BO GE FI AN 1 2 PG AQ 3 5 4 7 6 ROMA CB 8 9 10 11 NA 12 17 18 13 14 16 15 BA 15 Lake Bagno dell’Acqua (Pantelleria, Sicily region) 19 PZ ■ Tecto-volcanic depressions and calderas CA 22 20 PA RC 21 Volcanic edifices 1 2 3 4 5 6 7 8 9 10 11 ORCIATICO-MONTECATINI AMIATA-RADICOFANI LATERA-BOLSENA CIMINI VICO TOLFA-MANZIANA-CERITE BRACCIANO-SACROFANO COLLI ALBANI MEDIA VALLE LATINA S.VENANZO-CUPAELLO ROCCAMONFINA Age (years) onset end 4,100,000 1,000,000 720,000 1,310,000 410,000 2,500,000 600,000 600,000 680,000 420,000 600,000 200,000 <130,000 900,000 90,000 <40,000 6,000 <80,000 55,000 Volcanic edifices 12 13 14 15 16 17 18 19 20 21 22 PONZA GROUP MONTE GUARDIA VENTOTENE ISCHIA PROCIDA PHLEGRAEAN FIELDS VESUVIUS VULTURE IEOLIAN ISLANDS ETNA USTICA Age (years) onset end 2,500,000 1,230,000 >1,000,000 150,000 <50,000 60,000 400,000 600,000 1,300,000 500,000 750,000 Ages of onset of activity, and time-span involved, of the main Italian volcanic edifices 1,070,000 480,000 1306 D.C. 1538 D.C. 1944 D.C. 110,000 PRESENTE PRESENTE 130,000 Some Italian volcanoes have calderas, or tecto-volcanic depressions, definitions that in the past were distinct, but which today are considered synonymous. Lake Arquà (Euganean Hills, Veneto) They are vast depressions inside the original crater caused by the violent emanation of large quantities of gases, partial emptying of the magma chamber, and consequent collapse of the central part of the volcano above the vent due to modified mechanical conditions underground. Large Italian volcanic lakes are Bolsena, Vico and Bracciano. Lake Bolsena. Once known by its Latin name of Lacus Volsiniensis, this is the largest Italian volcanic lake (114 km2, 151 m deep). Located in upper Latium, near Viterbo, it has two islands, Bisentina (0.17 km2) and Martana (0.10 km2). Its origin was due to the outflow of large quantities of magma during the early stages of the history of the Vulsino Volcanic District (576-500,000 years ago) which, according to the most reliable models, caused the top of the partially emptied magma chamber to collapse and the above area in turn to collapse 16 Sinkhole lakes in volcanic areas Sinkhole lakes are associated with volcanic activity. They are created by the collapse of a generally circular area, between a few and one hundred metres in diameter, caused by the formation of shallow cavities underground. Thanks to their particular shape, sinkholes often fill with water, giving rise to ponds or small lakes. The original cavities giving rise to sinkholes may be produced by poorly coherent sediments migrating to the bottom of palaeo-karstic cavities, which gradually fill with water. Others may form as a result of chemical dissolution - closely connected with volcanic activity since the aggressive substances responsible for this phenomenon are rich in carbon dioxide and hydrogen sulphide. Renato Funiciello Solution greatly affects limestone, which is widespread in Italy, giving rise to karstic cavities. Among the many bodies of water that have formed in this way, there are those near Rome (Lago di Puzzo, or Lago di Leprignano, Lago Nuovo and Lago di Giulianello); in the Piana di S. Vittorino (Rieti), the best-known of which is the Lago di Paterno; at the foot of the Lepini mountains (Laghetti del Vescovo); and near Caserta (Lago di Vairano, Lago di Corree). Lastly, in the Acque Albule area (Bagni di Tivoli, near Rome), a series of dolinas have become small lakes, some of which are still active. Examples are the Lago di S. Giovanni, Laghetto Regina and Laghetto delle Colonnelle: these sulphur springs are called Albulae because of their milky colour. Laghetti del Vescovo, at the foot of the Lepini mountains (Latium) inwards. The distribution of alternating volcanic and lacustrine deposits shows that, over time, the lake underwent deep modifications associated with the complex evolution of the Vulsino District. The first great eruptions released about 50 cu.km of magma - a far greater amount than the catastrophic eruption of Vesuvius in 79 AD which destroyed Pompeii (about 1 cu.km). This phenomenon occurred after large quantities of lava scoriae, deposited in Lake Bolsena (Latium) the northern circum-caldera area (360352,000 years ago), were followed by ignimbritic scoriae alternating with lava and lake materials. Recent underwater archaeological findings of Bronze Age settlements show that the lake level remained the same (294 m a.s.l.) for a long time. In the Iron Age, it rose rapidly for reasons still unknown, up to the highest level allowed by the natural point of flooding (inlet) at 306 m a.s.l.. Later, to reclaim land for agriculture, the Etruscans lowered its level to 303 m by cutting a rock spur that dammed the flow of the river Marta, in an area that is still called “Sasso Tagliato” (Cut Rock). The current level of the lake (303.5 m) is maintained by a masonry inlet and sluice-gates constructed in the Middle Ages in the port of Marta, at the outlet. Lake Vico. Lake Vico, or Lacus Ciminus, is the highest Italian volcanic lake (510 m). According to legend, it was produced by the club Hercules thrust into the ground to challenge the local population. Nobody was able to remove it. When Hercules pulled it out, water flowed and filled the valley, thus creating the lake. Lake Vico is actually a partly filled-in caldera which underwent several collapses due to the settling of types of ignimbrite like Tufo Rosso a Scorie Nere (red tuff with black scoriae). When the eruptions of the whole Vico area ended (80,000 years ago), underground water and rain gradually filled the caldera. The lake is surrounded by the Monti Cimini, Monte Fogliano (965 m) and Monte Venere (851 m). Lake sediments reveal that, for several millennia, the lake was larger, and its water reached the rim of the caldera, Monte Venere then being a peninsula in the north-eastern part of the lake. It is still not known whether the Etruscans or the Romans modified its structure by lowering the water level by 20 m through an artificial underground canal. What we do know is that the canal was built before the Cassia Cimina, during Roman Imperial times. In the 1500s, 17 18 the Farnese family re-opened the canal (today’s Rio Vicano), lowered the level of the lake by another 3 m, and regulated it with a weir that is still in place. The lowering of the water level provided farmland and favoured the settlement of farmers and shepherds. Today, the lake covers 12 km2 and is 49.5 m deep. Lake Bracciano. This lake is contained inside a caldera in the Sabatino Volcanic District (Rome), in central-northern Latium. With its area of 57 km2, it is the second largest in Latium and the eighth largest in Italy. This lake, which the Romans called Lacus Sabatinus, is round, with a maximum depth of 165 m and a volume of over 5 billion m3. Its shores host the villages of Bracciano, Anguillara and Trevignano, placed at regular intervals along the perimeter. About 800,000 years ago, a series of volcanic eruptions gave rise to the Sabatino volcanic system and, when they ceased, the craters filled with water and formed several lakes, the largest of which is Lake Bracciano. This vast depression formed as a result of the combined action of local faults and a large magma chamber. The latter, which was probably a few km below the surface and fed all the craters of the district, gradually emptied, and its top collapsed along distension faults which, in turn, produced massive lava flows. In successive stages, the area which today is a lake, became a large caldera, forming over a period between 400,000 and less than 150,000 years ago. Its only natural outlet is the stream Arrone, now dammed, and only used when in Lake Bracciano (Latium) flood. Otherwise, the water flows into the river Tiber. In Roman times, the lake provided drinking water. Trajan’s ancient aqueduct was restored by Pope Paul V in the early 1600s, and is therefore called “Acqua Paola”. It carries a type of water known for its low mineral content, which is taken to the fountain decorating the square on the top of the Janiculum hill in Rome. ■ Crater lakes Volcanic lakes are ideally shaped for collecting water. Their shape and impermeable bottom deposits, which prevent water from seeping in depth, give rise to crater lakes. Generally, crater lakes form in maars, volcanic craters produced by phreatic and phreato-magmatic explosions associated with interactions between the watertable and upwelling magma. The word maar (in German sea) originated in the Eifel area in northern Germany, where there are many such lakes. When magma comes into contact with underground water, energy is produced by the sudden change in the state of the water - from liquid to vapour - which causes it to increase in volume about 100 times. Magma rapidly wells upwards and is fragmented into minute elements the size of ash particles. The limestone basins of the Apennine chain provide the water supply that interacts with magma causing this type of eruption. Interactions may take place at various depths, and even surface Two types of volcanic lake: a caldera lake inside a crater, and a dammed lake, at the foot of a volcanic edifice 19 waters may contribute in marshes and shallow lakes. Lake water itself may favour an explosion. However, this hypothesis contradicts the evidence whereby most volcanic systems of this type are produced by a single explosion, and their craters are therefore called monogenic (having a single origin). Instead, there are maars Lake Nemi (Latium) where multiple eruptions occur over time (polygenic), and therefore the water of crater lakes plays an active role. This is why the level of crater lakes and the chemico-physical characteristics of their waters are carefully monitored. Morphologically speaking, maars are typically funnel-shaped, due to the destruction by explosions of pre-existing rock formations composed of deposits deriving from previous volcanic activity. Maars are volcanic formations that must be kept under control in the assessment of volcanic risk, because of their nature and genesis. The particular shape of maar craters makes them vulnerable to collapses and landslides. In addition, the phreato-magmatic explosions that produce them give rise to pipes which, although filled with various types of deposits, link the deep sub-volcanic areas to the surface. These conduits can be filled with gases which accumulate and flow through. The most dangerous gas is carbon dioxide, and its accumulation under the surface of maar lakes may produce sudden gas releases, like the one which recently occurred in Lake Nyos in Cameroon (1984), as well as lake water overturn and flooding. Maar lakes must therefore be continually checked for seismic activity, gas flow and flank stability. In Italy, lakes belonging to this category are the lakes of Mezzano in the Vulsini chain, Martignano and Monterosi in the Sabatini chain, Albano and Nemi in the Alban Hills, Monticchio in the Vulture, and Averno in the Phlegraean Fields. 20 Lake Mezzano (Latium) Lake Mezzano. Known by the Romans as Lacus Statoniensis, this lake lies in the Caldera of Latera which, in turn, is contained within other calderas. The crater hosting Lake Mezzano formed at the margin of the Caldera del Vepe, the most recent of the monogenic Latera calderas. The crater is a maar, which originated with the last explosion occurring inside the caldera about 160,000 years ago. The materials released were mainly fine ash and falling blocks of lapilli which accumulated around the crater, producing its margins. Although the inner margin 21 22 Reclaimed volcanic lakes If volcanic areas filled by lakes are reclaimed, they provide fertile land and lake silt rich in minerals. Many volcanic lake basins have therefore been exploited in this way by man. In the Vulsini mountains, the areas of Lagaccione, Latera and Montefiascone once hosted lakes. In the 17th and 18th centuries, in the Sabatini mountains, the marshy lakes Stracciacappa (in the past known as Lago di Straccio) and Baccano were reclaimed. In 1828, the Presidenza delle Acque e delle Strade (the authority in charge of roads and water supplies) opened a tunnel between lakes Martignano, Bracciano, Stracciacappa and Baccano. In his “Viaggio nel Lazio: la Tuscia e l’agro pontino” (Travels in Latium: the Tuscia area and the Pontine agricultural land) (1815-1818), the Italian mineralogist and geologist Giovanni Battista Brocchi wrote: “The inn at Baccano is located in a basin completely surrounded by reliefs, which was once Arnaldo Angelo De Benedetti a lake and perhaps, in ancient times, a crater. For as long as man can remember, it has been partially filled with water, which was released by creating an outlet that drained the water […] which previously used to stagnate in that location.” Other fossil lakes are those of Cese, Morto, Riano and Polline. In the Alban Hills, there once were lakes Ariccia, Castiglione, Gabii, Laghetto a Pavona, Prata Porci, S. Giuliano, Valle Marciana and Regillo. The last one was drained in the 17th century, and the plain that now covers the area is called Pantano secco, or dry marsh. In the Phlegraean Fields (Campania), although the Lake Agnano was reclaimed in 1870, seventy-five springs with temperatures up to 75°C have survived. North-west of the lake, near Astroni, historical maps show that there were once lakes called Grande, Cofaniello Piccolo, or Lago di Mezzo, and Cofaniello Grande, which are now very small. is very steep, almost vertical, the outer margin slopes gently, never exceeding an angle of 10-15°. The lake (area = 0.5 km2, maximum depth = 31 m) has an outlet (Fosso Olpeta) that crosses the caldera and flows into the river Fiora. Lake Martignano. Located east of Lake Bracciano, at 207 m a.s.l., this lake was known by the ancient name of Lacus Alsietinus. Although it has an area of 2.4 km2, it is very deep in proportion (60 m). The Alsietinus aqueduct, which was built in 2 BC to exploit the water of the lake that lacked natural outlets, supplied Augustus’s Naumachia (an artificial body of water Lake Martignano (Latium) surrounded by rows of seats, where ancient Roman spectacles resembling naval battles were performed) - at the foot of the Janiculum hill, together with Caesar’s Gardens, private farms, and the Janiculum fountain. The crater hosting the lake formed after at least three phreato-magmatic eruptions (there are three overlapping eruptive units, separated by palaeo-soils or erosive surfaces showing quiescent volcanic activity), and is considered, at present, the last active centre of the Sabatino Volcanic District. Underwater explorations in the lake have revealed that the area has been inhabited by man since Neolithic times. A wooden structure and evidence of fires 32 m below the current level of the lake show where its margin was, and large fossil oak trunks indicate that the lake underwent great changes in level over time. Lake Monterosi. Known in ancient times as Lacus Janulae, this lake is found in the Sabatino Volcanic District at 276 m a.s.l. It has an area of 0.3 km2 and a diameter of 600 m, but is only 7 m deep. It is also historically known as the place where, in 1155, Frederick I, called Barbarossa (Redbeard), refused to hold the stirrup of Pope Hadrian IV’s horse, thus causing a diplomatic incident. The lake formed after a single phreato-magmatic eruption of medium intensity. The area originally occupied by Lake Ariccia (Latium) Lake Albano. This lake (Lacus Albanus) is in the Alban Hills (293 m a.s.l., covering an area of 6 km2), 15 km from Rome. It is the deepest Italian crater 23 24 lake (-165 m). It is a polygenic maar and was therefore created by more than one explosion. Stratigraphic analysis of the volcanic centre of Albano, prompted by seismic risk for the city of Rome, has revealed that at least seven explosive eruptions occurred at intervals ranging between 70,000 and 29,000 years ago. The exact date of the last explosion is still unknown. Lake Albano is the most important and recent volcanic centre of the Alban Hills and morphologically belongs to the “lithosome of the Via dei Laghi”, i.e., the volcanic edifice created during the last phreato-magmatic phase of the volcano. In ancient times, the inhabitants of this area continually moved to various altitudes inside the rim of the crater as the level of the water in the lake changed. In 394 BC, the Romans built an artificial outlet about 1,200 m long, that both provided the area with water by means of wells, and drained it at a height of about 70 m from the lowest point of the crater rim. Recent analysis has shown that, during the Middle Bronze Age (1700-1350 BC), the lake frequently flooded the area near Rome, destroying human settlements. The artificial outlet built by the Romans is therefore considered the first modern hydraulic work in the world especially constructed to reduce volcanic risk. From the geological viewpoint, the volcanic strata inside the crater are made up of a bottom layer of volcanic material before the explosion of the maar, which have been completely removed within the crater area. Overlying this bottom layer are the stratified deposits of the seven eruptions, alternating with palaeo-soil that formed during quiescent periods. At present, the lake level is a few metres below the flood line of the draining outlet. The chemicophysical characteristics of the lake are constantly monitored due to the hazard posed by the great flux of carbon dioxide (CO2) from deep inside the area. Possible accumulation of CO2 at the bottom of the lake may cause the lake water to rise suddenly and jeopardise the towns nearby. Lake Nemi. The second largest lake of the Alban Hills, this lake has an area of 1.6 km2 and a depth of 37 m. Called Lacus Nemorensis by the Romans, it is 316 m a.s.l. and was produced by explosive eruptions that occurred about 150,000 years ago. The lake basin is composed of deposits of the multiple phreato-magmatic eruptions that produced the crater itself. The crater rim, which lies on top of deposits of previous volcanic activity, is made up of 2 overlapping ancient craters. Oval in shape, it is oriented along the meridian line. Ancient waste dumps have been found along its banks, marking the presence of Neanderthal man, as well as tombs going back to the Iron Age. In the 1930s, the lake level was lowered by 22 m, in order to raise two Roman galleys of the 1st century, which were later restored and displayed nearby. Unfortunately, in 1945, the retreating German army burned them down, and today their remains and two scale reconstructions are preserved Lake Albano (Latium) Lake Nemi (Latium) 25 26 in the Lake Museum. The shallow water of Lake Nemi, compared with that of the nearby Lake Albano, may be due both to its long inactivity and slow accumulation of lake deposits, and to the relative weakness of its eruptions, the deposits of which lacked sufficient energy to be propelled to a distance and therefore accumulated in the crater reducing its volume. The lake is drained by a tunnel (1,653 m long, for a total difference of 12.5 m) probably built by the Ariccia population. It was restored in 1927-28, when the lake was partially emptied. the eastern portion of the basin shows that the eruption was not powerful enough to send its products beyond the 500 m separating the crater rim from the bottom of the lake. Lakes Monticchio. The two small Monticchio lakes (600 m a.s.l.) also known as the “Vulture twins”, lie in the double central crater of the Vulture volcanic edifice, along its western flanks, near the towns of Rionero and Melfi (Basilicata). The Lago Grande (Large Lake) (area = 0.4 km2; maximum depth 38 m) and the Lago Piccolo (Small Lake) (area, 0.1 km2, maximum depth 35 m) are separated from each other by a strip of land about 216 m across, and formed 110,000 years ago. The entire volcanic edifice of the Vulture was created over a period ranging from 600,000 and 130,000 years ago, and is still affected by seismic activity and considerable fluxes of carbon dioxide. The deposits associated with the formation of the two lakes are composed of melilite pyroclasts the particle size of lapilli, in dune-like layers (Case Agostinelli, at most 4 m thick) outcropping to the west. The absence of the same deposits in Lake Averno. This lake is found in the Phlegraean Fields volcanic region, between Monte Nuovo and Monte Grillo. It is elliptical in shape and has a maximum depth of 35 m. The lake basin is an extinct volcano that was created about 4,000 years ago. In 38-36 BC, the dense forest surrounding the area was dramatically modified by Marcus Agrippa, the Roman statesman, who cut the trees down and converted the lake into a naval port (Portus Iulius). In 1538, the eruption of the Monte Nuovo changed the shape of the lake completely. The name Averno derives from the Greek Aornon, meaning “area without birds”, giving rise to the legend that no bird could fly across the lake and live because of its poisonous sulphurous vapours. The lake was also mentioned by Homer and Virgil, who represented it as the entrance to hell. It was thought to be home to the Giants and of the population of the Cimmerians, who were associated with the so-called “Catacomb” culture and were believed to live in caves and flee the sunshine. The Greeks later identified this area as that described by Homer in the Odyssey. The lake was thought to be bottomless. Lago Piccolo di Monticchio (Basilicata) and the abbey of San Michele Lake Averno (Campania) 27 Hydrochemistry SILVIA ARISCI · LAURA LEPORE · MICHELA ROGORA From the hydrological viewpoint, lakes are standing waters which gather in basins within the Earth’s crust, and which are not fed by the sea. The sloping funnel-shaped area surrounding a lake collects rain and is called its catchment basin. The watershed is the highest level in the catchment basin, so that any water falling beyond it does not flow into the lake. Catchment basins greatly affect aquatic ecosystems, as their area determines the volume of collected water, their mineralogical composition influences the basic chemistry of lakes, and their plant cover influences the division of flowing and evapo-transpiring water. Drained water Lake Bracciano (Latium) is charged not only with organic and inorganic matter, but also with pollutants produced by the main uses to which they are put (agricultural, industrial, urban) in the area. In addition to water flowing into the lake from the catchment basin, there are underground springs from the watertable and, in volcanic lakes, seepage of groundwater from the volcanic edifice, which has a very particular chemistry. In this case, the lake may be the emerging portion of a larger catchment basin. The geographical position of lakes affects their physical behaviour: those in northern Europe freeze in winter and cannot exchange oxygen with air; those in tropical and temperate areas have warm surface layers that float on deeper layers, hindering water mixing and oxygenation of the bottom. Alpine lakes are at their highest level in late spring or summer, when snow melts; lakes in central Italy are at fullest in winter, when rain is more abundant. The Italian average precipitation is 100 cm, and evaporation is similar. This means that the rain and snow falling on Italian standing waters replenish, on average, only loss due to Lake Vico (Latium) 29 30 The properties of water Silvia Arisci · Laura Lepore · Michela Rogora Water is the essence of life on Earth and the basic chemical component of any living organism. It also regulates the metabolism of lakes, with its particular properties like density, high thermic capacity and behaviour in its different states - liquid, solid and gaseous. The unique properties of water are due to its molecular structure. The formula of water consists of two atoms of hydrogen and one of oxygen. The three atoms are bonded, with the oxygen atom between the two hydrogen atoms. However, the three atoms do not lie in a straight line the two hydrogen atoms bend towards each other. This three-dimensional structure is therefore unsymmetrical, the oxygen atom having a partially negative charge (δ-) and the two hydrogen atoms partially positive charges (δ+). Opposite the hydrogen atoms are two electronic clouds of negative electrification, which attract the hydrogen nucleus of an adjacent water molecule to form what is called a hydrogen bond. When water freezes and becomes solid (ice), its molecules form an ordered structural lattice: one water molecule collects its nearest four neighbours and arranges them about itself in a tetrahedral configuration. The peculiarity of ice is that the intermolecular distance is higher than in the liquid state, and therefore density in a solid state is lower than in a liquid state. Maximum density is obtained at 3.98°C and pressure of one atmosphere. Below this temperature, density decreases again, until freezing point (0°C) is reached. The most important consequences of this phenomenon are that, in winter, lake surfaces develop floating ice sheets, rather than freezing solid from top to bottom, and that the temperature of the water below remains almost constant, due to the low thermal conductivity of ice itself, which prevents heat from being released into the atmosphere. Water density varies with temperature, pressure and salinity. It increases with pressure (1 atmosphere every 10 m in depth) and with the increased specific weight of dissolved substances. Hydrogen bonds between molecules also determine the high specific heat of water, i.e., the quantity of heat one gram of water needs to increase its temperature by 1°C. The high thermal capacity of water explains why thermic variations near large lakes are unlikely, and why extensive bodies of water can release great quantities of heat accumulating in warm periods into the atmosphere. Strong intermolecular interactions influence the viscosity of water, i.e., the great resistance water opposes to motion within it (775 times that of air). δ+ O LAKE BASIN VOLUME DEPTH AREA EMIS. FLOW REC. RATE km2 km3 m km2 m3/sec years Garda 2350 50.35 346 370 59.5 Iseo 1842 7.60 251 62 59.4 4 Como 4572 22.50 410 146 158.0 4 Maggiore 6559 37.50 370 212 297.0 4 10 0.46 165 6 - 47 - 15 Albano Nemi 11 0.03 34 Trasimeno 376 0.59 6 124 0.9 21 Bolsena 273 9.20 151 114 2.4 120 41 0.26 12 0.5 17 147 5.05 165 57 1.2 137 0.004 38 - 110 Vico Bracciano Monticchio Grande 4 49.5 1.6 27 0.4 Characteristics of main subalpine and some volcanic lakes in central Italy Compared with subalpine lakes, volcanic lakes have far longer recharge rates, due to the weak hydrological conditions caused by their origin. Lakes Bolsena, Vico and Bracciano, for instance, lie at the top of their volcanic edifices, and their catchment basins are therefore extremely small, especially when compared with their volume. O H δ− evaporation. Excess water, i.e., that drained by outlets, corresponds to the supply provided by the basin, so that lakes fed by extensive basins, like Alpine ones, have large outlets, and lakes supplied by small basins, like volcanic ones, have small outlets. The theoretical recharge rate is the ratio between the volume of water in the lake and the quantity of water drained by an outlet in one year. The recharge rate is thus an index of the capacity of the lake to drain through its outlet some of the pollutants flowing into the basin. The table below compares the main morphometric and hydrological characteristics of the largest Italian lakes (Garda, Iseo, Como, Maggiore) with those of some volcanic lakes. O H ■ Temperature and oxygenation of lakes O O O A water molecule and its tetrahedral configuration in solid state (ice) From the thermic viewpoint, the volcanic lakes of central Italy are temperate. In winter they are homoiothermic (i.e., they have a relatively uniform temperature from surface to bottom), and therefore all the water has the same density. In spring, the sun heats only their upper layers of water, as shown by the fact that infrared light, thermically the most efficient, can only penetrate the topmost 31 water layers. Wind can give rise to currents that distribute heat at deeper levels. As summer sets in, the temperature rises and wind-generated currents can overturn water to the bottom of the lake only at night and if the lake is not very deep. When density differences are too great for the wind to maintain homoiothermic conditions, temperature stratifies vertically between May and October. In this period, the water is divided into three layers: ● A warm upper layer (epilimnion) ● A cold, deep layer (hypolimnion) ● A thin layer separating the epilimnion from the hypolimnion (thermocline, or metalimnion), in which water undergoes sharp temperature changes. As temperature decreases in autumn, the surface water cools and mixes with deeper water, giving rise to winter mixing, a situation that maintains homoiothermic conditions in the lake between January and March, and also provides a certain chemical homogeneity. This phenomenon is particularly important for the oxygenation of lake waters, especially deep layers where dissolved oxygen is gradually consumed by biological decay, giving rise to initial depletion (hypoxia) to total lack of oxygen (anoxia). These conditions make life in the lake impossible for most organisms. Dissolved oxygen is essential for life in lakes. Its main source is the passage of oxygen through the air-water interface, and also by biological production (photosynthesis). Oxygen can return to the atmosphere through diffusion, be consumed through respiration by aquatic organisms, or be oxidised by chemical processes. The balance between these exchanges produces the spatial distribution and time variations of oxygen in a body of water. In some lakes, deep water never circulates with the other layers. This condition, called meromixis, is common to several volcanic lakes. In meromictic lakes, isolation of the deepest layers, oxygen depletion, and processes of anaerobic decay produce compounds like hydrogen sulphide (H2S) (which produces a typical, Lake Bracciano (Latium) unpleasant odour), ammonia (NH4+) and methane (CH4). Bottom sediments turn black or grey, and organisms living at the bottom of lakes are negatively affected. ■ Other gases and ion compounds Temperature (C°) 0 Depth (m) 32 5 10 15 20 25 0 5 10 15 20 25 30 February April June August October December 35 40 Yearly temperature distribution in water column of Lago Grande of Monticchio (Basilicata) 30 From the chemical viewpoint, lakes are open areas interacting with the atmosphere at the surface, the underlying bedrock, and water supply underground and the surrounding area. Gases in the atmosphere, especially oxygen and carbon dioxide, dissolve in water according to Henry’s law, which states that the amount of gas absorbed by a liquid is proportional to the partial pressure of the gas upon the liquid, by means of a constant value (k) called Henry’s constant. The concentration of dissolved gases decreases as temperature increases, and is influenced by chemical and biological processes occurring in liquid. Carbon dioxide (CO2) is supplied to lakes through exchange with the atmosphere at the surface, in rain, which collects CO2 as it falls, and is released by animal respiration. Carbon dioxide is very soluble in water and forms carbonic acid, which in turn dissociates, raising the concentration of hydrogen ions (producing bicarbonate and carbonate ions). These chemical reactions, each of which is controlled by a balancing constant, are called carbonate and bicarbonate balance, and determine the pH value of water at a given 33 temperature (generally between 7 and 9). The carbon dioxide system therefore serves as a buffer, because any change in pH will cause a shift in the system that will offset that change. Water can dissolve not only gases, but also organic and inorganic matter which is either polar or subject to polarisation. Salinity varies greatly and depends on interactions with the atmosphere, drainage from the surrounding area, and exchange with sediments inside the lake. The composition of natural water is controlled by the bedrock and soil of the catchment basin, precipitation, crystallisation and evaporation, and biological processes inside the body of water (production-respiration). Volcanic lakes have far higher salinity than other Italian subalpine lakes. Their total salinity ranges between 7000 and 11000 µeq/l - much greater than, for instance, that of Lake Maggiore (3000 µeq/l) Salinity is computed from the cations (positively charged ions) of alkaline and alkaline-earth metals like calcium (Ca++), magnesium (Mg++), sodium (Na+) and potassium (K+), and anions (negatively charged ions) like carbonate (CO3--), bicarbonate (HCO3-), sulphate (SO4--) and chloride (Cl-). According to their salinity, natural waters are divided into two groups: soft water, with low salinity when it erodes igneous rocks (poorly soluble), and hard water, containing large amounts of alkaline and earth metals deriving from the underlying calcareous bedrock (soluble). Natural water, the ion spectrum of which is given by the bedrock of its catchment basin, is typically rich in calcium 34 Lake Bracciano (Latium) Lake Albano (Latium) 35 36 and bicarbonate, and the relationships between ions are: Ca++>Mg++=Na+>K+ for cations and HCO3-> SO4-->Cl- for anions. In central Italian volcanic lakes, this order is only partially maintained. Supply from the sea is very important for these lakes, which have higher concentrations of sodium and chlorine than standing waters far from the coast. In volcanic lakes, in addition to marine aereosol, the main source of solutes is bedrock and soil runoff from the catchment basin, which supply bicarbonate, calcium, magnesium, sulphate and sodium. Among anions, bicarbonate dominates, followed by chloride compounds, which in volcanic lakes are more frequent than sulphate compounds. These lakes generally have a good buffer system, which produces minimal variations in pH values at various depths. Cations like calcium, magnesium, sodium and potassium have similar concentrations, and sodium is slightly higher in lakes Bolsena and Bracciano. Lastly, nitrates are barely present in volcanic lakes (<10 µeq/l), although they are found in high concentrations in subalpine lakes (50-60 µeq/l). Nitrates mainly derive from precipitation which, in northern Italy and densely populated areas, is particularly rich in nitrogen oxides. Volcanic lakes are rich in organic nitrogen produced by a complex cycle of metabolic assimilation of atmospheric nitrogen by heterotrophic bacteria. They transform nitrogen into ammonia, which either returns to the atmosphere or enters the plant-herbivore-carnivore food-chain. Decomposition then triggers the cycle again. µeq/l 0 1000 2000 3000 4000 MAGGIORE 5000 ■ Algal nutrients: phosphate, nitrogen and silica In addition to the inorganic constituents mentioned above, other inorganic compounds are particularly important in lake waters. They are usually referred to as nutrients, because they are exploited in various ways by algae and micro-organisms. Silica (SiO2) is essential to diatoms, a group of algae that use this compound to produce their frustules (siliceous shells), thus regulating its dissolved concentration. Dissolved silica in lakes undergoes great seasonal variations, i.e., it accumulates in winter and decreases dramatically in spring, when diatoms bloom. Phosphorus, in its highly oxidised forms like orthophosphate (PO4---), plays an active role in biological cycles as a nutrient for lake organisms. Phosphorus, like silica, undergoes seasonal and spatial variations inside a lake: in the upper portion, where photosynthesis occurs, it decreases when algal development peaks; in summer, it increases in deep water, due to decay of biological material sinking down from the upper layers. Nitrogen, in the form of nitrate (NO3-), nitrite (NO2-), ammonia (NH4-) and organic nitrogen, is an essential nutrient for several organisms. In lakes, the dynamics of nitrogen and phosphorus are associated with biological processes, and therefore with consumption by algae and bacteria. The 6000 Ca++ Mg++ ALBANO Na+ BRACCIANO NEMI K+ HCO3NO3- VICO BOLSENA Average ion concentrations in some volcanic lakes and in Lake Maggiore SO4-Cl- Lake Mezzano (Latium) 37 passage between the various nitrogen compounds is regulated by microorganisms. In summer, when temperature and light are sufficient for intense algal growth, the production of nitrogen is controlled by the availability of nutrient compounds. In particular, as it is present in lower quantities than those required by algae, it thus limits their growth. According to nutrient contents, lakes are divided into trophic levels, from oligotrophic (few nutrients and low algal productivity), mesotrophic (an intermediate stage), to eutrophic (very productive lakes rich in nutrients). Trophic conditions are closely associated with the chemico-physical characteristics of lake waters and the biocoenoses populating them. Excessive nutrient enrichment (especially by phosphorus) due to human disposal, is called eutrophication. This process diminishes water quality and affects the use of the lake as a source of drinking water, for irrigation, bathing, and other uses. In the 1970s, results of monitoring analyses on Latium volcanic lakes by the CNR (Consiglio Nazionale delle Ricerche - Italian Research Council) had revealed low concentrations of phosphorus and low productivity of the lakes, most of which were classified as oligotrophic. Recent increases in phosphorus caused by human activities, has since positioned the lakes at the highest trophic level. The quality of their water has therefore deteriorated (see chapter on conservation and management). 38 µg/l 0 100 200 300 400 ALBANO BRACCIANO BOLSENA NEMI 3000 MONTICCHIO GRANDE MONTICCHIO PICCOLO Lake Bracciano (Latium) surface bottom Average concentrations of total phosphorus (in µg/l) at the surface and bottom of some volcanic lakes 39 ■ Traces of heavy metals 40 41 Another typical characteristic that distinguishes volcanic lakes from other bodies of water is their heavy metal content. The heavy metal contents of large subalpine lakes (like Lake Maggiore) are far lower, almost undetectable. The water of volcanic lakes is generally rich in dissolved metals, especially boron and strontium, both of which are indicators of geothermic activity. Other commonly found metals, like iron, manganese and zinc, which come from the underlying bedrock, are far more frequent in lakes Bolsena, Lago Grande of Monticchio (Basilicata) Albano and Monticchio than in the Maggiore. The reason for this characteristic lies in the lithology of the basins, generally composed of incoherent lithoid tuff and effusive rocks. These porous, permeable rocks facilitate runoff by rain and enrich lake water with metals. BOLSENA MONTICCHIO GRANDE 130 m 20 m 9 9 6 0 15 182 14 5 Boron 390 395 86 94 53 64 4 - Barium 35 38 13 22 84 289 10 10 Copper - 4 - - - 0.4 0.5 0.2 Iron - 25 - 75 46 5900 8 4 Manganese - 21 - 105 99 3572 1 5 Zinc - 20 - - 2 2 1 1 42 41 4 4 3 3 1 - 420 450 753 893 479 630 208 227 Lithium Strontium 160 m 0m 35 m MAGGIORE 0m Alluminium Lake Nemi (Latium) ALBANO 0m 360 m Average concentrations (in µg/l) of main cations and anions in some volcanic lakes and in Lake Maggiore Phytoplankton GIUSEPPE MORABITO At first sight, volcanic lakes appear to be special environments, as they have the characteristic shape of a crater with high rims, so that they actually look like large bowls.However, examination of samples of their waters and of the microscopic algae (phytoplankton) contained in them will not surprise limnologists and naturalists greatly. Over the seasons, volcanic lakes are not subject to the extreme physical and chemical conditions that would enable only some selected algal species to survive. On the contrary, these habitats are usually inhabited by the same species that normally colonise completely different kinds of lakes. However, some Lake Mezzano (Latium) of these species find the ideal living conditions that make them dominant precisely in volcanic lakes, due to the optimal combination of physical and chemical parameters that is usually found in such lakes. This section analyses the factors that lead to the development of planktonic algae in lakes, highlighting the main characteristics of volcanic lakes associated with several variables, and aims at providing a general overview of the habitats in which these algae live. Readers will then understand why some types of phytoplankton can be dominant in volcanic lakes. ■ Factors regulating the growth of phytoplankton There are many variables controlling the growth of phytoplankton in lake water, and they interact in often baffling ways. Yet, like all plant species, also phytoplankton depend on certain light and temperature conditions, as well as sufficient nutrients. Ceratium hirundinella 43 45 44 Lake Vico (Latium) Dinoflagellates of the genus Peridinium Temperature and light. Analyses by phytoplankton experts reveal that, in lake environments, there are two essential conditions for the life of single algal species - temperature and sunlight. Just as on land, also in aquatic ecosystems temperature and light must fall within the values compatible with the life of plant species, and the range of these values may differ between species. In addition, sunlight penetrating the water column produces thermic energy and gives rise to layers of water with differing thermic characteristics. Seasonal temperature variations in lakes follow a regular cycle, alternating periods of total water overturn and periods of water stratification. Differences in temperature produce differences in water density, and these small variations drastically affect the life of algae which, having a slightly higher density than water, must contrast their natural tendency to sink towards the bottom of the lake. The survival of phytoplankton in lakes therefore relies on mechanisms that enable them to slow sedimentation down. For instance, algae can resort to adaptations such as changing shape to increase their cell area, forming colonies, or regulating their position inside the water column by means of propelling devices (flagella or mobile bristles), gas-filled vacuoles, or the synthesis of compounds with low specific weight. As regards sunlight, phytoplankton behave in various ways: some algae thrive in broad sunlight on the surface, whereas others prefer to live a few metres below. Both the quantity and quality of sunlight are important, and depend on the nature of particles in suspension. These particles are like coloured filters that select some wavelengths of the light spectrum, endowing the lake with precise optical features and shades of colour, generally ranging between blue and green, according to the quantity of growing phytoplankton. Lakes can be brown or yellowish in areas with great quantities of complex plant organic matter. Phytoplanktonic biocoenoses adapt to the qualitative variations of the light spectrum by selecting species with photosynthetic pigments (molecules that capture light energy to carry out photosynthesis), which can better exploit the available light in the environment. Early limnologic studies describe Italian volcanic lakes as very transparent, places where light - in qualitative and quantitative terms - is not a limiting factor for the growth of phytoplankton. The excellent optical quality of water is an original characteristic of these lakes, and it is probably associated with their morphology. Lakes are often found at the top of volcanic edifices, where water from rivers is scarce compared with the area of the body of water. 46 Large quantities of water carrying suspended solids are therefore unlikely to flow into the lake and alter the optical properties of its waters. Lake transparency enables light to penetrate deep down into the water column. For instance, in the late 1960s, a limnologic study by the Istituto Italiano di Idrobiologia of Pallanza (today Istituto per lo Studio degli Ecosistemi of the CNR) showed that lakes Bolsena, Bracciano and Vico had euphotic areas (of, relating to, or constituting the upper layers of a body of water into which sufficient light penetrates to permit the growth of green plants) down to 30 m. Some algae find these conditions in volcanic lakes particularly favourable for their development. In volcanic lakes, this species replacement is also affected by the morphology of the basin, which influences the processes and quantity of algal nutrients available. Hence, when water overturn occurs in shallow volcanic lakes like Monterosi and the Lago Grande of Monticchio, even the mineralised inorganic nutrients sedimented at the bottom of the lake can return to the surface, to be consumed by plankters. By contrast, deep volcanic lakes are unlikely to undergo complete water overturns, and inorganic nutrients deposited at the bottom remain inaccessible to phytoplankton. Nutrients. The availability of nutrients, especially limiting ones, is perhaps the third factor that most affects algal growth. Since the early 1970s, research focusing on eutrophication has indicated phosphorus as the main limiting nutrient of phytoplankton in lakes. Most volcanic lakes have evolved from original oligotrophic conditions to mesotrophic or even eutrophic conditions. These modifications have also deeply altered the original phytoplankton communities, which have lost their oligotrophic-loving species which were replaced by others better adapted to the new environmental conditions. The general introduction of the previous section provided a limnologic analysis of volcanic lakes, and described the conditions algae encounter in colonising them. Summarising briefly, an alga in a typical volcanic lake is in an environment where temperature undergoes profound seasonal variations. In summer, this produces layers of water all greatly differing in temperature from one to another. These lakes are also transparent, so that sunlight can penetrate to considerable depths, their water overturn is very slow, and supply from the catchment basin is scarce. Nutrient availability varies according to the trophic state and morphology of the basin, just as in other lakes. Lago Grande of Monticchio (Basilicata) Cyclotella comensis ■ Structures of phytoplanktonic associations 47 The specific components of phytoplanktonic associations undergo seasonal modifications caused by changes in environmental conditions. In late winter, when the water column is mixing and light becomes more intense, diatoms start developing. These are typical pioneer algae, which grow rapidly in the water still free of other algal groups thanks to their great capacity for nutrient assimilation. Diatoms have a siliceous cell wall called frustule which contains their cells - making them very heavy - and therefore causing them to sink rapidly to the bottom, far from the upper layers illuminated by the sun. This is why diatom growth is closely associated with the period of maximum lake water turbulence, because water mixing prevents them from sinking. In Italian volcanic lakes the dominant diatom groups are the Centrales (discshaped) - as emerges from analyses conducted on lakes in Latium. According to the scientific literature, in the years prior to trophic deterioration, the dominating genus was Cyclotella, with species C. comensis, C. kuetzingiana and C. ocellata. In several basins, when higher trophic conditions set in, Cyclotella species were replaced by Stephanodiscus species (S. parvus, S. minutulus, S. hantzschii), another genus of Centrales which, unlike Cyclotella, prefers habitats with higher nutrient contents. The alternation of these two genera was clearly highlighted by palaeolimnological research carried out on volcanic lakes in Latium, which reconstructed trophic evolution by analysing diatom frustules in lake sediments. The research also identified the community of the diatom components in lakes Nemi and Albano since the Late Pleistocene, and revealed variations of diatoms in eutrophic and oligotrophic waters in periods even prior to human activity. They were probably due both to climatic events that caused the water temperature to rise, and to periods of geothermal activity which influenced the availability of algal nutrients. Unfortunately, palaeo-limnological information about the structure of past phytoplanktonic associations can only be obtained from algae that leave fossil traces, i.e., diatoms and golden algae (Chrysophyceae). According to palaeolimnological data about recent algal communities, the latter count only a few species in volcanic lakes. Generally speaking, both diatoms and golden algae are spring phytoplankters, not only in volcanic lakes. Environmental conditions change greatly between spring and summer, when thermic stratification occurs. The nutrients that had started to mix in late winter and had supported the spring development of diatoms, have almost completely disappeared from the epilimnion. Cyanophyceae iIndividuals/ml 48 Diatoms Cryptophyceae Chlorophyceae 60000 50000 40000 30000 20000 10000 0 J Stephanodiscus minutulus F M A M J J A Seasonal trend of the main phytoplanktonic groups in Lake Nemi (1982) S O N D 49 The blue-green alga Planktothrix rubescens The species Planktothrix rubescens typically forms dense deposits in the lake thermocline, i.e., the layer of water in which the highest temperature gradient is measured during summer water stratification (temperature declines at least 1°C with each metre of increase in depth). P. rubescens is a frigo-stenotherm (resisting only slight changes in temperature) which thrives at around 15°C. It is also sciaphilous, and therefore prefers dim, poorly illuminated areas of the lake. From the trophic viewpoint, P. rubescens lives in environments rich in nutrients. Chlorophyll (µgl-1) 0 0,5 1 0 10 Giuseppe Morabito The combination of cold, dim and nutrient-rich water occurs in the thermocline of deep lakes, between 10 and 15 metres in depth, where light has already diminished, and the temperature is quite low. The temperature gradient causes differences in water density, which favour the accumulation of organic matter in the thermocline, where it is decomposed and mineralised. P. rubescens is also favoured by special photosynthetic pigments, which colour it red and enable it to make efficient use of even low light radiation. It uses its intracellular gas vacuoles to regulate its position in the water column. 1,5 2 2,5 15 20 25 0 Depth (m) 50 5 10 15 20 25 30 35 40 45 °C 5 Planktothrix Temperature Relationship between Planktothrix rubescens and water temperature as depth increases Nonetheless, the thermocline - where organic matter remains for some time, due to temperature and density gradients - can store nitrogen and phosphorus. Differences in water density in the thermocline are physical barriers to the sedimentation of particle-sized organic matter, which is mineralised as it moves through this layer of water, and can therefore produce deposits of nutrients used by growing algae. Obviously, not all algae can use these reserves and, in this season, the numbers of diatoms decrease for three reasons: firstly, because of thermic stratification and the ceasation of the Planktothrix rubescens turbulence which kept them afloat; secondly, because the silica used to form their frustules has been consumed and the lack of this element now limits their growth; and thirdly, because they have become prey to herbivorous zooplankton, especially water fleas (Cladocera), which drastically reduce spring algae, giving rise to a seasonal phenomenon whereby water is particularly clear, due to algal removal by zooplankton. Diatoms therefore leave the water free for the development of other algae but which? In the 1930s and ’40s, research on the pelagic (open water) flora of lakes Albano and Nemi revealed that the most successful algae in summer phytoplanktonic associations in volcanic lakes are dinoflagellates. In particular, one of the most common is Ceratium hirundinella, whose fascinating shape is typically found in lake phytoplankton. There are two key factors giving it competitive advantage, one being its flagella, which enable it to swim in the water column in search of the best conditions for its development. Thanks to this capacity for independent movement, Ceratium can migrate through well-illuminated surface water and carry out photosynthesis, and then move to the thermocline to feed. The other decisive factor lies in its size, which enables it to store large amounts of nutrients whenever it can. This is particularly helpful in environments poor in nutrients, like Italian lakes prior to the 1960s. 51 52 Volcanic lakes in which trophic conditions have deteriorated show a decline in the total numbers of dinoflagellates, which are replaced by cyanobacteria (blue-green algae) in summer algal communities. The species which are most successful in volcanic lakes depend on the type of environment. Deep volcanic lakes have almost exclusively been colonised by Planktothrix rubescens, and shallow lakes host taxa of chroococcoid cyanobacteria (for istance Microcystis, Woronichinia, Merismopedia). Planktothrix rubescens is a bluegreen alga that gathers in filamentous Cyanobacteria of the genus Woronichinia colonies, and is one of the bestknown species, due to its occurrence worldwide. Its blooms, which turn lake waters red, are also very well-known because they are the clearest mark of deteriorated trophic conditions in many European deep lakes. Its ecophysiological characteristics therefore also allowed it to colonise Italian deep lakes successfully (see p. 50). A water column that is clear down to great depths is the optimal ecological niche for the development of P. rubescens. The transparency of volcanic lake water guarantees an euphotic zone deep enough to include the thermocline, which further enhances colonisation by this species in volcanic lakes like Albano and Nemi. Shallow volcanic lakes seldom have a stable thermocline and great transparency because they are subjected to total overturn by strong winds. Turbulence can affect the deepest areas of the lake, sediments may move to the upper layers of water, and the water becomes more turbid. In addition, shallow lakes may have high water temperatures even deep down. These conditions therefore hinder the growth of Planktothrix and favour a particular type of cyanobacteria, the Chroococcaceae, which form colonies of irregular shape (Microcystis), round (Woronichinia), or rectangular (Merismopedia). Their cells are contained in a gelatinous matrix, enabling them to float easily and return to the surface after water overturn. Most Chroococcaceae can withstand high temperatures and radiation, and may therefore colonise the upper layers of water very successfully. As these bodies of water are quite shallow, when water overturns, great quantities of nutrients sedimented at the bottom float to the surface, giving rise to cyanobacterial blooms that look like large mats. Blooms can often be seen in the Lago Grande of Monticchio. The development of cyanobacteria in volcanic lakes has also affected the food-chain, bringing, for example, Eudiaptomus padanus etruscus - a frequent copepod in these habitats before eutrophication - to complete extinction. Microcystis wesenbergii Let us now summarise the main topics analysed in this section. Firstly, the dynamics of phytoplanktonic communities in lakes is regulated by several physical, chemical and biotic factors, the relative importance of which depends on the seasons and the environment. Among the variables determining the composition of algal species, recent analyses on phytoplankton ecology have shown that physical factors are the most important in determining species alternation. This is why the description of phytoplankton in volcanic lakes focused on the adaptations of algal species to light and temperature variations. However, this is not the only reason, because the physical environment of volcanic lakes has characteristics that distinguish these habitats from other lakes and make them suitable for colonisation by algae which, nonetheless, are not exclusive to this type of freshwater. The scientific literature on the phytoplankton of volcanic lakes has highlighted the fact that many of these environments are very vulnerable to man’s influence. Their slow hydrological recharge favours the quick accumulation of algal nutrients, and modified chemical characteristics give rise to sometimes profound changes in the structure of phytoplanktonic communities, with the development of species that may severely compromise how humans use these waters. Typical examples are the blooms of potentially toxic cyanobacteria which have recently occurred in volcanic lakes in Latium. 53