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Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali Progetto Biodiversità: Integrare lo sviluppo del territorio con la conservazione della biodiversità in provincia di Trento. Testo approfondito ABSTRACT The exponential increase of the human population, the environmental changes associated with it, and global warming are causing a dramatic rate of extinction and re-distribution of plant and animal species. In response to these trends, there is urgent need for studies of biodiversity and factors affecting it at the global, regional and local level. In addition, the scientific approach to conservation has shifted from the protection of single species or sites to preservation of overall biodiversity. There is reasonable agreement in the international scientific community about the main measurable components of biodiversity. These include: (1) diversity of genes and of characters; (2) diversity of species; (3) diversity of ecosystems; (4) functional diversity, i.e. diversity of ecological processes. To date, quantitative studies of biodiversity have been few, often at coarse spatial scales, and based on only one component of biodiversity. The few studies conducted in Italy are often below the international scientific standards. The Trento region still lacks any quantitative study of biodiversity. Project BIODIVERSITÀ is aimed at filling such gaps. Main objectives of the project will be to: 1. estimate the four biodiversity components, integrating them with vulnerability indexes; 2. build statistical models aimed at predicting field-recorded biodiversity estimates from topographic, vegetational and land-use characteristics; 3. apply the predictive models and field-recorded estimates to the whole Trento region, so as to create a GIS map of regional biodiversity; 4. highlight potential biodiversity hotspots within the Trento region; 5. Build statistical models capable to predict the loss of biodiversity associated with environmental changes driven by local development, and propose sustainable development guidelines aimed at integrating economic, touristic, selvicoltural, and agro-pastoral development with the conservation of biodiversity. Data used to build biodiversity estimates will include: 1. distribution databases of birds, mammals, reptiles and amphibians, completed by fish data from an in-press publication; 2. intensive population studies of 11 predatory species and their main prey, chosen as indicators of environmental health of the main regional macro-habitats, sampled at various levels of their trophic pyramid; 3. progressive censuses and sessions of ringing of migrant birds, so as record estimates of “transient biodiversity”. Results of the project will be elaborated so as to be potentially integrated within the P.A.T. management systems, in agreement with the units deputed to such duties. Results will be published on international scientific journals and made public through conferences, an internet site, books for the public and a museum exhibition. 1 Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali Progetto Biodiversità: Integrare lo sviluppo del territorio con la conservazione della biodiversità in provincia di Trento. Testo approfondito 1. INTRODUCTION 1.1. What is biodiversity? The exponential growth of the human population and the environmental changes deriving from it are causing a rate of extinction of animal and plant species judged to be 1000 times higher than the one characteristic of the current geological period. In addition to such “biodiversity crisis”, global warming is predicted to cause major redistribution of organisms on the planet. Such processes make the study of biodiversity and of the factors affecting it extremely urgent on a global, regional, and local scale. In association with such events, in the last decades the scientific approach to conservation has shifted from the protection of single species or sites to the definition of wider, more diffuse targets, which include the whole biological diversity of an area. The importance of preserving the whole biological diversity of an area has been justified through various instrumental or ethical criteria, associated with the economic, scientific, educative, ecological, spiritual, recreational and esthetical value of biodiversity. Biological diversity, or “diversity of life”, commonly defined as biodiversity, is a complex concept which has assumed in time scientific, socio-political, and economic meanings. Biodiversity can be defined as the genetic, taxonomic and ecosystem variety in living organisms of a given area, environment, ecosystem or the whole planet. It must be noted how this is only one of the possible definitions. In general in ecology, the term “diversity” is composed of two components: richness, the absolute number of categories in a sample (e.g. number of species), and evenness, the degree of uniform frequency of sample units within each category (e.g. the number of individuals within each species). For example, a very diverse community will be composed of a high number of species, each one represented by a similar number of individuals. When this concept is applied to biodiversity, we are faced with the additional problem of incorporating a measure of relatedness, or genetic similarity, among the elements of the sample system. For example, a community composed of two butterfly species will be less diverse than a community composed of one butterfly and one vole species. In an ideal world, biodiversity would be measured by one unique index incorporating richness, evenness and genetic similarity. In the real world such an index does not exist; the complexity of biodiversity makes it a conceptual entity, difficult to define quantitatively and measurable only in terms of some of its components. If there is disagreement about quantitative definitions of biodiversity, more consensus exists about the hierarchical levels of measurement of biodiversity. With little variation, biodiversity has been subdivided into the following components: genetic diversity and/or character diversity; diversity of species or of higher taxonomic categories (e.g. genera or families); diversity of habitats and ecosystems. These three categories are related to the spatio-temporal pattern of occurrence of biodiversity elements. They ignore all those ecological processes which cause the observed pattern (Gaston 1996a), where by ecological processes we intend all those activities associated with the interactions among organisms and among organisms and their environment. Such fourth component of biodiversity has been defined as “functional biodiversity”, i.e. the diversity of all those processes, such as genetic flow and nutrients cycle, which determine the distribution pattern of biodiversity elements, such as individuals, populations, or species. To date, most biodiversity studies have been organized following the above cited hierarchical structure or some variation of it. Below, we briefly review the most commonly employed methods of data collection and analysis within each of the four biodiversity components cited above. 2 Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali Progetto Biodiversità: Integrare lo sviluppo del territorio con la conservazione della biodiversità in provincia di Trento. Testo approfondito 1.2. Methods of biodiversity measurement Genetic diversity and character diversity Measuring genetic diversity in a direct way at the level of chromosomes, enzymes and DNA is practically impossible for studies at the regional level, both in terms of manpower and laboratory analyses and of economic costs. New techniques have thus been recently developed to provide indirect measures of genetic diversity, based on the diversity of characters among species, genera, and families. We define as characters all those biochemical, morphological, or behavioural traits on the basis of which organisms can be taxonomically separated. Such differences, at the heart of the science of taxonomy, are considered to be the exterior, phenotypic expression of information encoded into the gene pool of an organism. Phenotypic characters diversity can thus be used as an indirect estimate of genetic diversity. As it is generally impossible to count all the characters of all the sample organisms, the standard method of measurement of character diversity consists of predicting the distribution of such characters among organisms based on their genealogy, i.e. based on their phylogenetic tree. Such analyses yield diversity estimates which take into account the evolution history, the phylogenetic distance, and thus the character convergence or divergence among species or among other taxonomic categories. The complex cladistic techniques used for such analyses, not detailed here for lack of space, were initially proposed by Vane-Wright et al. (1991), and progressively refined by Williams et al. (1991, 1994), Weitzman (1992) and Faith (1992, 1994) (review in Williams and Humphries 1996). Obviously, the absence of phylogenetic trees for many organism groups precludes the usage of cladistic models. In these cases the diversity of higher taxa, such as genera or families, is employed as an indirect estimate, or surrogate, of character diversity. Diversity of species The species is considered by most authors as the fundamental currency of biodiversity. On this line, the species has been defined as the base-unit of biodiversity, species diversity as the main expression of biodiversity and species extinction as the main manifestation of the crisis or loss of biodiversity. To date, most studies were thus conducted employing species diversity as the main estimate of biodiversity; in particular, in most cases the simple species richness has been used as an estimate of biodiversity. The usage of species richness instead of species diversity is caused more by practical, logistical and economical reasons than theoretical ones: in fact, it is usually practically impossible to census all individuals of all study species for studies at the regional level. Hereafter, we use species richness as a surrogate for species diversity. Methods to measure species richness are of two kinds: 1. sample-based methods: samples and estimates of species richness are collected within each area; 2. indirect methods, often defined as “surrogates”: these are estimates of species richness based on the measurement of one or more predictive variables. Indirect methods are usually based on one of the three following predictive variables: 1. environmental variables: the diversity of habitats or ecosystems is used to predict species richness; 2. indicator groups: the diversity or richness of certain indicator groups, often predators such as raptors or carnivores, is used as an indirect estimate of the overall species richness; 3. higher taxa: the richness of higher taxonomic groups, such as genera or families, is used to estimate species richness. 3 Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali Progetto Biodiversità: Integrare lo sviluppo del territorio con la conservazione della biodiversità in provincia di Trento. Testo approfondito Diversity of habitats or ecosystems Habitat or ecosystem diversity is sometimes recorded in the field, but nowadays is usually recorded through topographic or satellite maps detailing vegetation or land-uses. Diversity is measured as the presence and frequency occurrence of each habitat or ecosystem per unit space, estimated through diversity indexes. Such measures often include estimates of the topographic and geophysical diversity of the sample area, as these are often key factors regulating biodiversity. Functional diversity Ecosystem diversity has been recently advocated as one of the few measurable components of biodiversity incorporating an estimate of the so called “functional diversity”. More direct measures of functional diversity have been recently proposed. These consist of subdividing species on the basis of the trophic role of organisms within the community, such as herbivores, parasites, meso-predators, or top-predators. Functional diversity is then expressed as the number of trophic functions recorded in the sample. 2. PROJECT BIODIVERSITÁ In the past decades there has been a steady rise in the demand by public opinion for “transparent” political actions, based on criteria as objective as possible, aimed at maximising the action efficacy while minimising the economical expenditure of the tax payer. In addition, the steep rate of biodiversity loss at the global level has increased the pressure for sustainable development, based on environmentally-friendly policies. As a response, there is higher urgency for public administrations to base their actions on technical, quantitative knowledge, in line with the standard level of scientific knowledge of other European countries, and supplied by independent sources, such as universities and museums. Notwithstanding such trends, very few biodiversity studies have been carried out in Europe, and only a handful of them were conducted in Italy. Most studies were carried out at a very large spatial scale, using only species richness as a biodiversity estimate and ignoring all its other components (genetic, character, functional and ecosystem diversity); very rarely were such studies conducted at a local scale and with sufficient detail. Finally, in many cases such studies lacked the necessary scientific rigor which would have allowed them to be published on international journals; their results were thus confined to local reports, not easily accessible to the scientific community and the general public. This situation on one side reflects the recent birth of biodiversity as a formal science with a repertoire of standard study methods, and on the other is caused by the economic and practical difficulty of accomplishing such type of research. In agreement with the general national situation, the Trento region still lacks any biodiversity assessment based on advanced scientific criteria and in line with the standard methodologies set by the international scientific community. This project aims to fill these gaps, by means of a study aimed at sampling all the four main components of biodiversity. To this end, numerous complementary techniques of data collection and analysis will be employed, with the final objective to develop a GIS biodiversity map of the whole Trento region. Such map will allow to: • integrate plans of regional development with the conservation of natural resources; • simulate the loss or increase of biodiversity associated with potential environmental changes cause by local development policies; • outline the priority areas, sites or reserves for conservation action; 4 Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali Progetto Biodiversità: Integrare lo sviluppo del territorio con la conservazione della biodiversità in provincia di Trento. Testo approfondito • optimise the management of present reserves. In particular, aim of the project will be to: 1. record biodiversity estimates at various hierarchical scales of organization (individuals, species, genera, families, ecosystems); 2. incorporate indexes of vulnerability or conservation priority into such biodiversity estimates; 3. build statistical models aimed at predicting field-recorded biodiversity estimates from topographic, vegetational and land-use characteristics, so as to integrate biotic and abiotic components of ecosystems; 4. validate the predictive models by applying them to samples statistically independent from the one used to build the models; 5. apply the predictive models and field-recorded estimates to the whole Trento region, so as to create a regional biodiversity map; 6. highlight areas of highest biodiversity and/or conservation priority, the so called “biodiversity hotspots” (Myers 1988, 1990, Myers et al. 2000); 7. build statistical models easy to apply for the non-expert and capable to predict the loss or increase of biodiversity associated with environmental changes driven by local development (e.g. land-use changes, such as the currently ongoing conversion of Alpine meadows to scrubs and then forest); 8. propose databases to be integrated in the projects of environmental analysis and assessment conducted by the Autonomous Province of Trento (e.g. forestry studies, environmental impact assessment analysis, etc.). 3. METHODS AND EXPECTED RESULTS 3.1. Data collection on the field Project BIODIVERSITÀ aims to estimate biodiversity using vertebrates as model organisms. This choice is caused by the following reasons: (1) censusing the invertebrate component of biodiversity is rarely possible over large areas; (2) there is in-depth knowledge about the distribution of vertebrates (and scarce information about the distribution of invertebrates) within the Trento region; (3) vertebrates are often perceived in a more positive way by the public and are thus a better tool for environmental education (Jacobson and McDuff 1998). Data collection in the field will be organised on the basis of the three following aspects. Atlases and databases of vertebrate distribution Research carried out in last fifteen years has culminated in the creation of large databases on the distribution of all species within the vertebrate classes of Birds, Mammals, Reptiles and Amphibians in the Trento region. Data on fish distribution will be acquired by access to a database currently in press. Such databases will be completed within 2001 through data collection in the few areas which previously received little coverage. Such update will allow the publication of the distribution atlases, used as tools to record species richness within each quadrat of the 1: 25 000 IGM topographic maps (n = 81 quadrats). Intensive population studies Atlas work will allow usage of presence-absence data from sample quadrats; this approach yields important data for large areas but with little detail: for example, a species may be present in a quadrat 5 Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali Progetto Biodiversità: Integrare lo sviluppo del territorio con la conservazione della biodiversità in provincia di Trento. Testo approfondito but with only one non-breeding individual. There is thus high need for complementary estimates of environmental quality, based on the density and breeding success of species in the quadrat. (Van Horne 1983). As recording density and productivity estimates for all species in each quadrat is not feasible, such data will be obtained only for some sample species. These species and the areas where they will be studied were chosen through the following procedure: 1. we first identified the main macro-habitats within the Trento region. We preferred to chose samplespecies on the basis of habitats because: a. the main aim of the procedure was to find indicators of habitat quality; b. habitats are one of the fundamental units on which to base plans of conservation actions; c. habitat conservation is a more efficient way to protect overall biodiversity than the conservation of single species. 2. We chose one, or more than one top-predatory species within each macro-habitat. We preferred to give emphasis to top-predators because: a. being at the top of the food-pyramid, predators reflect perturbations at the lower levels of the pyramid and may thus be reliable indicators of ecosystem health; b. richness of predatory species is generally strongly and positively correlated to the richness of prey species in the same area; c. top-predators are often vulnerable, conservation-priority species, thus their conservation is already an integral part of a strategy of biodiversity preservation; d. top-predators exert a strong fascination on the general public and are thus optimal models for the popularisation of issues of biodiversity conservation. 3. For each predator, we selected one or more of its main prey species, whose abundance will be recorded in the same areas where predators will be studied. This will allow to sample biodiversity at various levels of the trophic pyramid within each macro-habitat. 4. For each system “macro-habitat – top-predator – prey-species”, we selected random study plots within the Trento region. All the study plots were selected so as to include a minimum area of 100 km2 or a minimum sample of 25-30 breeding pairs of the predatory species. This minimum sample will allow us to build statistical models predicting the distribution of predatory and prey species, and reapply the models to the whole Trento region. Hotspots of “transient” biodiversity There is a local component of biodiversity given by the flow of migrating birds passing through the Trento region every year. This component, which could be defined as “transient” biodiversity, is difficult to estimate through the above cited methods, and causes suddenly high biodiversity values within small areas and time frames. To quantify such component, we will carry out progressive visual and acoustic censuses, and sessions of mist-netting and ringing of migrant birds at various sites suspected to be key areas for passage and stop-over of migrants. This work, often conducted within nature reserves (“biotopi provinciali”), will also allow to explore the importance of current reserves as biodiversity hotspots. 3.2. Expected results and methods to accomplish them 1) Record biodiversity estimates at various hierarchical scales of organization. Genetic and character diversity will be analysed by means of cladistic, phylogenetic techniques applied to atlas data (review in Williams and Humphries 1996). Species diversity will be investigated by using data from distribution atlases and from intensive population studies, so as to estimate both species richness and 6 Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali Progetto Biodiversità: Integrare lo sviluppo del territorio con la conservazione della biodiversità in provincia di Trento. Testo approfondito evenness. Diversity of migrant species will be analysed separately, using data from the ringing stations and from the progressive censuses within local reserves (“biotopi”). Diversity of habitats and ecosystems will be quantified through a geographic Information System (GIS), accessing the available GIS databases: the forest cover map of Servizio Foreste (1999), and the “Corine” land-use map. Such data will be integrated with those from very recent aerial photographs. Finally, functional diversity will be estimated through the techniques reviewed by Martinez (1996), using data from both distribution atlases and intensive population studies. 2) Incorporate indexes of vulnerability into biodiversity estimates. Various vulnerability estimates will be integrated with the biodiversity ones. The main vulnerability indexes which will be used will be extracted from the IUCN classification, the Bern and Bonn Conventions, the “Habitats” Directive of the EU and the categories of conservation priority proposed by BirdLife International for birds. 3) Build statistical predictive models. For each quadrat of the 1: 25 000 IGM grid (n = 81 quadrats) we will record through a GIS variables related to: (1) topography (e.g. altitude, mean slope, ruggedness indexes), (2) vegetation, (3) rate of local development (e.g. length of paved roads), and (4) land-use. These will be used as independent predictor variables in statistical models aimed at predicting biodiversity values recorded in each sample quadrat. This type of analyses usually yield reliable models, capable to predict biodiversity values with reasonable accuracy. Employed models will include univariate and multivariate analyses, mainly multiple and logistic regression, discriminant analysis, and general linear models (GLM). The integration of environmental and geophysical variables with biodiversity estimates will allow the construction of more complete estimates, incorporating both the living (biotic) and non-living (abiotic) component of the ecosystem, and thus related to the overall “biological integrity” of an area. 4) Validate the predictive models. The above cited models will be built with sub-samples of the available data. Their predictive power will be tested by applying them to the rest of the originally available sample, thus applying them to a statistically independent sample, a standard procedure in this sort of studies. 5) Create a regional biodiversity map. Direct estimates and predictive models will be applied to the whole Trento region. Various biodiversity maps will be initially created, one for each main biodiversity component. An overall biodiversity map will be created by employing principal component analysis (PCA) to incorporate all biodiversity and vulnerability estimates in one only index. 6) Highlight biodiversity hotspots. We will highlight areas characterised by high biodiversity and simultaneous high abundance of vulnerable species. Such biodiversity hotspots will need more indepth studies on a finer spatial scale, following a commonly applied hierarchical procedure of study, which involves the identification of priorities first at coarse levels and then at progressively finer spatial scales. 7) Propose environmental simulation tools. Equations derived from the above models will be proposed as potential tools to simulate the biodiversity loss or increase caused by potential environmental changes in the future, such as those caused by incentive schemes of the EU Common Agricultural Policy. The models will allow to chose among available options the least harmful for biodiversity. 8) Propose tools to be integrated within the PAT management systems. Based on the results and simulations obtained, we will propose tools, such as models and databases, which can be integrated within the PAT projects of environmental study and/or analysis (e.g. forestry studies, environmental impact assessment analyses, etc.). 7