Middle-Sized Carnivores in Agricultural Landscapes [1 ed.] 9781611223682, 9781611220339

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Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved. Middle-Sized Carnivores in Agricultural Landscapes, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved. Middle-Sized Carnivores in Agricultural Landscapes, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

ANIMAL SCIENCE, ISSUES AND PROFESSIONS

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MIDDLE-SIZED CARNIVORES IN AGRICULTURAL LANDSCAPES

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ANIMAL SCIENCE, ISSUES AND PROFESSIONS

MIDDLE-SIZED CARNIVORES IN AGRICULTURAL LANDSCAPES

LUÍS M. ROSALINO AND

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CARLA GHELER-COSTA EDITORS

Nova Science Publishers, Inc. New York

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Copyright © 2011 by Nova Science Publishers, Inc. All rights reserved. No part of this book may be reproduced, stored in a retrieval system or transmitted in any form or by any means: electronic, electrostatic, magnetic, tape, mechanical photocopying, recording or otherwise without the written permission of the Publisher. For permission to use material from this book please contact us: Telephone 631-231-7269; Fax 631-231-8175 Web Site: http://www.novapublishers.com NOTICE TO THE READER The Publisher has taken reasonable care in the preparation of this book, but makes no expressed or implied warranty of any kind and assumes no responsibility for any errors or omissions. No liability is assumed for incidental or consequential damages in connection with or arising out of information contained in this book. The Publisher shall not be liable for any special, consequential, or exemplary damages resulting, in whole or in part, from the readers‘ use of, or reliance upon, this material. Any parts of this book based on government reports are so indicated and copyright is claimed for those parts to the extent applicable to compilations of such works.

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Independent verification should be sought for any data, advice or recommendations contained in this book. In addition, no responsibility is assumed by the publisher for any injury and/or damage to persons or property arising from any methods, products, instructions, ideas or otherwise contained in this publication. This publication is designed to provide accurate and authoritative information with regard to the subject matter covered herein. It is sold with the clear understanding that the Publisher is not engaged in rendering legal or any other professional services. If legal or any other expert assistance is required, the services of a competent person should be sought. FROM A DECLARATION OF PARTICIPANTS JOINTLY ADOPTED BY A COMMITTEE OF THE AMERICAN BAR ASSOCIATION AND A COMMITTEE OF PUBLISHERS. Additional color graphics may be available in the e-book version of this book.

LIBRARY OF CONGRESS CATALOGING-IN-PUBLICATION DATA Middle-sized carnivores in agricultural landscapes / editors: Lums M. Rosalino, Carla Gheler-Costa. p. cm. Includes index. ISBN H%RRN 1. Animal ecology--Mediterranean Region. 2. Animal ecology--Brazil. 3. Carnivora--Adaptation--Mediterranean Region. 4. Carnivora--Adaptation--Brazil. I. Rosalino, Lums M. II. Gheler-Costa, Carla. III. Title: Middle sized carnivores in agricultural landscapes. QH150.M53 2011 599.7'1755--dc22 2010036436

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CONTENTS

Preface

vii Luís Miguel Rosalino and Carla Gheler-Costa

Chapter 1

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Chapter 2

Adaptation of Mesocarnivores (Mammalia: Carnivora) to Agricultural Landscapes in Mediterranean Europe and Southeastern Brazil: A Trophic Perspective Luciano M. Verdade, Luis Miguel Rosalino, Carla Gheler-Costa, Nuno M. Pedroso and Maria Carolina Lyra-Jorge Factors Affecting Small and Middle-Sized Carnivore Occurrence and Abundance in Mediterranean Agricultural Landscapes: Case Studies in Southern Portugal Filipe Carvalho, Ana Galantinho and António Mira

Chapter 3

Fruits and Mesocarnivores in Mediterranean Europe Luís M. Rosalino and Margarida Santos-Reis

Chapter 4

Ecology of European Badgers (Meles meles) in Rural Areas of Western Switzerland Emmanuel Do Linh San, Nicola Ferrari, Claude Fischer and Jean-Marc Weber

Chapter 5

Chapter 6

Chapter 7

1

39 69

83

Middle-Sized Carnivores in Mosaic Landscapes: The Case of Biscay (SW Europe) Cristina Rodríguez-Refojos and Iñigo Zuberogoitia

105

Predicting the Spatial Distribution of Four Sympatric Species of Mid-Sized Carnivores in an Amazonian Deforestation Frontier Fernanda Michalski, Carlos A. Peres and Jean Paul Metzger

127

A First Approach to the Comparative Ecology of the Hoary Fox and the Crab-Eating Fox in a Fragmented Human Altered Landscape in the Cerrado Biome at Central Brazil Frederico G. Lemos, Kátia G. Facure and Fernanda C. de Azevedo

143

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vi Chapter 8

Contents Western Mediterranean Landscapes: Opportunities and Challenges for Carnivore Conservation Margarida Santos-Reis

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Index

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161 183

PREFACE Luís Miguel Rosalino1,2 and Carla Gheler-Costa3 1

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Centro de Biologia Ambiental, Universidade de Lisboa, Faculdade de Ciências de Lisboa, Edifício C2, Campo Grande 1749-016 Lisboa, Portugal 2 Laboratório de Ecologia, Turismo e Sustentabilidade (LETSISLA), ISLA-Lisboa - Instituto Superior de Línguas e Administração, Quinta do Bom Nome, Estrada da Correia, 53 - 1500-210 Lisboa, Portugal 3 Laboratório de Ecologia Animal, Departamento de Ciências Biológicas / ESALQ / Universidade de São Paulo, Caixa Postal 09, Piracicaba, SP 13418-900, BRASIL

Nowadays, habitat loss is one of the main threats to terrestrial vertebrates‘ survival. Due to the decreasing of continuous natural areas, and consequent habitat loss, how animals use these remaining patches, is becoming a central question for the conservation of these species. The loss of the original land covers can promote a change in the composition, diversity and behaviour of the native fauna, as well as constrain the community structures. The outcome of these processes of habitat fragmentation due to human agro-forestry management is a landscape where patches of autochthonous/native vegetation are immersed in a matrix of human shaped landscapes. Although some carnivores are sensitive to fragmentation, some species can benefit with the expansion of agriculture and/or silviculture, especially generalists mesocarnivores, which are able to take advantage of, for example, the food supply that some of these landscapes offer. However, it is not clear if this benefit is in a medium- or long-term, or just a numerical short-term response to an eventual local resources surplus. Furthermore, the importance and role of those agricultural matrix for species conservation is not consensual, although it is essential to incorporate them in conservation plans, especially in areas where they are widespread, shaping the countryside for thousands or hundreds of years. Motivated by this need to understand how mesocarnivores are adapted to agro and/or silvicultural landscapes, as well as to share experiences and knowledge on the impact that this human shaped environments have on several middle-sized carnivores groups, in August 2009 we organized a symposium in the 10th International Congress of Mammalogy (IMC-10), hold in Mendoza, Argentina, entitled "Middle-sized carnivores in agricultural landscapes". This

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viii

Luis M. Rosalino and Carla Gheler-Costa

symposium had the participation of researchers from Portugal, Brazil, South Africa, Switzerland and United Kingdom and was attended by a large number of student and researchers from different world regions interested in questions regarding mesocarnivores living in agriculture landscapes throughout the world. Due to the high interest showed by all the symposium participants in this subject, as well as the lack of a tool which compiled information on this topic, the natural consequence would be to publish a text book focused on mesocarnivores relation with agricultural landscapes. While most of the symposium speakers have agreed to be involved in this project, we felt the need to aggregate information regarding other geographical areas (e.g., Spain) or focused on different Human intervention schemes (e.g., open pasturelands in the Brazilian ‗arc of deforestation‘). Therefore, this book aims to contribute to the discussion regarding the relations between middle sized carnivores and agrosilvo-pastoral landscapes, by presenting case studies of how some mesocarnivores are using these altered landscapes, in Europe and South America, as well some reviews on the intrinsic values of these habitats for mesocarnivores conservation. In the first chapter is presented a review regarding the use of trophic resources by mesocarnivores in agricultural landscapes of Mediterranean Europe and Southeastern Brazil, focusing on the possible role of the surplus of food available in these environments (i.e., increase in the carrying capacity) on the increase in numbers of these carnivores in opposition to the mesopredator release hypothesis. Some case studies from southern Portugal focused factors affecting small carnivore occurrence and abundance in Mediterranean agricultural landscapes are described in Chapter 2. Overall, the study show that the maintenance of a sustainable mosaic embracing Montado (cork oak woodland), shrubland and open land areas, may allow higher carnivore species richness and abundance by enhancing connectivity between crucial areas. It‘s concluded that the implementation of this kind of agricultural practices considering the landowners needs (including cattle, sheep, goat and pig raising and game) is the key issue to achieve the main carnivore conservation goals in Southern Portugal. In Chapter 3 is presented a review on fruit consumption by mammalian carnivores in the Mediterranean region and the role of these vertebrates as seed dispersers. Is shown that generalist predators, such as red fox, stone marten, Eurasian badger and common genet consumed more diversified fruits, including those produced in orchards, olive groves and vineyards, with a tendency to increase the consumption towards eastern Mediterranean areas. Moreover, it was confirmed the seed disperser character of many of mesopredator species (e.g., genets), including those fruit species produced by man (e.g., grapes), and some considerations are presented concerning the role of fruits in mesocarnivores conservation and the possible impact of the dispersion of non-native seeds. Another review is presented in Chapter 4. Here data on the ecology of Eurasian badgers in four different agricultural areas of western Switzerland (located at different altitudes and characterized by varying climatic conditions and human intensity of land use) were analyzed. It‘s demonstrated that badgers cope well with human origin resources, since maize and other cereals (e.g., wheat) are indeed the dominant food items. Moreover, badgers breeding setts were consistently located in wooded areas (often restricted to patches in agricultural landscapes), indicating that the availability of favorable sett sites is probably one of the most important factors to consider when dealing with conservation of central European badger populations inhabiting agricultural landscapes.

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Preface

ix

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.

In Chapter 5 an analysis of the trends of the carnivore guild inhabiting the Biscay Province in northwestern Spain (southwestern Europe) is presented. This area has been subject to a huge landscape change induced by man, where most of the ―pristine‖ habitats have been destroyed and nowadays production forests have become the dominant landscape unit. In this scenario, where the other land covers (including agriculture patches) are highly fragmented, carnivores have responded differently. While generalist species (e.g., red fox) seem to be increasing their range and numbers, specialists (e.g., wildcats) are apparently becoming scarcer, and could face regional extinction if the landscape continues to change in the same direction (intensification of production forests). However, while some invasive species (e.g., American mink) seem to be occupying the vacant niches left by some native specialists, these man shaped landscape are being recolonized by one of the more emblematic mustelids: the Eurasian otter. Returning to South America, in Chapter 6 is presented another case study. This time authors used distributional temporal data series (obtaining by camera trapping, direct and indirect sighting records) on four sympatric mesocarnivores species and environmental factors related to their spatial distribution in a highly fragmented landscape of northern Mato Grosso, southern Brazilian Amazonia. It‘s showed that land-cover, distance to water sources and distance from urban centres were highly influential on carnivores‘ occurrence. The current levels of deforestation in that area may result in the expansion of the distribution of species less susceptible to Human presence and habitat changes (e.g., crab-eating fox), and the reduction of the range of forest-dependent species such as the ocelot and the margay. The study concludes that mesocarnivore conservation in this southern America region is dependent upon greater efforts in environmental law enforcement and education programs to maintain the remaining forest cover. In Chapter 7 are described several aspects of the hoary fox (Lycalopex vetulus) and crabeating fox (Cerdocyon thous) ecology in cattle farms in Goiás state, Center-West Brazil. The ecological adaptations of this mesocanids to a fragmented landscape in the Cerrado was accessed by studying these carnivores food habits, habitat use, foraging group size and parental care, as well as the interactions between both species, domestic mammals (domestic dogs and cattle) and man. Foxes revealed different hunting strategies, with crab-eating foxes foraging more frequently in pairs, while hoary foxes were primarily solitary hunter. Moreover, these differences also include diet diversity and habitat use, since hoary foxes were located more in grazed pasture and seem to be more focused on termites‘ consumption, while crab-eating foxes use more diversified resources (habitats and food). Both species seem to be negatively affected by man induce perturbations (mainly dogs, roads, and illegal poisoning), but the conversion of natural areas of Cerrado into agro-pastoral areas seems to affect less hoary foxes. Finally in Chapter 8 is discussed the idea that to achieve success conservation of middlesized carnivores in man shaped environments is crucial to consider not only the understanding of the ecological processes but also its spatial and social contexts. This reasoning is exemplified by analyzing the mesocarnivore guild inhabiting spatial and temporal heterogeneous landscapes of south western Europe. Namely, is discussed the opportunities created by the current multi-use landscapes of southern Portugal, the adaptation of individuals to the development trend of modern human societies, and the impacts of the Human intervention schemes on the carnivore community. Moreover, carnivores‘ conservation options in a highly interventioned area, such as southern Iberian landscapes, are presented.

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We sincerely hope that the ideas and examples presented in this book, together with the discussion about the role of agricultural landscapes on middle-size carnivore conservation and population adaptation to the characteristics of man-shaped environments, can be a starting point to a broader discussion regarding: the effective function of these environments for mesopredators; and the management schemes that might maximize carnivore biodiversity and conservation, without compromising farms‘ profits.

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In: Middle-Sized Carnivores in Agricultural Landscapes ISBN: 978-1-61122-033-9 Editors L. M. Rosalino and C. Gheler-Costa, pp. 1-38 © 2011 Nova Science Publishers, Inc.

Chapter 1

ADAPTATION OF MESOCARNIVORES (MAMMALIA: CARNIVORA) TO AGRICULTURAL LANDSCAPES IN MEDITERRANEAN EUROPE AND SOUTHEASTERN BRAZIL: A TROPHIC PERSPECTIVE Luciano M. Verdade¹, Luis Miguel Rosalino², Carla Gheler-Costa¹, Nuno M. Pedroso² and Maria Carolina Lyra-Jorge1

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¹ Laboratório de Ecologia Isotópica, CENA / Universidade de São Paulo, Caixa Postal 96, Piracicaba, SP 13400-970, BRASIL ² Centro de Biologia Ambiental, Faculdade de Ciências da Universidade de Lisboa, Ed. C2, Campo Grande, 1749-016 Lisboa, PORTUGAL

ABSTRACT The conversion of natural ecosystems into silvo/agriculture systems has been occurring for millennia or centuries throughout our planet. While for many carnivores species this landscape transformation can have negative conservation consequences, for others it could represent a window of opportunity. Therefore, we aimed to review the use of agricultural ecosystems of Mediterranean Europe and Southeastern Brazil by carnivores, as well as how these predators use the surplus of food available in those menshaped environments. Our review shows that most of these studies were carried out in mono-silvicultural landscapes of Southeastern Brazil and in mixed-agricultural landscapes of Southern Europe. Moreover, while Neotropical species prey more upon small vertebrates, Mediterranean species tend to consume more often invertebrates and fruits. We discuss the possible role of this increase in habitat carrying capacity on the enhancement in numbers of these carnivores in opposition to the mesopredator release hypothesis. We believe that mesocarnivores are adapting to take advantage of a trophic resource enhancement opportunity window created by agro-systems practices, which increase the overall landscape carrying capacity as agriculture is re-shaping the landscape for thousands of years. Finally, we discuss the possible conservation value of agricultural landscapes

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1. INTRODUCTION Large-sized carnivores experienced massive population declines due to habitat destruction and human hunting pressure (Crooks, 2002). The former is predominantly associated with agriculture expansion (Marino, 2003), whereas the second is mostly related with livestock depredation (Blanco and Cortés, 2007) and illegal trade of skin or other body parts (e.g., Shepherd and Nijman, 2008). Large home ranges, naturally low population densities and a low intrinsic growth rate make large carnivores more prone to these anthropogenic impacts (Crooks, 2002; Caughey, 1977, 1994). This is the case of large felids such as the tiger (Seidensticker et al., 1999), the African and the Asian lion (Chardonnet, 2002) and the jaguar (Hoogesteijn and Mondolfi, 1992), as well as other large carnivores like the brown bear (e.g., Palomero et al., 1997) and the wolf (Mech, 1970). There are evidences that the extinction of top predators can have an indirect and negative effect on prey species, by increasing the presence of other smaller predators (Wright et al., 1994; Palomares et al., 1995; Palomares et al., 1996; Crooks and Soulé, 1999; Terborgh, 2000; Gehrt and Clark, 2003), which were no longer subjected to a predatory/competition constrain by larger carnivores. However, this interpretation has been subjected to some criticisms (e.g., Litvaitis and Villafuerte, 1996), and other alternative hypotheses have also been proposed. Some authors have argued that more generalist predators are favoured when natural landscapes are fragmented by agriculture, even in the secular absence of larger predators due to an increase in food resources which they can use (Oehler, 1995; MoránLópez et al., 2006; McDougall et al., 2006; Sánchez-Hernandéz et al., 2001; Tabeni and Ojeda, 2005). Agricultural landscapes are mostly mosaics formed by a matrix of an agroecosystem (the landscape element with a high connectivity whose area exceeds the area of all other elements combined – Forman, 1995) permeated by remnant patches of native vegetation. Agroecosystems are cultivated areas of domestic plant species for economic purposes. The permanence of wild species in agricultural landscapes depends basically on the matrix permeability and on the resources available, both in the agroecosystems and in the remnant patches of native vegetation (Gascón et al., 1999). In general, the displacement of native ecosystems by agriculture affects species composition and consequently the ecosystem structure and functioning (Oehler and Litvaitis, 1996). However, some species – in especial mesocarnivores (intermediate body-size mammalian carnivores – Buskirk and Zielinski, 2003) – apparently benefit from this land use change (e.g., Laurance, 1994; Litvaitis and Villafuerte, 1996; Chiarello, 1999; Gehring and Swihart, 2003; Dotta and Verdade, 2007) due to a possible increase in their food resources (Moguel and Toledo, 1999; Faria et al., 2006; Acharya, 2006). Agriculture has been developed for the last thousands of years (Diamond, 1997, 2005), affecting not only the spatial structure of the landscape but also inducing temporal heterogeneity (Wiens, 2000). Time scale should then be considered in order to evaluate the effects of agriculture on the patterns of abundance and distribution of biodiversity (Balée, 2006). In this study we review the use of food resources by mesopredators in agricultural landscapes from Mediterranean Europe and Southeastern Brazil, as a way to address adaptive processes of mesopredators to Human changes in the landscapes. Agriculture based on deforestation has been implemented in both regions but in different time scales. In southern

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Adaptation of Mesocarnivores…

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Europe this landscape shaping activities induced by man started some millenniums ago. For example, in the island of Crete (Greece - Eastern Mediterranean) 4000-3000 BC the landscapes nearby the city of Kommos already consisted of a intricate mosaic of cultivated fields and orchards interspersed with semi-natural woodlands (Blondel and Aronson, 1999). In Southeastern Brazil, agriculture transformation started only recently (in the last 200 to 300 years – Dean, 1995). Although such a different time scale could act as a diversion variable, this agricultural transformation of the landscape in both geographical regions have similarities: the type of agriculture, together with its management schemes, implemented in Southern America have a southern European origin, since they were introduced in the New World by European colonial settlers and immigrants. Moreover, ecologically similar species of mesopredators are found in both (Jaksić and Delibes, 1987). Although sharing a common set of evolutionary features the families herein considered vary considerably in their life history strategies. Felids are highly efficient solitary predators hunting mostly on vertebrate prey by ambush. On the other hand, herpestids and canids developed well structured social systems. Ursids, procyonids and viverrids developed threeclimb abilities. However, the former are large-sized omnivores, the intermediate are middlesized omnivores and the later are middle-sized carnivores. At last, mustelids have a slender body shape with a relatively high metabolic rate and a great variation in body size and food habits (García-Perea et al., 1996; Macdonald, 2001; Nowak, 2005).

2. STUDY AREAS

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2.1. Mediterranean Region The Mediterranean region is defined by the Mediterranean Sea basin. It‘s delimited by the Euro-Siberian region to the north (with the Palaeartic deciduous and coniferous forests interspersed with steppes), the Saharo-Arabian in the south (influenced by the Sahara desert) and the Irano-Turanian region on the east, which includes high steppes (Blondel and Aronson, 1999). It is composed by diverse ecosystems, adapted to the transitional regime between cold temperate and dry tropical climates that, in a geological scale, can be considered young due to the relatively recent appearance of a Mediterranean climate (Suc, 1984). This climate, characterized by hot, dry summers and cool humid winters, has one defining trait: unpredictability. The Mediterranean region‘s current biodiversity also comprises species whose core distribution is located in the other biogeographical regions, which led to its inclusion in the list of the world‘s 25 biodiversity hotspots for conservation priorities (Myers et al., 2000). This rich species diversity and high number of endemisms are a result of the conjunction action of three factors: biogeography, geology and history (Blondel and Aronson, 1999). Although the first two shaped biodiversity and the species natural history, the long lasting Human history in the region seems to be the most important constraining/facilitating factor. Its continuum contact with the vast land of Eurasia and Africa promoted species contact and interaction with a strong influence on species evolution. In addition, its climatic unpredictability created opportunities for specializations (Blondel and Aronson, 1999). However, the Human factor is a constant since the end of the late Pleistocene, with the region

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being the birth place of the largest and most powerful civilizations of the Old World (e.g., Sumerians, Phoenicians, Greeks, Romans, Ottomansand Sarrasins) and the focus of northern invaders (e.g. Vandals and Visigoths), who have impacted ecosystems everywhere in the basin leading some authors to consider that a ‗coevolution‘ has shaped the interactions between Mediterranean ecosystems and humans through constantly evolving land use practices (di Castri, 1981 in Blondel, 2006). Naveh and Dan (1993 in Blondel and Aronson, 1999) suggested that humans have a direct impact in the Mediterranean landscape for at least 50.000 years leading to an intensive landscape redesign. However, the most intense ecosystem transformation occurred 10.000 years ago, when hunters/collectors began the domestication of animals and cultivation of plants to survive. Since then, pastoralists and agriculturalists were responsible for major deforestation and soil erosion (Blondel and Aronson, 1999). Nowadays, agro-silvo-pastoral Mediterranean landscapes include orchards, fruit farms, olive yards (intensive and extensive), cereal fields, pastures for cattle breeding, or production forests (e.g., Eucalyptus and Pinus), among others. Nevertheless remnants of deciduous (e.g., Quercus spp.), conifers (e.g., Pinus spp.) and riverine (e.g., Populus spp.) forests, matorral (e.g., Erica spp., Cistus spp., Arbutus spp.), tomillares and Phyrgana patches (e.g., Thymus spp., Genista spp.), steppes, grasslands and wetlands can still be found throughout the Mediterranean basin (Blondel and Aronson, 1999). This long lasting landscape shaping has produced intense modifications in the ecosystems with the transformation of native communities, including the extinction or declining of several species (e.g. Brown bear Ursus arctos and Iberian-lynx, Lynx pardinus populations in Portugal, respectively – Cabral et al., 2005), the invasion of some (e.g., Egyptian mongoose Herpestes ichneumon – Delibes, 1982) and the adaptation of others (e.g., Eurasian badger, Meles meles - Rosalino et al. 2005). Many species with high conservation profiles are associated with traditional land uses and with the human-maintained semi-natural habitats, what can be considered extreme cases of adaptation (Sirami et al., 2008). For example, great bustard Otis tarda, a globally threatened species (Alonso et al., 2003), depends greatly on agricultural areas for survival in the Iberian Peninsula (Lane et al, 2001). Other examples of the importance of man-shaped landscapes are the oak forests (cork and holm), typical Mediterranean forests and the major remaining wood-pasture systems of Europe. They represent a sustainable agro-silvo-pastoral system well adapted to the environmental restrictions of the Mediterranean region (low edaphic and climatic potential) (Pinto-Correia, 2000), important to wildlife conservation (Diáz et al., 1997; Cabral et al., 2005), including endangered species such as the polecat (Mustela putorius) (Santos-Reis et al., 1999) and the wildcat (Felis silvestris) (Virgós et al., 2002).

2.2. Southeastern Brazil The Southeast is the most populated and developed region of Brazil. As a consequence, this region presents the most dramatic environmental problems of this country including a massive deforestation of its major original biomes, the Atlantic Forest and the Cerrado. The former originally covered 12% of the country from 3º S to 31º S, and from 35º W to 60º W, mainly extending along the Brazilian coast (92%), but also reaching into Paraguay (Cartes and Yanosky, 2003; Huang et al., 2007) and Argentina (Giraudo, 2003).

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Possibly due to its large latitudinal range, the Atlantic Forest has a high number of endemic species even higher than some parts of the Amazon Forest (Silva and Leitão Filho, 1982; Peixoto and Gentry, 1990; Barros et al., 1991; Joly et al., 1991; Brown Jr. and Brown, 1992). For this reason, the Atlantic Forest is considered prioritary for conservation among tropical forests (Conservation International, 2008a). According to Joly et al. (1991), the Atlantic Forest is composed by three distinct formations: coastal lowland, steep and plateau forests. In Southeastern Brazil the steep forest formation is predominant. The Cerrado lato sensu varies from savanna to forest formation due to the physicalchemical characteristics of the soil and the frequency of natural fire (Coutinho, 2002; Ruggiero, 2002). Possibly due to its continental location in South America, the Cerrado has a high diversity of fauna and flora including species from the Amazon and the Atlantic forest as well as from the semi-arid Caatinga at its northeast and the Pantanal at its southwest. However, it has a relatively smaller number of endemic species (Durigan et al., 2007). Both biomes are characterized the typical climate of Southeastern Brazil which varies from humid subtropical (> 1,200 mm, 20ºC in Southern São Paulo) to semi-arid tropical (< 900 mm, 24ºC in Northern Minas Gerais) (Projeto Cactáceas do Brasil, 2010). The Atlantic Forest tends to cover the more humid areas closer to the seashore, whereas the Cerrado tends to cover the drier continental plateau (Huber, 1987). Although native South Americans used to make slash-and-burn agriculture in areas of the Atlantic Forest, only after the arrival of the first Europeans in the 1,500‘s deforestation became dramatic (Dean, 1995). By its turn, the Cerrado used to be predominantly inhabited by native hunters and gatherers and suffered a massive deforestation only in the late 20th Century due to the expansion of agroindustry in Brazil which also affected the remaining parts of the Atlantic Forest (Dean, 1995). As a result, there is currently only 12% of the Atlantic Forest (Ribeiro et al., 2009) and 20% of the Cerrado remaining in Brazil (Conservation International, 2008b).

3. METHODS In this study we reviewed the diet of species of the order Carnivora from the Mediterranean Europe (six families, 17 species) and the Southeastern Brazil (four families, 21 species), with special reference to mesocarnivores. Reviewed studies were divided into five silvo-agricultural landscapes types: intensive and extensive exotic pasture, mono- and mixedagriculture and mono-silviculture. We considered mono-cultures those where an agricultural or silvicultural matrix covers at least 75% of the landscape. Mixed-cultures were defined as those below landscape composition percentage (Agnelli and de Marinis, 1995).We calculated the percentage of diet studies implemented in each of the landscapes units considered in our review. In order to gather prey items presence/absence data on carnivores diet in a systematic approach, we clustered data into 12 food items categories: fruits, other plant materials, microinvertebrates, macroinvertebrates, fish, amphibians, reptiles, passerine- and nonpasserine birds, small, middle and large-sized mammals. Again, we calculated the percentage of diet studies mentioning the consumption of these 12 prey items by carnivores in the silvoagriculture landscapes of Southwestern Brazil (A) and Mediterranean Europe.

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Luciano M. Verdade, Luis Miguel Rosalino, Carla Gheler-Costa et al.

4. RESULTS We found 151 articles containing information of diet of Carnivora species in agricultural landscapes of the Mediterranean Europe and the Southeastern Brazil in South America, published between 1974 and 2007 (Table 1). Most of these studies were carried out in monosilvicultural landscapes of Southeastern Brazil and in mixed-agricultural landscapes of Southern Europe (Figure 1). However it‘s interesting to confirm that all of the studied species use, at least, one type of agro/forest system, which can be an indication that this species are managing to subsist (with more or less conflicts) in man shaped ecosystems. However, these studies do not cover all, or at least most, of the agricultural landscapes in both regions, which could bias our analysis. For example, while there is some information on the use of extensive pastures by carnivores in Brazil, especially regarding livestock depredation by jaguars and pumas, information about areas of intensive agriculture and livestock production is scarce (Table 1).

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A

B Figure 1. Percentage of carnivore diet studies implemented in different landscapes from Southwestern Brazil (A) and Mediterranean Europe (B) [MAg: Agricultural monocultures (> 75% of the landscape matrix); MSi (Silvicultural monocultures (> 75% of the landscape matrix); Mal (Mixed agricultural landscapes (< 75% of the landscape matrix); Pin: Intensive exotic pastures (> 75% of the landscape matrix); Pex: Extensive exotic pastures (> 75% of the landscape matrix)].

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Adaptation of Mesocarnivores…

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The Mediterranean carnivore guild includes 17 species, only three (18%) of which are considered large carnivores (European lynx Lynx lynx, wolf Canis lupus and brown bear Ursus arctos). In South eastern Brazil we identified 21 species, and again only four (19%) can be included in the large carnivores group, using the same patterns as for European communities (jaguar Panthera onca, Puma Puma concolor, maned wolf Chrysocyon brachyurus and giant otter Pteronura brasiliensis). Small vertebrates predominate in the diet of Neotropical carnivore species, including large felids (Figure 2). On the other hand, the Mediterranean species tend to consume more often invertebrates and fruits, which complement their consumption of vertebrate preys, resulting in a tendency to have a more diverse diet. Also relevant is the relatively high consumption of birds by middle-sized carnivores in both regions (Figure 2).

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A

B Figure 2. Percentage of diet studies mentioning the consumption of a particular prey item by carnivores in Southwestern Brazil (A) and Mediterranean Europe (B) [Fru: Fruits; Opm: Other plant materials; Ima: Macro invertebrates (> 10g), Imi: Micro invertebrates (< 10g); Fis: Fish; Amp: Amphibians; Rep: Reptiles; Apa: Passerine birds; Anp: Non-passerine birds; Msm: Small-sized mammals (< 500g); Mme: Medium-sized mammals (500 - 5,000g); Mla: Large sized mammals (> 5,000g)].

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Table 1. Diet and habitat use by Carnivora (Mammalia) in Mediterranean Europe and Southern South America Taxon REGION / Taxon

Conserv. status IUCN

Diet Fru

Environments Opm

Ima

Imi

Fis

Amp

Rep

Apa

1 2 3

1

2

1 6

18 19 23

6 20

Anp

Msm

Mme

Mla

MAg

MSi

Mal

Pin

Pex

2

1 2 3

2 3

2 6

1 2 6 7

8 9

6 8 9 10 11 12 13 14 15

16 17

1 6 8 18 20 21 23 24

8 21 23

9

6 9 10 11 12 13 14 25 10 11 10 11 14

22

SOUTHERN BRAZIL

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Felidae Panthera onca

NT

Puma concolor

NT

Leopardus pardalis

LC

Puma yagouaroundi

LC

Leopardus tigrinus

NT

Leopadus weidii

LC

Canidae

2

2

26

26

26

2

23

23

21

23

1

3 4 5

16 16

26

16

Taxon

Conserv. status

Diet

Environments

REGION / Taxon

IUCN

Fru

Opm

Chrysocyon brachyurus

NT

27

27

Cerdocyon thous

LC

28

Lycalopex vetulus

DD

32

Ima

Imi

Fis

Amp

27

28

28 29

32

28

28

Rep

Apa

6 27

Anp

Msm

Mme

Mla

6 27

6 27

27

9

28

28

28

32

32

32

9

MAg

30 31

MSi

Mal

6 9 10 11 13 14 15

27

9 10 11 12 13 14 15 29

28

14

32

Pin

Pex

16

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Mustelidae Eira barbara

LC

Conepatus semistriatus

LC

Galioctis vittata

LC

31

9

10 11 12 13 14

16

9 10 11 14 15 31

16

Table 1. (continued) Taxon

Conserv. status

Diet

REGION / Taxon

IUCN

Fru

Galictis cuja

LC

Lontra longicaudis

DD

Ptenoura brasiliensis

EN

Environments Opm

Ima

Imi

Fis

Amp

Rep

Apa

Anp

Msm

Mme

Mla

MAg

MSi

Mal

9

9 10 11 12 14 9 10 11 13

33

27

33

33

33

33

33

33

33

9

27

27

27

27

27

27

27

9

31

9 10 11 12 13 14 15

9

30 31

9 10 11 12 13 14

Pin

Pex

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Procionidae Procyon cancrivorus

LC

Nasua nasua

LC

MEDITERRANEAN EUROPE Felidae

27

16

16

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Taxon

Conserv. status

Diet

REGION / Taxon

IUCN

Fru

Lynx pardinus

CR C2a(i)

Lynx lynx

NT

Felis silvestris

LC

51 52

LR/cd

96 112

Environments Opm

Ima

Imi

Fis

Amp

Rep

Apa

135

Anp

Msm

Mme

Mla

125 130 131 135

130 131 135

124 125 128 129 130 131 135 21

125 130 131 135

129 133 134

23 63A

19 20 23 24 178 63A

178

114 51 52 70

63A

75

116 51 52 70 75 82 106

115 116 117 51 52 70 75 82 96 106

70 82

115 116 117 51 52 70 75 82 96 106

115 116 117 51 52 70 75 82 96 106

115 116 117 51 52 70 75 82 106

82

96

49 50 68 81 99

50 68 99

49 50 73 68 81 99 112

MAg

MSi

Mal

Canidae Canis lupus

68

41

50 81

Pin

Pex

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Table 1. (continued) Taxon

Conserv. status

Diet

Environments

REGION / Taxon

IUCN

Fru

Opm

Ima

Imi

Fis

Vulpes vulpes

LC

43 45 47 48 51 52 54 58 64 65 83 87 90 94 96 100 105 107 124

43 45 54 64 83 87 90 94 100 107

65 89

43 45 47 48 51 52 54 58 64 65 83 87 89 90 94 96 100 105 107 124

90

Mustela nivalis

LR/lc

67

Mustela erminea

LR/lc

79 158

Amp

Rep

Apa

Anp

Msm

Mme

Mla

MAg

45 51 52 65 83 90 96 100 107

47 48 58 64 65 83 87 90 94 96 105

45 48 51 52 54 58 64 65 83 87 90 94 96 100 105 107

43 45 47 48 51 52 54 58 64 65 83 90 94 96 100 105 107

43 45 47 48 51 52 58 64 65 83 87 90 94 100 105 124

45 48 58 64 65 83 90 105 107

58

67 96

67

67 96

67 96

67

79

79 44 41 42 43 45

160

MSi

Mal 45 51 52 94 105 107 113

Mustelidade 96

162

79 162

67

67

79

53 160

Pin

Pex

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Taxon

Conserv. status

Diet

Environments

REGION / Taxon

IUCN

Fru

Opm

Mustela putorius

LR/lc

93 179

93 179

Mustela vison

LR/lc*

143

Mustela lutreola Martes foina

EN A1ace LR/lc

Martes martes

LR/lc

Ima

Imi

Fis

Amp

Rep

Apa

Anp

Msm

Mme

93 144 179

93 144 179 184

93 144 179 184

93 144 151 179 184

93 144 151 179 184

121 126 127 139 143 183 184

121 126 127 139 143 184

93 144 145 151 179 184 121 126 139 143 144 184

121 126 127 139 183

121 126 127 139 143 144 183 184

121 127 139 143 144 183 184

143 144 184

35 45 78 83 107

35 40 45 69 78 83 104 105 107 108

69 104

35

35 40 45 69 83 104 107 108

35 40 69 83 104 107

121 126 127 139 143 144 183 184 120 35 40 45 69 78 83 104 105 107 108

35 40 45 69 78 83 104 105 108

35 45 69 83 107

39 55

34 39 55 57 96

34 39 55 57 96

55

55

144 184

143 144

118 35 40 45 69 78 83 88 104 105 107 108 34 39 55 57 96

55 96

121 35 40 45 69 78 83 104 105 107 108 34 39 55 96

Mla

MAg

MSi

Mal

150 182

93 148 149 152 179 119 119

182

119 35 40 69 80 98 104 105 107

55

Pin

Pex

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Table 1. (Continued) Taxon

Conserv. Status

Diet

REGION / Taxon Lutra lutra

IUCN NT

Fru

Environments Opm

Ima 153 154 155 157 165 166 168 171 172 173 174 175 176 177

Imi 153 154 155 157 161 163 165 166 168 169 171 172 173 174 175 176 177

Fis 96 15 3 15 4 15 5 15 7 16 1 16 3 16 5 16 6 16 8 16 9 17 1 17 2 17 3 17 4 17 5 17 6 17 7

Amp 153 154 155 157 161 163 165 166 168 169 171 172 173 174 175 176 177

Rep 96 153 154 155 157 161 163 165 166 168 169 171 172 173 174 175 176 177

Apa 155 157 161 165 166 171 172 173 174 175 177

Anp 155 157 161 165 166 168 171 172 173 174 175 177

Msm 96 155 157 161 163 165 166 168 173 174 175 176 177

Mme

Mla

MAg

MSi

Mal 164

Pin

Pex

Taxon

Conserv. status

Diet

REGION / Taxon Meles meles

IUCN LR/lc

Fru 36 38 42 47 54 59 66 71 74 76 77 91 95 96 97 101 124

Opm 36 54 59 74 76 77 97

LR/lc

72

72

LR/lc

37 46 51 52 55 56 60 62 96 102 103 105 109 110

37 55 60 85 86 102

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Ursidade Ursus arctos Viverridae Genetta genetta

Environments Ima

Imi 36 38 42 47 54 59 66 71 74 76 77 91 95 96 97 101 124

Fis 36

Amp 36 42 47 66 77 91 95 97 101

Rep 36 42 54 59 66 71 77 95 96 97 101

Apa 36 47 59 71 77 91 101

Anp 36 38 42 59 66 71 74 76 91 95 96 97 101

Msm 36 38 42 47 59 66 71 74 76 77 91 95 96 101

Mme 36 38 59 66 71 76 77 91 95 97 101 124

72 85 86 102 103

37 46 51 52 55 56 60 62 70 85 86 96 102 103 105 109 110 111

46 56 10 3

46 56 60 85 86 103 110 111

37 46 51 52 55 56 60 62 70 85 86 96 102 103 109 110 111

37 46 56 60 70 102

37 46 51 52 55 56 60 62 70 85 86 96 102 103 105 109 110 111

37 46 51 52 55 56 60 62 70 85 86 96 102 103 105 109 110 111

37 46 51 52 55 56 60 70 86 105 102 103 111

Mla 36 74 91

MAg 36

MSi

Mal 38 42 47 71 76 101

72

92

46 55 56 102

46 51 52 55 70 102 103 105

Pin

Pex

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Table 1. (Continued) Taxon

Conserv. status

Diet

Environments

REGION / Taxon

IUCN

Fru

Opm

Ima

Imi

Fis

Amp

Rep

Apa

Anp

Msm

Mme

Mla

Herpestidae Herpestes ichneumon

LR/lc

105

61 63 86

84 86

61 63 85 86 105

63

63 84 85 86

61 63 84 85 86 105

61 84 86 105

61 63 85 105

61 63 84 85 86 105

61 63 84 85 86 105

84

MAg

MSi

Mal

Pin

Pex

105

Fru: Fruits; Opm: Other plant materials; Ima: Macro invertebrates (> 10g), Imi: Micro invertebrates (< 10g); Fis: Fish; Amp: Amphibians; Rep: Reptiles; Apa: Passerine birds; Anp: Non-passerine birds; Msm: Small-sized mammals (< 500g); Mme: Medium-sized mammals (500 - 5,000g); Mla: Large sized mammals (> 5,000g); MAg: Agricultural monocultures (> 75% of the landscape matrix); MSi (Silvicultural monocultures (> 75% of the landscape matrix); Mal (Mixed agricultural landscapes (< 75% of the landscape matrix); Pin: Intensive exotic pastures (> 75% of the landscape matrix); Pex: Extensive exotic pastures (> 75% of the landscape matrix). 1 Taber et al., 1997; 2 Leite and Galvão, 2002; 3 Crawshaw, 2007; 4 Conforti and Azevedo, 2003; 5 Soisalo and Cavalcanti, 2006; 6 Mantovani, 2001; 7 Hass and Valenzuela, 2002; 8 Mazzolli, 2000; 9 Dotta, 2005; 10 Gargaglioni et al., 1998; 11 Talamoni et al., 2000; 12 Silva, 2002; 13 Tozetti, 2002; 14 Lyra-Jorge et al., 2008; 15 Hulle, 2006; 16 Chiarello, 1999; 17 Mazzolli et al., 2002; 18Sunquist et al., 1989; 19 Murray and Gardner, 1997; 20 Chinchila, 1997; 21 Miranda et al., 2005; 22 Trolle and Kéry, 2003; 23 Wang, 2002; 24 Meza et al., 2002; 25 Lopes and Mantovani, 2005; 26 Azevedo, 1996; 27 Motta-Júnior et al., 1996; 28 Gatti et al., 2006; 29 Beisiegel, 1999;3 0 Gheler-Costa et al., 2002; 31 Briani et al., 2001; 32 Dalponte, 1997; 33 Pardini, 1998;34 Agnelli and de Marinis, 1995; 35 Alegre et al., 1991; 36 Balestrieri et al., 2004; 37 Ballesteros et al., 2000; 38 Barea-Azcón et al., 2001; 39 Bermejo and Guita, 1996; 40 Bertolino and Dore, 1995 ; 41 Blanco and Cortés 2007; 42 Boesi and Biancardi, 2002 ; 43 Boldreghini and Pandolfi, 1991; 44 Bounous et al., 1995 ; 45 Brangi, 1995 ; 46 Calviño et al., 1984; 47 Canova and Rosa, 1993; 48 Cantini, 1991 ; 49 Capitani et al., 2004 ; 50 Carreira and Petrucci-Fonseca, 2000; 51 Carvalho and Gomes, 2001; 52 Carvalho and Gomes, 2004 ; 53 Ceña and Ceña, 2000; 54 Ciampalini and Lovari, 1985; 55 Clevenger, 1995 ; 56 Cugnasse and Riols, 1984; 57 de Marinis and Massetti, 1995; 58 Debernardi et al., 1991 ; 59 del Bove and Isotti, 2001 ; 60 Delibes, 1974; 61 Delibes, 1976; 62 Delibes, 1977; 63 Delibes et al., 1984; 63A Jobin et al., 2000; 64 Fais et al., 1991 ; 65 Fedriani, 1996; 66 Fedriani et al., 1998 ; 67 Fragoso and Santos-Reis, 2000; 68 Gazzola et al., 2005; 69 Gil-Sánchez, 1996; 70 Gil-Sánchez, 1998; 71 Kruuk and de Kock 1981; 72 Lagalisse et al., 2003; 73 Llaneza et al., 2000; 74 Lucherini and Crema 1995; 75 Malo et al., 2004.; 76 Marassi and Biancardi, 2002; 77 Martín et al., 1995; 78 Martinoli and Preatoni 1995 ; 79 Martinoli et al., 2001 ; 80 Masseti, 1995; 81 Mattioli et al., 2004;82 Moleón and Gil-Sánchez, 2003; 83 Padial et al., 2002; 84 Palomares, 1993; 85 Palomares and Delibes, 1991a; 86 Palomares and Delibes, 1991b; 87 Pandolfi and Bonacoscia, 1991 ; 88 Pandolfi et al., 1996 ; 89 Pandolfi et al., 1991 ; 90 Papageorgiou et al., 1988; 91 Pigozzi, 1991; 92 Posillico et al., 2004 ; 93 Prigioni and de Marinis, 1995; 94 Prigioni and Tacchi, 1991; 95 Revilla and Palomares, 2002; 96 Rivera and Rey, 1983 ; 97 Rodríguez and Delibes, 1992; 98 Rondinini and Boitani, 2002; 99 Roque et al., 2001; 100 Rosa et al., 1991; 101 Rosalino et al., 2005; 102 Rosalino and Santos-Reis, 2002; 103 Ruiz-Olmo and López-Martín, 1993; 104 Ruiz-Olmo and Palazon, 1993; 105 Santos et al., 2007; 106 Sarmento, 1996; 107 Serafini and

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Lovari, 1993; 108 Such and Calabuig, 2003 ; 109 Torre et al., 2003; 110 Virgós et al., 1996 ; 111 Virgós et al., 1999 ; 112 Vos, 2000; 113 Yom-Tov et al., 2007; 114 Lozano et al., 2003; 115 Aymerich, 1982; 116 Ragni, 1981 ; 117 Gil-Sánchez et al., 1999; 118 Bonesi and Palazon, 2007; 119 Ceña et al., 2003; 120 Bravo and Bueno, 1999; 121 Vidal-Figueroa and Delibes, 1987; 124 Fedriani et al., 1999; 125 Delibes et al., 2000; 126 Bueno and Bravo, 1985; 127 Palazón and Ruiz-Olmo, 1997; 128 Palma, 1980; 129 ICN, 2003 ; 130 Calzada, 2000; 131 Beltrán and Delibes, 1991; 133 Castro and Palma, 1996; 134 Fernández et al., 2006 ; 135 GilSánchez et al., 2006; 136 Stahl et al., 2002 ; 137 Stahl et al., 2001a; 139 Bueno, 1994; 140 Molinari et al., 2001; 143 Bravo, 2002; 144 Lodé, 1999; 145 Lodé, 2000; 148 Marcelli et al., 2003; 149 Rondinini et al., 2006 ; 150 Zabala et al., 2005; 151 Virgós, 2002; 152 Mestre et al., 2007; 153 Clavero et al., 2005 ; 154 Clavero et al., 2006 ; 155 Pedroso and Santos-Reis, 2006; 158 Remonti et al., 2006; 160 Azcón and Duperon, 1999; 161 Sales-Luís et al., (2007); 162 Ruiz-Olmo et al., 1997; 163 Beja, 1991; 164 Bifolchi and Lodé, 2005; 165 Prigioni et al., 2006a; 166 Prigioni et al., 2006b; 168 Clavero et al., 2004; 169 Ruiz-Olmo, 2006; 171 Beja, 1996; 172 Adrián and Moreno, 1986; 173 Adrián and Delibes, 1987 ; 174 Callejo and Delibes, 1987; 175 Ruiz-Olmo et al., 1989 ; 176 Arcá and Prigioni, 1987; 177 Prigioni et al., 1991; 178 Stahl et al., 2001b; 179 Prigioni and De Marinis, 1995; 182 Fournier et al., 2007; 183 Ruiz-Olmo 1987; 184 Lodé 1993.

18

Luciano M. Verdade, Luis Miguel Rosalino, Carla Gheler-Costa et al.

While canids (Chrysocyon brachyurus, Cerdocyon thous and Lycalopex vetulus) are the most generalist species in Southeastern Brazil (Figure 2), in the Mediterranean Region mustelids, viverrids and herpestids, share this generalist trophic behavior with canids (Table 1). Large-sized carnivores such as jaguars, pumas and wolves are frequently associated with large-sized livestock depredation (i.e., cattle and sheep). Nevertheless, smaller species can also be associated with small-sized livestock (i.e., chicken and fish) depredation (Table 1) (e.g., Genet consumption of ducks - Rosalino and Santos-Reis, 2002; Eurasian otter consumption of stocked fish – Freitas et al., 2007).

4. DISCUSSION

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4.1. Mesopredator Release The present results suggest a possible competition between large- and middle-sized Carnivora species as both have small-sized mammals as preys. In addition, there seems to be some consumption of middle-sized mammals (including carnivores) by large-sized species in Brazil, and by many species in Europe. In such circumstance the local extinction of top predators could decrease competition for food items concurrently as it also decreases predation pressure over medium-sized carnivores. These two processes combined could result on an increase in the fitness of mesopredators, due mainly to an increment in the natality rate (as a result of higher food resource availability) as well as a decrease on mortality rate as a consequence of absence of top predators. These two aspects corroborate the mesopredator release hypothesis (Wright et al., 1994; Palomares et al., 1995, 1996; Crooks and Soulé, 1999; Terborgh, 2000; Gehrt and Clark, 2003), which was defined by Prugh et al. (2008) as ―the expansion in density or distribution, or the change in behavior of a middle-ranked predator, resulting from a decline in the density or distribution of an apex predator‖. However, the actual ecological relevance of these aspects is not clear, especially in manaltered and fragmented landscapes. In these environments mesopredator outbreaks can be the result of the disappearing of top predators, due to their need of wider areas, some of which not fragmented, and their higher probability of conflict with man (use of similar resources – e.g. livestock depredation by wolves and jaguars) leading to a higher persecution levels (Prugh et al., 2008). However, fragmentation can also enhance the resources available to mesopredators, especially those directly related with human activities such as pet food, crops and crop pests.

4.1. Increase in Carrying Capacity Some evidences of the increase food availability in agro-forest systems are already available. For example, Silva et al. (2008) showed that agricultural units of a Mediterranean cork-oak landscape recorded the highest abundance and richness levels of ground beetles, which are regularly included in the diet of southern European mesocarnivores (e.g., Rosalino et al., 2005; Santos et al., 2007). Moreover, in southeastern Brazil, sugarcane plantations also support higher densities of rodents than pristine landscapes (Gheler-Costa, 2006). Such

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rodents are an important resource for many carnivores (Rocha et al., 2004; Tófoli et al., 2009). Furthermore, the simple existence of agricultural patches, specially those devoted to fruit production, increase the environmental carrying capacity, in particular to those species highly adaptable in using fruits as a trophic resource, often highly energetic (e.g., olives). For example, a study reviewing fruit consumption by carnivores in Mediterranean Europe showed that more than 20% of fruits eaten by predators such as the red fox, stone marten, pine marten, Eurasian badger and the common genet where originated in orchard or fruit trees plantations (e.g., grapevine Vitis vinifera, loquats Eriobotrya japonica, persimmon Diospyros sp.) (Rosalino and Santos-Reis, 2009). These evidences seem to point out that human altered landscapes (by agriculture or forestry) can be an important food source for more generalist species per se, allowing more foraging opportunities, due to higher trophic resources availability (increasing the landscape carrying capacity), even in the presence of top predators, when comparing with some type of more natural habitats. Actually, the simple fact that most of the studied species use, at least, one type of agro/forest system (see figure 1) seems to indicate that they are using available resources present in those areas, most likely as surplus of food resources. This ecological adaptation can develop in short/medium time scale, with species starting to use new resources made available by human activities. For example, since the beginning of the 1980‘ fish farming has been implemented as an important industry in Portuguese estuaries, producing fish species which are not naturally abundant in those areas (Freitas et al., 2007). Recent studies have determined that 60% of prey consumed by otters in fish farming areas are produced marine fish species (Freitas et al., 2007), and that fish farms had a 76% visiting rate by otters, indicating that otters are using a surplus food resource, nowadays highly available, but probably less common 30 years ago. We thus believe that mesocarnivores are adapting to take advantage of a trophic resource enhancement opportunity window created by agro-systems practices, which increase the overall landscape carrying capacity, some of which are shaping the landscape for thousands of years (e.g., Mediterranean Region) (Pinto-Correia and Vos ,2004).

4.3. Conservation Value of Agricultural Landscapes It is possible and necessary to promote good management practice that allows a balanced promotion of biodiversity and maintenance of production, integrating conservation with production objectives. This can be achieved through several approaches. For example, the main problem when dealing with monocultures is the habitat homogeneity. Maintaining landscape connectivity is difficult when dealing with agricultural landscapes, especially in intensive management schemes. The presence of water courses (rivers and streams) with riparian vegetation, constitute areas of high biodiversity (Naiman et al., 2005), including not only typical aquatic species (e.g. Eurasian otter and European mink in Europe; neotropical otter in Brasil), but also more terrestrial carnivores that use these corridors as refuge, access to prey (e.g. fish, crayfish, birds), routes for juvenil dispersion, and often as the only habitats that enable successful reproduction in a intensive agriculture landscape (e.g., Virgós, 2001; Matos et al., 2009). Therefore, the maintenance of an ecological flow and improvement of the riparian vegetation in this water courses constitute an important contribute to improve biodiversity. The example of the legal obligation regarding maintaining buffers of vegetation

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Luciano M. Verdade, Luis Miguel Rosalino, Carla Gheler-Costa et al.

(30m) along the water courses in Brazil is, therefore, one to maintain and promote (Metzger et al., 2010). Another example is based on the fact that ecological corridors can be accomplished by the conservation or reimplementation of patches of adequate habitat for certain species, or species guilds. This can be achieved by maintaining or creating hedges or walls for smaller carnivores (Macdonald et al. 2007), or larger patches of favorable habitat for all carnivores, in general. Traditionally, hedgerows are part of agricultural landscape in Europe, as well as a cultural element of the landscape structure and diversity. Therefore, these structures may act as important commuting and hunting routes, allowing the survival of some species which use these areas as refuges while feeding on the contiguous agricultural fields. In some cases the fields' boundary areas comprise the most species-rich prey community (Tattersall et al., 2002) resulting in ideal hunting grounds for carnivores, where protective cover is associated to higher prey density. Often, these corridors do not have to be prime habitats. Regarding the Iberian lynx, a species considered highly specialist in habitat requirements (and prey), Palomares et al. (1991) stated that these corridors can even support moderate habitat degradation due to human activity. However, this species uses mixed landscapes, with Mediterranean woods and scrubland, and avoids intensive plantations and agriculture (Rodríguez and Delibes, 1990). Agricultural landscapes are also commonly used as grazing pastures. The establishment of areas without pasture, mainly in extensive exploration schemes, allows the growth of grasslands and scrubland mosaic, which could promote some prey species availability (e.g., wild rabbit – Monzón et al., 2004), and consequently be used by several carnivore species that exploit those food resources (Verdú et al., 2000; Ferrer and Negro, 2004). The importance of this mosaic is well expressed in the Iberian cork and oak woodland (Quercus suber and Quercus rotundifolia), the ―montado‖ in Portugal and ―Dehesa‖ in Spain. These are mixed farming landscapes around extensive woodlands, interspersed with patches of scrubland, grassland and cultivated fields. The montado/dehesa is one of the best examples for the balance between biodiversity and human activity (Pinto-Correia and Vos, 2004), providing both economic value (from the extensive cattle raising, the cork extraction of the Q. suber, mushrooms collecting, ecotourism) and natural value (which encompass a wide range of carnivores, including the wildcat, polecat, genet, stone marten and Iberian lynx). Another important aspect regarding increasing carrying capacity for biodiversity is the presence of small and medium-sized water reservoirs. These aquatic systems, common in the Mediterranean region, where river flows are irregular and many dry out during summer, constitute a water source for cattle and, sometimes, irrigation for agriculture. The role of these systems in the ecology and conservation of carnivores is still somewhat unknown. However, it is obvious that they constitute resources for mammalian carnivores (among others), namely water and prey availability. For example, Basto et al. (in press) detected that the abundance of American crayfish (Procambarus clarkii) was one of the most important variables positively associated with the Eurasian otter‘s use of these smaller reservoirs. Moreover, in more open agriculture landscapes, such as arable steppes or pasturelands, the vegetation developing in these reservoirs margins can also provide some refuge for mesocarnivores. The positive association of the reservoirs‘ use with the availability of refuges was already referred in other studies (e.g. Elliot, 1983; Prenda et al., 2001). Ideally, the maintenance or creation of these water bodies, usually associated with livestock production, should be accompanied by a moderate/low use by cattle, which is rather difficult. Higher cattle pressure promotes a high

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disturbance around the reservoir and reduces cover, and consequently refuges, for carnivore species. Conservation of pristine habitats, still occurring in agriculture landscapes, is also an important way of ensuring the maintenance of biodiversity in these human altered habitats, especially because they can offer protective cover which is often lacking in the agricultural fields. Most of these patches remain in the agricultural landscape due to difficulty of implementing agriculture practices in those terrains. Valleys and steep slopes are among these areas. However, they have been greatly reduced in the last decades (Tscharntke et al., 2005), maybe due to the increase of machinery capability and pressure for soil use. Presently, in some countries where intensive agriculture has led to an alarming level of ecological degradation, an increasing number of efforts are now being made to restore agricultural landscapes and enhance biodiversity by implementing agro-environmental programs, such as introducing semi-natural habitats and field margins into farmland (Jeanneret et al., 2003).

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4.4. Adaptive Processes of Mesopredators to Agricultural Landscapes Time in evolutionary processes should be counted in number of generations of the population in question and not in years (Simpson, 1949). In addition, genetic adaptive changes tend to be faster in changing environments (Levins, 1968). Alterations in food resources availability and spatial-temporal heterogeneity tend to act as selection forces towards individuals. On a single generation time scale, individuals tend to respond to such alterations by behavioural changes (Tuyttens and Macdonald, 2000) which can be called ―acclamation‖. Individuals‘ success in such circumstance depends basically upon their phenotypic plasticity (Relya, 2004). On the other hand, along a certain number of generations a population tends to respond to environmental changes by genetic changes (Dobzhansky, 1970). Populations‘ success in such circumstance depends basically upon their genetic variability (Sinclair et al., 2006). Both acclamation and adaptation to agricultural landscapes should involve changes in two basic behavioural-ecological processes: feeding ecology and use of space. The former is related to changes in food resources availability whereas the later is related to the land use change in spatial terms where, for instance, shelters quality and availability may alter. Mesocarnivores seem extremely plastic both in terms of diet and space use. The present results (high number of generalist species in both European and South American carnivore guilds) corroborate the former whereas their apparently higher abundance in agricultural landscapes (e.g., Dotta and Verdade, 2007) corroborates the later. The extinction or absence of top predators – usual in agricultural landscapes – may potentiate these characteristics. The present results suggest that mesocarnivores may benefit from agricultural landscapes: directly by the increase in the abundance of fruits (Rosalino et al., 2009) or indirectly by the increase in the abundance of seeds or green matter (plant biomass) and consequent increase in rodents‘ abundance (Gheler-Costa, 2006). Systematized studies about the use of space by mesocarnivores in agricultural landscapes should be prioritized as information about it is still scarce. We need to better understand the possible impact of land shaping mechanisms associated with Human activities (e.g. agro-environmental schemes, habitat management) in order to promote the increase of conservation value of agricultural landscapes, namely regarding carnivores.

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4.5. Possible Ecological Imbalances and the Domestication of Nature The extinction of top predators and consequent ―mesopredator release‖ has been considered to result on an increase in bird predation and their consequent population decline (e.g., Soulé et al., 1988; Crooks and Soulé, 1999; Galetti et al., 2009). Although the present results indicate a consistent consumption of birds by middle-sized carnivores there is no evidence that this consumption is higher in agricultural landscapes and/or in the absence of top predators. However, in general these studies are based on short-term surveys where both dependent and independent variables are circumscribed on a smashed time frame. Thus, there is not enough information available about the possible consequences of mesopredator release on a long-term basis. Middle-sized carnivores are also considered as reservoir for pathogens that can cause diseases in domestic carnivores (i.e., dogs and cats) as well as in humans (Whiteman et al., 2007). There seems to be indeed a lot of contact between domestic and wild carnivores (e.g., predation, physical contact with urine and feces etc, hybridization) (Oliveira et al., 2008). It is possible that such contact increase in the absence of top predator as they prey on both domestic and wild smaller carnivores. However, in this case domestic animals should be excluded from conservation units and even from unprotected natural areas as they compete with wild carnivores for food resources (Lepczyk et al., 2003; Bonnaud et al., 2007; Campos et al., 2007). As small-sized carnivores adapt to anthropogenic environments we might expect a certain process of domestication occurring in agricultural landscapes. The concept of domestic or domesticated nature depends on the human perception of nature (Descola, 1986). However, no matter how subjective the concept of nature and natural can be, the fact that small-sized carnivores may depend on anthropogenic habitats to succeed may become their – and ours - most complex conservation problem.

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Chapter 2

FACTORS AFFECTING SMALL AND MIDDLE-SIZED CARNIVORE OCCURRENCE AND ABUNDANCE IN MEDITERRANEAN AGRICULTURAL LANDSCAPES: CASE STUDIES IN SOUTHERN PORTUGAL Filipe Carvalho, Ana Galantinho and António Mira

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Unidade de Biologia da Conservação. Universidade de Évora, Pólo da Mitra, 7002-554 Évora, Portugal Grupo de Ecossistemas e Paisagens Mediterrânicas – Instituto de Ciências Ambientais e Agrárias Mediterrânicas, Universidade de Évora - Núcleo da Mitra, Apartado 94, 7002-554 Évora, Portugal

ABSTRACT Carnivores have a key role in ecosystems and their populations are declining at an increasing rate. Habitat loss and degradation through agricultural practices are among the most serious menaces affecting carnivore survival. Agricultural landscapes are dominant in Western Europe and in several places worldwide are changing quickly into more intensive practices. Therefore, it is important to understand how small carnivore occurrence and abundance is affected by these changes in order to find ways to manage agricultural and grazing systems in a sustainable way, allowing both biodiversity and production to co-exist. We present three case studies in Natura 2000 areas, mainly covered by private agricultural land (livestock, cereal crops, oak, pine and eucalyptus plantations) in southern Portugal. In areas of dominant traditional agro–silvo–pastoral systems (montado) small and middle-sized carnivores tend to occur in a mosaic of montado and shrubs. We found that the occurrence of one of the forest species was positively related with the density of trees and shrubs, soil organic matter content, and Shannon‘s index of vegetation vertical diversity. The presence of livestock and extension of game-estate areas also seem to influence carnivore occurrence. In areas of extensive cereal crops, the presence of shrubs and age of forest plantations play an important positive role for the carnivore community. Mainly our results suggest that maintaining a sustainable mosaic embracing montado, shrubland and open land areas, may allow higher

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Filipe Carvalho, Ana Galantinho and António Mira species richness and abundance by enhancing connectivity between crucial areas. The implementation of this kind of agricultural practices considering the landowners needs is the key issue to achieve the main carnivore conservation goals in Southern Portugal.

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INTRODUCTION Few areas of the Earth have not yet been affected by human activities (Sunquist and Sunquist, 2001). Worldwide, thousands of hectares of land have been converted to agriculture due to the increasing human population demands. One of the greatest challenges of the 21st century is food security since in 2008 nearly 15% of the world‘s population was considered undernourished and diminishing by 50% the poverty and hunger, by 2015, requires a greater investment in agriculture in developing countries (OECD-FAO, 2009). Currently, 30 countries around the world require external assistance for cereal supplies as a result of natural disasters, conflict or insecurity, and economic problems (GIEWS-FAO, 2009). About 4.3 billion of the 13.2 billion hectares of the world‘s land surface are moderately to highly suitable as rain-fed cropland and accounting for currently cultivated cropland (around 1.4 billion hectares) and for forested and urban/protected areas, 1560 million hectares are still available for cropland expansion (OECD-FAO, 2009). In the past the agricultural expansion has been the main driving force of ecosystem change and since the middle of the 20th century the technological and economic incentives allowed for a quicker conversion and intensification of agriculture causing a pervasive decline in farmland biodiversity in recent decades (Benton et al., 2003; Groom and Vynne, 2006). Presently, the decline of biodiversity associated with agricultural practices is mainly due to the intensification of agriculture, the abandonment of marginally productive but high nature value farmland, and the changing scale of agricultural operations (Henle et al., 2008). Agricultural practices (including crop, livestock farming and timber plantations) are one of the three main causes of habitat fragmentation, degradation and loss (Groom and Vynne, 2006) which ultimately is the highest threat to terrestrial vertebrates and biological diversity (Crooks 2002; Michalski and Peres, 2005). Most of the agricultural practices destroy much of the original vegetation cover leading to changes in the community structure and composition, diversity and behaviour of the native fauna (Lyra-Jorge et al., 2008). Nevertheless, the effect of agricultural practices on biodiversity depends upon the characteristics of the landscape and on the requirements of the existing species. Agricultural landscapes can be very different depending on soil, water, climate, slope and human management (Henle et al., 2008). On the other hand, a specialist species may become locally extinct due to the loss or fragmentation of a particular habitat, while a generalist species may benefit by the opportunities a new habitat may bring such as supplementary food and loss of specific competitors or predators (Crooks, 2002; Gehring and Swihart, 2003; Acosta-Jamett and Simonetti, 2004). A focus on protecting places that simultaneously present high percentage of endemic species and greater areas of habitat destroyed is needed to reconcile biodiversity conservation and world development in the present context of increasing demands for agricultural expansion and intensification which endanger the conservation of biodiversity worldwide. In this sense the Mediterranean Basin is one of the 25 Global Biodiversity Hotspots (Myers et al., 2000; Cuttelod et al., 2008). This region besides having an extremely high diversity of

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flora and fauna is also a mosaic of natural and cultural landscapes (Cuttelod et al., 2008). For thousands of years, human activities and natural ecosystems have coexisted in the Mediterranean in a complex way originating high environmental diversity and fragmentation (Maiorano et al., 2006). Currently most of the remaining ecosystems are highly managed and therefore conservation efforts must be directed to this mosaic of mostly hostile habitat, spatial and temporal heterogeneous created by agricultural activities (Macdonald and Rushton, 2003). In an effective conservation process, human activities should be considered as part of the system since traditional agriculture and pasture allow for high levels of biodiversity (Maiorano et al., 2006). In the Mediterranean Basin, agroforestry systems have shaped the landscape and the vegetation, being able to overcome the highly variable Mediterranean climate (Joffre et al., 1999). One of the most representative habitats that contributed for the hotspot designation of the Mediterranean Basin is the open arable area of South Iberia, also known as pseudosteppes. This habitat found in Portugal and Spain comprises the major winter quarter for European migratory birds (Moreira et al., 2005). This high bird diversity depends on low intensive cereal croplands where a rich and abundant assemblage of insects is commonly found. In the last decades, many of these areas have been facing an agricultural intensification mainly in the most productive land, being the remaining areas abandoned, with the consequent shrub encroachment and afforestation with exotic species (Moreira et al., 2005; VanDoom and Bakker, 2007; Reino et al., 2009). The major consequences of these management practices (or the lack of them) may be the decline of most of the winter migratory birds in Europe, once their resting area used during the winter migration is lost and predation pressure increases associated to the changes in land use (Shapira et al., 2008; Pita et al., 2009). The montado (equivalent to the Spanish dehesa) is the dominant agro-silvo-pastoral system in South-western Iberian Peninsula and 33% of the worldwide cork oak range occurs in Portugal (Pinto-Correia, 1993). This traditional landscape is dominated by cork (Quercus suber) and holm (Quercus rotundifolia) oak trees that emerge from a grassland or shrubland matrix. In the montado although there is a repetition of similar elements, the different densities of trees and livestock, the rotational system of exploration and the different development of the vegetation allow for heterogeneity to exist in an extent necessary for maintaining a high biodiversity (Pinto-Correia and Vos, 2004; Rosalino et al., 2009). The combination between tree stands and grassland areas also enhances higher levels of plant and animal biodiversity (Ramirez and Díaz, 2008). This complex system of Mediterranean landscape remained multifunctional at several levels, nevertheless, in recent decades the montado system has gone through a process of intensification and consequent simplification of land use (Pinto-Correia and Vos, 2004). The increase of tree diseases related with alterations in soil conditions and destructive management practices (e.g. understory removal), the abandonment of the traditional explorations and the reforestation using exotic species (e.g. Eucalyptus spp.) endangers the traditional balance between the montado components that previously assured the maintenance of high levels of biodiversity (Pinto-Correia and Vos, 2004; Santos-Reis et al., 2004). Understanding and predicting the distribution, abundance and persistence of wild species in highly valuable habitats such as pseudosteppes and montado is a key element in conservation planning (Macdonald and Rushton, 2003).

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Filipe Carvalho, Ana Galantinho and António Mira

Although mammalian carnivores have a historic resilience as a group, today many of these species are threatened with extinction (Gittleman et al., 2001). Even if several species are not considered presently threatened, most of the carnivores face a hard human persecution due to their known predatory behaviour, especially in agricultural areas where livestock production and game activities are a common practice (Ginsberg, 2001; Macdonald, 2004; Michalski and Peres, 2005). Mammalian carnivores are wide-ranging and low density species and consequently are very sensitive to habitat loss and fragmentation (Oehler and Livaitis, 1996; Livaitis, 2001; Crooks, 2002; Riley et al., 2003; Lyra-Jorge et al., 2010). The activity patterns of carnivores are influenced by many factors, including habitat fragmentation, temperature variations, limitations of the visual system, risk of predation, competition, social and thermoregulatory behaviour (Zielinsky, 2000). Carnivore‘ conservation must be a worldwide goal, due to their unique function in ecosystems as regulators of the trophic cascades (Gittleman et al., 2001; Mangas et al., 2008). Carnivores can be considered umbrella species due to their generally high home ranges that encompass the vital areas of several other species and therefore habitat management for carnivores may assure the conservation of high levels of biodiversity (Hilty et al., 2006). Surveys regarding carnivores are time-consuming and expensive, which has discouraged many researchers to study these species, especially the big carnivores that have larger home ranges (Sunquist and Sunquist, 2001). However, despite these logistical constraints, the ecological features of carnivores make them good models, if we aim to get a first glance on the effects of landscape fragmentation on wildlife. Indeed, they are excellent tools, if we want to evaluate the responses to fragmentation at different landscapes scales (local, regional and global) (Oehler and Livaitis, 1996). On the other hand, small sized carnivores like the majority of the species occurring in Portugal due to their small home ranges and higher densities can be good surrogates to evaluate the effect of habitat changes on carnivore community, when compared with larger species. Some forest species as the American martens (Martes americana) have already been studied for these purposes with good and encouraging results in other regions (Hargis et al., 1999). In this chapter, we will try to present and discuss the response of small and middle-sized carnivores to the two main types of agricultural activities that shape the landscape in southern Portugal: 1) the montado as an agro-silvo-pastoral ecosystem; and 2) the Mediterranean arable land as a mosaic of crops. These two landscapes are particularly important due to their extent, high biodiversity and high degree of threat by the trend in agricultural intensification accordingly to the European Union. Three case studies will be described in order to disentangle the major factors affecting carnivores‘ occurrence and abundance in these two important agricultural landscapes. These case-studies are particularly important because the data were gathered in Natura 2000 sites mainly composed by private property. In these lands active agricultural management is directly related to local livelihoods in a rural socio-economic context dominated by extensive agricultural practices since many generations ago in Alentejo, southern Portugal. The first two case-studies are based on papers in preparation (Carvalho et al, in prep; Carvalho and Mira, in prep) and the third case study is based on a recently published paper (Galantinho and Mira, 2009).

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CASE STUDIES

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Forest Plantation Effects on Middle-Sized Carnivore Assemblages in an Open Arable Landscape: A Preliminary Approach One of the most important agricultural landscapes prevailing in South Portugal is the open arable land, which is a result of extensive cereal croplands campaigns initiated in the last century (see Figure 1) (Moreira et al., 2005). During these campaigns, holm and cork woodlands were replaced by large open areas with cereal croplands. Consequently, several small and middle-sized carnivores considered habitat specialist as wild cats (Felis silvestris) and genets (Genetta genetta), which depend largely on woodlands and shrublands for resting and hunting sites, have been declining from those areas. On the other hand, generalist predators as feral dogs (Canis familiaris) and foxes (Vulpes vulpes) became predominant in these landscapes based on their capacity to explore all kind of habitats and food resources (Mangas et al., 2008; Pita et al., 2009). In recent years, mainly after Portugal joined the European Union in 1986, due to the consequent new funding policies promoting production and agriculture intensification (e.g. cereal crops, livestock, and tree plantation) several changes have occurred in this landscape (Wade et al., 2008). In the last two decades, some of the economic incentives have lead to new forest plantations with umbrella pines (Pinus pinea) and holm (Quercus rotundifolia) and cork oaks (Quercus suber). Although tree plantations enhanced edge effects on otherwise open areas, these new plantations should be considered as causing soft edge effects when compared with the hard edge effects caused by old plantations already established in southern Portuguese landscapes (Reino et al., 2009). The main goal of this study was to evaluate the effects of afforestation, since the early nineties, on the occurrence and abundance of small and middle-sized carnivore assemblages in the Special Protection Area (SPA) of Castro Verde. Secondly, we aimed to know which are the main landscape features influencing the occurrence of middle-sized carnivores in this kind of environment. The landscape is flat and dominated by an agricultural mosaic of cereal, fallow and cultivated fields originated by rotational dry cereal cultivation (Figure 1). Until few years ago, tree cover was largely restricted to some old eucalyptus (Eucalyptus spp.) plantations and open holm oak (Quercus rotundifolia) woodlands grazed by livestock. The climate is typically Mediterranean, with hot and dry summers (average temperature of 24ºC) and cold winters (average temperature of 9ºC), the majority of the rainfall (500-600mm) occurring between October and March (Reino et al., 2009). In 2005, we sampled carnivores through ‗sign surveys‘ twice (summer and autumn) during one hour and half in each of the 60 sampling units located inside or in the surroundings of the study area (Figure 1). Fifty sampling units included edges between forested (of different ages and types) and open agricultural land, and ten consisted only in open agricultural areas (agricultural mosaic of cereal, fallow and ploughed fields). We chose a circular sampling unit of 300 m radius (~28 ha) to achieve a good balance between behavioural ecology of the species inhabiting the area, the size of the core areas of their known home ranges, the aims of the study and the available resources (Wilson and Delahay, 2001).

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Figure 1. Study area in Castro Verde Special Protection Area. The location of the 60 sampling units and a detail image of the main habitat occurring in the area are also presented.

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Table 1. Description and summary statistics of environmental variables (selected during exploratory analysis) used to examine carnivore responses to landscape structure and configuration in the study area Acronym DAMS DIRT_ROADS ROADS RIP_AREAS SHRUB REC_MT_SHRUB OLIVE_GROVES PINE_FORESTS PASTURELAND URBAN_AREAS REC_FOREST WATER_STR FOREST_AGE FOREST_TYPE DIVERS

Description Proportion of area with dams (ha) Proportion of area with dirt roads (ha) Proportion of area with roads (ha) Proportion of area with riparian areas (ha) Proportion of area with shrubland (ha) Proportion of area with recent montado with shrubs (ha) Proportion of area with olive groves (ha) Proportion of area with pine forests (ha) Proportion of area with pastureland (ha) Proportion of area with urban areas (ha) Proportion of area with recent forestation (ha) Length of water streams per plot (m) Age of forests inside each plot a (categorical) Type of forests inside each plot b (categorical) Shannon diversity index

a

Mean ± S.D 0.19 ± 0.07 1.51 ± 0.11 0.88 ± 0.35 0.79 ± 0.24 4.34 ± 1.39 0.77 ± 0.62 0.39 ± 0.17 9.06 ± 2.12 63.39 ± 2.41 0.15 ± 0.12 17.47 ± 2.79 210.62 ± 13.90 2* 2* 1.21 ± 0.04

Age of forests: 1 – without any type of forests (n =10); 2 – recent forests (< 5 years; n = 26); 3 – young forests (between 5 and 15 years; n = 11); 4 – Old forests (> 15 years; n = 13). b Type of forests: 1 - without any type of forests (n= 10); 2 – oak forests (n = 24); 3 – eucalyptus forests (n = 11); 4 – pine forests (n = 15). * For categorical variables the mode are presented.

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Relative abundance indexes (RAI) for each species were estimated considering the total number of signs counted per each hour of search (Long et al., 2008). Scats of all species detected were identified based on their shape, size, location and smell. Any ambiguous identification of scats and footprints was discarded from data analysis (Barea-Azcón et al., 2007; Beja et al., 2009; Mangas et al., 2008). In each sampled plot, surveys were done along linear structures in the landscape such as dirt roads, fences, field edges and riverbanks, where signs of most small carnivore species are commonly found (Mangas et al., 2008). Other prominent structures in the landscape such as rocky areas and old trees were also surveyed to avoid false absences, because some species (e.g. genet) usually use them for latrine deposition (Espirito-Santo et al., 2007). On each sampling unit, several variables (Table 1) related to land use and landscape metrics (e.g. proportion of pastureland, urban areas and length of water streams) were computed using a GIS program Arcview 3.2 (ESRI, 1999) and the Patch analyst 2.2 extension (Eikie et al., 1999). Moreover, we analysed two categorical variables (both with four levels) concerning forest type and age of forest plantations. We compared the relative abundance and species richness of small and middle-sized carnivores between seasons using the Man-Whitney test. Furthermore, we compared those parameters among different types and ages of existing forests through analysis of variance (ANOVA) and/or nonparametric tests (Kruskal-Wallis test), whenever the Levene‘s test for homogeneity was violated (Sokal and Rohlf, 1997). Finally, we ran a multivariate redundancy analysis (RDA) for all species of carnivores on which reliable statistics analysis were possible (more than 10% of presences registered) (Zuur et al., 2007). All non collinear (R > 0.7) explanatory variables were considered in this analysis (Tabanick and Fidell, 2001). Throughout the 180 hours of signs search during the summer and autumn of 2005, we observed 388 signs belonging to eight carnivore species: red fox, Eurasian badger (Meles meles), Eurasian otter (Lutra lutra), stone marten (Martes foina), Egyptian mongoose (Herpestes ichneumon), genet, domestic cat (Felis catus) and domestic dog. The carnivore assemblage was diverse and included almost all the common carnivore species that usually occur in southern Portugal (see e.g. Santos et al., 2008; Matos et al., 2009; Pita et al., 2009). The majority of the signs found were footprints (78.65%) followed by scats (19.01%) and less frequently others signs such as fur, skulls and prey remains (2.34%). The mean number of species registered per sampling area was 2.28 ± 1.21 (Mean ± Standard Deviation) of which 1.30 ± 1.06 were wild species and 0.98 ± 0.34 were domestic. The most common and abundant species were red foxes (RAI = 0.60 ± 0.01) and domestic dogs (RAI = 1.17 ± 0.17), matching respectively 28% and 54% of the signs recorded. Because of that only all predators, red foxes and feral dogs were used in comparison analysis. The remaining species occurred in less than 25% of the sampling sites, being the common genets (RAI = 0.02 ± 0.01) and feral cats (RAI = 0.02 ± 0.01) the least common. No significant differences (p > 0.05) in relative abundance and species richness were found between seasons, neither for all predators and for the red fox and the feral dog considered individually (Man-Whitney test). However, the values were higher during the autumn season, perhaps because the rainy periods have lead to a mud accumulation in dirt roads, being the footprints easier to found and identify. Nevertheless, most of the juvenile dispersion, depending on carnivore species, starts on final summer and reaches the peak in autumn (Virgós et al., 2001; Matos et al., 2009), which may also contributed for these results for the wild species. Concerning the type and age of the forest plantation, we did not find

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significant differences for all predators‘ species, red foxes and feral dogs (Kruskal-Wallis test). The type of forest was expected to influence the diversity and the abundance of carnivores in the study area, because forests of exotic species are known to harbour less biodiversity than oak forests (Virgós et al. 2002; Pita et al. 2009). However, in our study carnivores were more abundant in areas of exotic species (pine and eucalyptus forests). This apparent paradox may be explained by the age of the forest plantations. In fact, most of the eucalyptus forests sampled were the oldest in study area, and presented highly developed undercover vegetation which harbours a great diversity of food resources and shelter sites, not found in the surrounding open agricultural matrix (Mangas et al., 2008). Conversely, the majority of the oak forests were planted less than five years ago. These recent oak plantations are structurally very similar to the open agricultural matrix and therefore the above result was not surprising. However, the more diverse assemblages of carnivores were found on the few old oak forests existing in study area (results not shown), which may reflect the tendency for a higher diversity of carnivores in mosaic landscapes (Gehring and Swihart, 2003; Matos et al., 2009; Pita et al., 2009). Clearly, the results highlight the fact that oldest forest plantations have the great carnivore relative abundances in the study area. Indeed, a positive linear relationship between age forest plantation and carnivore abundance was found (result not shown). This relationship was already documented in literature concerning the Mediterranean context (e.g. Pita et al., 2009). The results of redundancy analysis (RDA) have shown that several other descriptors influence carnivore‘ community in SPA of Castro Verde: DIRT_ROADS, ROADS, FOREST_AGE, OLIVE_GROVES and URBAN AREAS (Figure 2). The RDA showed a positive influence of the FOREST_AGE (p 51 % (ha) GRASS Area with arable or grassland (ha) MONT1 Montado with a low c.cb < 30 % (ha) MONT2 Montado with a middle c.c 31-50% (ha) MONT3 Montado with a higher c.c > 51 % (ha) Oli Olive groves (ha) MORTMONT Percentage of dead trees in montado landscape (%) PCHAR Trees affected by charcoal canker (%) Soil type S_ARG Low unsaturated clayed soil (ha) S_LIT Dry soils (ha) S_IC Incipient soils (ha) Copyright © 2011. Nova Science Publishers, Incorporated. All rights reserved.

a

Mean (± S.D)c 1.91 ± 0.48 0.02 ± 0.01 11.18 ± 4.34 7052.79 ± 4572.05 1323.08 ± 1820.32 0.21 ± 0.09 22.27 ± 26.49 2.27 ± 6.83 4.31 ± 8.63 46.55 ± 28.17 17.64 ± 19.49 22.20 ± 22.07 27.85 ± 28.22 5.84 ± 13.92 0.29 ± 0.42 8.29 ± 5.74 47.87 ± 28.13 21.96 ± 25.31 1.67 ± 3.44

s.c = shrub cover; b c.c = crown cover; c S.D – standard deviation.

Prior to analysis, the spatial autocorrelation was tested for each carnivore species between sampling sites, using the Moran‘s I and accounted for its significance with a z-test computed with the script of Lee and Wong (2001) for Arcview (ESRI, 1999). We employed a Binary GLM (generalized linear models) analysis to predict the presence/absence of middle-sized carnivores using landscape metrics, land cover and soil type as explanatory variables (Table 2) (Hosmer and Lemeshow, 2000; Long et al. 2008). The capacity of the independent variables to discriminate between the presence and absence of middle-sized carnivores was examined by univariate regression during the exploratory data analysis phase (Wald test; p 0.70) (Tabachnik and Fidell, 2001). All the possible biological plausible two–way interactions, between significant or nearly-significant (p < 0.1) independent descriptors, were also considered (Hosmer and Lemeshow, 2000). During the Binary GLM models construction, variables selection was performed by the forward elimination procedure (Hosmer and Lemeshow, 2000). Evaluation of models fitting was based on the classification tables (proportion of cases correctly classified) and ROC curves (Manel et al., 2001).

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Table 3. Summary statistics of the final best models selected to predict the presence of middle-sized carnivore through scent stations in Monfurado study area Red fox Model AWMSI*MONT2 MONT2 AWMSI Intercept Genet Model DIVERS*PCHAR MAT1 Intercept Stone marten Model MONT1*NUMP AWMSI*MONT2 Intercept Badger Model PCHAR*S_ARG MORTMONT*OLI MORTMONT Intercept

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a

βa -0.119 0.256 4.844 -9.731

SE b 0.062 0.120 1.966 3.376

Wald c 3.708 4.550 6.070 6.641

Sig. d 0.054 0.033 0.014 0.010

1.026 0.054 -3.045

0.408 0.018 1.013

6.314 9.227 9.045

0.012 0.002 0.003

0.003 0.019 -2.400

0.002 0.009 3.376

3.729 3.959 9.970

0.053 0.047 0.002

-0.011 -5.596 5.162 2.154

0.005 2.573 2.635 1.332

5.606 4.730 3.839 2.618

0.018 0.030 0.050 0.106

β – Estimate regression coefficient; b S.E – Standard error; c Wald statistic; d Significance level.

Models cross-validation was done through a Jackknife procedure, with ―leaving-one-out‖ methodologies (Olden et al., 2002). A total of 600 scent-station nights were implemented. Using this methodology, we found six small and middle-sized wild Iberian carnivores: red fox in 55% of the sampling sites, genet (47.5%), stone marten (30%), (Meles meles) (27.5%), Egyptian mongoose (7.5%) and weasel (7.5%). We also recorded domestic dogs and feral cats (Felis catus) but they were not taken into account in this study. Statistical modelling was done only for the species recorded in more than 25% of the sampling sites (Table 3). The Moran I results show that there were no significant spatial autocorrelation for any of the species (-1< Z 0.05) (Lee and Wong 2001). The best Binary GLM models for all species correctly classified more than 75 % of cases and presented a useful adjustment by the ROC curve (useful accuracy = 0.7-0.9), except for the stone marten model which only showed a near acceptable accuracy (AUC stone marten = 0.67). Cross-validation validated all models (< 50 % error classification for p < 0.05), meaning that they can be used in other areas with similar landscapes. The red fox occurred more often in fragmented areas [significant positive association (+ β) with AWMSI] (Eikie et al., 1999). Foxes are able to survive in open landscapes, so they are less susceptible to forest loss than other small and middle-size carnivore species (Virgós et al., 2002). Moreover, larger and more vagile species, like the fox, are more prepared to cross and hunt in deforested areas (Gehring and Swihart, 2003). However, even for these species the presence of shrubs and trees are very important for shelter and den sites (Virgós et al., 2002). In fact, areas with 31% to 50% of tree canopy cover (MONT2) have a significant positive association with the presence of red fox in Monfurado. Nevertheless, the interaction between MONT2 and AWMSI had a negative influence in the presence of the red fox,

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probably meaning that the effect of the landscape fragmentation on fox occurrence depends on the montado patches area. This may reflect a possible threshold in the fragmentation degree, even for a generalist species (Pita et al., 2009). The presence of the genet was significantly associated to MAT1 (areas with shrub cover under 30%, see details on Table 2). Lower to moderate density values of shrubland are preferred by genets as shelter, resting and reproduction sites (Palomares and Delibes, 1988 and 1994; Virgós and Casanovas, 1997; Carvalho and Gomes, 2004; Galantinho and Mira, 2009). This habitat seems to have a great abundance of small mammals like rodents, and passerines that together represent the greatest amount of biomass in the diet of the genet in the Iberian Peninsula (Virgós et al., 1999; Rosalino and Santos-Reis, 2002; Carvalho and Gomes, 2004). Areas with a great diversity of vegetation and percentage of trees affected by charcoal canker are significantly associated with the presence of the genet. The index of plant structural diversity reflects the vertical structure of vegetation in the landscape, which is a crucial aspect for a forest carnivore as the genet. In fact, this carnivore has territorial habits, where faecal marking behaviour occurs normally in trees (cork and holm oaks), but also on the ground inside the core areas in a moderate density of shrubland (Palomares and Delibes, 1994; Virgós, 2001; Carvalho and Gomes, 2004; Galantinho and Mira, 2009). In Monfurado throughout the field work, we also found genet latrines in rocks, when no trees were found (Carvalho and Mira, in prep). The charcoal canker disease is derived by the contamination of a fungus denominated Nummularia regia (syn. Biscogniauxia mediterranea) and affects cork oak trees (Nugent et al., 2005). This fungus destroys the cork tree tissues (phloem and xylem), so the tree becomes dark (carbon colour) and hollow. Genets use those holes as shelter to rest, for reproduction and also for minimizing the risk of predation by dogs and red foxes (Palomares and Delibes, 1994; Santos-Reis et al., 2004). The presence of the stone marten was positively associated with two interactions: MONT1 (tree canopy cover below 30%) and NUMP (number of patches); MONT2 and AWMSI. Both interactions deal with predictors directly related with forest cover and habitat fragmentation. As for the red fox, the stone marten model seems to reveal the existence of a threshold in the rate between matrix (open land) and forests above which the species disappear, as already stated for other carnivores (Oehler and Livatis, 1996; Hargis et al., 1999; Hilty and Merenlender, 2004; Pita et al., 2009). Stone martens, despiste their forest preferences, can tolerate human presence, whenever the associated extra food resources exist, like fig trees, apple trees and wild roses (Hargis et al. 1999; Rondinini and Boitani, 2002; Santos-Reis et al., 2004). Moreover, stone marten seems to prefer areas where small forest patches are embedded in open areas. These areas may be favourable to stone martens, because an increase in the abundance and diversity of small mammals and passerines associated with clear cuts occurs and still permits movements between patches inside the matrix (Virgós, 2001; Rondinini and Boitani, 2002; Virgós and García, 2002; Virgós et al., 2002). The badger was positively associated to MORTMONT (the percentage of trees affected by diseases in montado landscape). This species digs near tree roots to find food (e.g. insects) which is an easier task around sick or dead trees (Moore et al. 1999). Furthermore, the places surrounding dead trees are areas with more organic matter which enhances the abundance of the soil fauna (Revilla et al., 2000; Rosalino et al., 2003, 2005b). However, when some tree diseases affect orchards and olive groves, their fruit production might be compromised. Fruits are a very important food resource for generalist carnivores like the badger. Actually, our data showed that the probability of occurrence of badger decreases in areas where the percentage of diseases

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increases associated with olive grooves. The decrease in fruit availability especially in some seasons as in winter (e.g. olives) can be crucial for the dispersion of young badgers (Revilla et al. 2000; Rosalino et al., 2005b). In the Mediterranean region, the densities of badger populations are low, when compared with the populations from North Europe (e.g. UK) probably due to the high xericity in southern Iberian Peninsula, where factors influencing location of den sites must be critical. The negative interaction between charcoal canker (PCHAR) and loam soils (S_ARG) may reflect a higher degree of human disturbance. In fact, the charcoal canker affects highly trees from genera Quercus sp., which result in an economical constraint in study area concerning cork production. The higher human presence due to this disease control may affect den site fidelity by badgers, even if the density of den sites should be higher due to a good soil for digging dens (Mickvicius, 2002; Rosalino et al., 2005a). However, badger may have to travel greater distances to find food safely (Revilla et al., 2001). Although the models obtained had high predictive power, we should interpret our results with caution and consider them as a tool which may state out trends about the way Iberian carnivores can be distributed in the montado landscape (Hilty et al., 2006). However, we think that these results are in concordance with others surveys realized in Iberian Peninsula for small and middle-sized carnivores (Virgós, 2001; Virgós et al., 2002; Mangas et al., 2008; Matos et al., 2009; Pita et al., 2009; Carvalho and Mira, in prep). One of the main results highlighted, is that the presence of large montado landscape areas, together with a moderate shrubland density interspersed with open lands is one of the best habitats for most of the Iberian carnivore species (Virgós, 2001; Pita et al., 2009). Montado landscapes depend on extensive human related activities and some of our results support the fact that human intervention is critical to maintain favourable conditions for carnivore occurrence (Ramirez and Díaz, 2008). Once identified the most significant factors influencing the presence of carnivores, this information should be incorporated in management plans aiming biodiversity conservation. We strengthen the fact that effective management actions in human-dominated landscapes should take place not only on the Natura 2000 sites but also outside these areas.

Human, Ecological and Livestock Influences on Genet (Genetta Genetta) Distribution on Mediterranean Farmland The genet is widely distributed and common in Portugal (Santos-Reis and Mathias, 1998), and classified as ‗‗Least Concern‘‘ in Europe (IUCN, 2007). Evaluating the relative importance of specific issues of local management on the distribution of wild species is important if we wish to integrate farmland activities in species conservation plans. An abundant and common forest species like the genet may be a representative of the small and middle size carnivore community in montado dominated landscapes and may be used as surrogate to model the influence of ecological and human related activities on this community as a whole. Although each carnivore species have specific characteristics and needs they share resources (e.g. food, space) where they co-exist. The presence/absence pattern of the genet in southern Portugal was evaluated considering natural landscape features and two important axes of rural management: livestock production and direct human influence. We aimed to identify main factors or groups of factors influencing the distribution of the genet and especially how management practices influence its presence on Mediterranean farmland. As in the previous case-study the fieldwork was conducted in Monfurado a Site included in

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the Mediterranean biogeographic region of Natura 2000 Network in southern Portugal (Figure 3). Thirty six homogeneous habitat patches were used as sampling sites. These sites were selected accounting for habitat heterogeneity (agro–silvo-pastoral use, shrub structure, tree density) and representativeness in the study area. On three occasions (summer, autumn/winter and spring), from June 2003 to June 2004, every patch was surveyed walking on a path of approximately 1,000 m long and 25 m wide searching for signs (faeces and tracks) of genet. Latrines were searched in old trees (Palomares, 1993), fallen tree trunks (Queirós, 1989), old decaying roofs and walls (Gomes and Giraudoux, 1992), large rocks (Roeder, 1980; Palomares and Delibes, 1994; Virgós and Casanovas, 1997), and raised ground (Palomares, 1993; Virgós and Casanovas, 1997). Twenty-eight explanatory variables were collected or derived from a 500 m radius sampling circular plot centred in each of the patches (see for more details Galantinho and Mira, 2009). The variables were grouped into three sets of descriptors: ecological, livestock, and human influence. Both livestock and human influence groups reflect different types of suspected perturbation. The ecological group included each of the most representative habitats (e.g. HMONT_HSHUB; MEADOWS), Shannon‘s diversity index (SDI), topographic variables (e.g. ITR; ELEV_RANGE; FLAT), soil organic mater (SOIL_OM) and trees affected by diseases (e.g. PCHAR). The livestock variables comprised density and time spent in the plots by sheep and cattle (e.g. SHEEP_D; SHEEP _T). The direct human influence accounted for distance to roads (DIST_ROAD) and villages (DIST_VILL), number of buildings present in the sampled areas (N_BUILD), area covered by small-game estates (SGAME_EST) and shrub cutting (SHRUB_CUT). Moran‘s I was used to test for autocorrelation in the genet data. The influence of each set of explanatory variables on the presence of the genet was assessed by the variation partitioning procedure proposed by Borcard et al. (1992), adapted to logistic regression models and extended to the three sets of variables. The best models for each set were chosen by information–theoretical model comparison (ITMC) (Burnham and Anderson, 2002). Regression coefficients were considered significant at the P < 0.15 level (Rawlings, 1988) and slightly higher P values were also used to account for variables of particularly important biological meaning (Hosmer and Lemeshow, 2000). From all the explanatory variables collected, fifteen were used to complete the procedure (Table 4). Genet‘s presence/absence data was not significantly autocorrelated. Genets were present in half the patches surveyed and its occurrence seemed to depend more upon the ecological factors (pure effect: 30.3%) than on grazing (pure effect: 8.5%) and direct human influence (pure effect: 4.8%) (total variance captured: 56.5%). High cover and the complexity of the vegetation strata, and organic content rich soils seem to be the most important ecological factors influencing the occurrence of the species (Table 5). These factors are often associated to dense shrub and tree cover (Moreno et al., 2007; Moreno, 2008). High understory cover and old trees are important for the placement of latrines in cork oak woodland (Espírito-Santo et al., 2007) and as resting places during the day (Palomares and Delibes, 1988). A dense shrub cover often offers to genets, a higher abundance of the wood mouse (Apodemus sylvaticus) which is considered their main prey, besides shelter against predators and competitors (Cugnasse and Riols, 1984; Palomares and Delibes, 1988, 1994; Virgós and Casanovas, 1997; Virgós et al., 2001). Although cork oak trees are one of the most important sources of profit for landowners, shrub areas are considered as potentially dangerous as highly inflammable and as obstacles in the fire extinguishing process.

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Table 4. Variables groups, abbreviations, description and summary statistics for significant explanatory variables selected during exploratory analysis and included in models for predicting occurrence of the genet in Monfurado. We present the mean ± standard deviation (Mean ± SD) of significant continuous variables and the number of patches per class of categorical significant variables with (1) (n = 18) and without (0) (n = 18) genet signs. The number of buildings is analysed as a categorical variable because it has only three possible values (Tabachnick and Fidell, 2001)

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The livestock models suggested that moderate grazing intensity by sheep favours the presence of the genet (Table 5). The vegetation structure and species composition of plants and arthropods in shrub and grass layers depends on sheep grazing intensity (Marrs et al., 2007; Woodcock et al., 2005). Low to moderate levels of livestock grazing increase arthropod diversity (see Debano, 2006). Arthropods (e.g. coleoptera, orthoptera, and arachnidae) are the second most frequently consumed prey by the genet after mammals, in Europe (Virgós et al., 1999; Rosalino and Santos-Reis, 2002). Small mammals, birds, and dung beetles are important food items for the genet and are highly available in montados associated with the presence of livestock (Virgós and Casanovas, 1997). Although moderate grazing may have a positive effect in an important food item, higher intensities of livestock have extremely negative effects in biodiversity (Fleischner, 1994; Dobkin et al., 1998; Krueper et al., 2003; Wheeler, 2008). Moreover, intensive stocking rates endanger tree renewal which is essential to the sustainability of the montado system once the sprouting young trees are consumed or destroyed. However, grazing prevents land abandonment and is an important source of income associated to the montado system in the study area. Therefore stocking rates compatible with conservation values must be defined for this and other montado areas. Rotational livestock grazing is a traditional management option recommended to provide habitat for those species requiring short swards and intense grazing, as well as for large, diverse invertebrate assemblages in taller, older swards (Dennis, 2003). The best human influence model suggested that the occurrence of the genet is lower in hunting estates (Table 5). Hunting estates are profitable associations that rely greatly on the contributions of its members and occasional hunters and most certainly on the numbers of game hunted. The main goal of hunting management is to maintain a profitable number of game species and individuals. Often, management goals in these areas include lowering predator numbers. In Portugal, legal predator control consists in trapping and killing foxes and Egyptian mongooses (Herpestes ichneumon). However, predator control actions can influence and threaten non-target species, like the genet, once they are inherently not selective and often raise negative effects on the overall carnivore guild composition and diversity (Duarte and Vargas, 2001; Virgós and Travaini, 2005). In fact, cage-traps have to be inspected daily to release non-target species otherwise they will die due to lack of food and water. Moreover, often people believe that all small and medium size carnivores are an eminent threat to game-species and livestock and therefore illegally kill all trapped animals. Other predator control actions are used in many estates although they have already been considered illegal for many years. Poison is one of the most dangerous techniques, because it affects many species of mammal, reptiles and birds. Therefore, environmental awareness campaigns directed to landowners, game-estate managers and hunters about the role of carnivores in the ecosystem should be a generalized measure. The legal process of attributing credentials for using traps should require more strict rules for their placement, checking frequency, and, a better and more effective surveillance system (Duarte and Vargas, 2001; Virgós and Travaini, 2005).

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Table 5. Best partial models for the ecological, livestock and human influence sets. Best partial models, selected based on ITMC are reported along with the effect sizes, their precision and associated P-values. The interpretation of the regression coefficient for categorical variables is based on the first category (indicator)

Globally considering the best models for each set of variables and the variation partition procedure, genet‘s presence was highly associated with management strategies, and indicators of the ecosystem‘s well-being (for more details on all the models compared see Galantinho and Mira, 2009). Soil organic matter content is the most significant predictor in both partial and full models. The maintenance of soil organic matter has been an important goal of several research activities and programmes concerning environment conservation to assure the sustainability and productivity of the natural and agricultural systems (Wolters, 2000). Certainly relevant variables were not analysed since 43.5% of the variance in the presence/absence pattern of the genet was not explained by the three sets of variables used. Probably, including prey availability and microhabitat characteristics would improve the explained variance of our models once predators have to assess and choose patches for prey availability (Stephens and Krebs, 1986) and microhabitat is known to be significant for the placement of latrines in the montado landscape (Espírito-Santo et al., 2007). Nonetheless, we explore issues regarding local farming management and soil conservation practices that are important for the sustainability of agro-silvo-pastoral activities and wildlife management in Mediterranean environments. This is especially relevant for landowners to understand and engage voluntarily in conservation strategies that can initially be considered by them economically disadvantageous (Jacobson et al., 2003) such as maintaining shrub areas and diminishing grazing intensity. However, developing strategies to involve landowners in

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conservation actions should and can go beyond economic incentives once many of them are emotional attached to the land they own. Moreover, pure economic incentives (e.g. subsidies, tax reduction) do not have a long-term existence and effectiveness while the effects of social commitment, recognition and problem avoidance can engage emotionally people and therefore last longer (Jacobson et al., 2003). Finally, univariate data analysis showed that the presence of the genet is positively related to the abundance of the other carnivores present in the study area considered together (Vulpes vulpes, Mustela nivalis, Mustela putorius, Martes foina, Meles meles and Herepstes ichneumon). Some of these species use the same habitats or resources, but not to the same extent or at the same time (see Fedriani et al., 1999; Mangas et al., 2005; Peris and Tena, 2005; Barrientos and Virgós, 2006). Consequently, measures related to the issues explored here might contribute not only to the conservation of the genet but also to the conservation of the whole carnivore community.

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CONCLUSION In this chapter, we aimed to understand the factors associated with the occurrence and abundance of several species of carnivores in agricultural landscapes of the Mediterranean region. Two main agricultural systems were studied: the open arable farmland and the montado. Our research has shown that the less diverse carnivore assemblages occur in the largest open areas, with small remnant forests patches. These areas are dominated by generalist species as red foxes and feral dogs, which may benefit from the fragmentation induced by agricultural activities due to their high adaptability. These two species are more vagile and able to travel greater distances, exploiting new habitats, and eating different food items (e.g. fruits and/or meat) (Gittleman et al., 2001; Gehring and Swihart, 2003; Dotta and Verdade, 2007). The overabundance of these two species may limit the abundance and diversity of other smaller sized and more specialized carnivores, due to competition or even occasional predation (Garrott et al., 1993). Moreover, the enhancement of red foxes and feral dogs will increase predator pressure on key species in Mediterranean environments like the wild rabbit (Oryctolagus cuniculus) or the Iberian hare (Lepus granatensis), which are relatively abundant in Monfurado and Castro Verde, respectively (Shapira et al., 2008). The decrease in rabbit density in Iberia over the last three decades is known to be a main factor contributing to the decline of specialized carnivores of conservation concern like the Iberian lynx and the wild cat (Monterroso et al., 2009; Sarmento et al., 2005, 2009). On the other hand, forest specialist species like genets and stone martens may face local extinction in these open areas in a short period (Hargis et al., 1999). These open areas may become impenetrable and limit the matrix permeability by hampering the easy access to food and resting sites and the dispersion of juveniles of species highly dependent upon forest areas (Lyra-Jorge et al., 2008). Consequently, the lack of sufficient number and size of permanent native forest patches will decrease the probability of persistence of these carnivores‘ populations (Crooks, 2002). In fact, we may find that the percentage of sample units where forest carnivores (genets and stone martens) are present is lower in the mosaic of open land and afforestation of Castro Verde than on the well conserved montado landscape in

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Monfurado. Probably the same trend occurs concerning carnivore abundance, although we cannot compare the abundances obtained between the two study areas once different field methodologies (transects and scent stations) were used in case studies 1 and 2 of this chapter. The montado landscape has always been described as a habitat highly bio diverse, due to its structurally complex vegetation and traditional management system resulting in a particularly resilient and rich mosaic landscape (Ramirez and Díaz, 2008; Pita et al., 2009). In fact, the combination of holm and cork woodlands, shrublands and open grasslands interspersed by riparian galleries enhances a great animal diversity. This combination leads to a complex trophic net, which starts on soil microfauna (e.g. insects, earthworms and spiders) eaten by several insectivorous like shrews, moles, passerines, reptiles and amphibians, which in part became prey of most small and middle sized carnivores like the ones studied in this chapter. In addition, the diverse vegetation structure together with scattered grassland areas harbours great abundances of wild rabbit and red-legged partridge (Moreira et al., 2005; Delibes-Mateos et al., 2007). Consequently, hypercarnivores (meat eaters) like Iberian lynxes, wild cats and weasels (Mustela nivalis) and omnivorous carnivores like genets, stones martens, badgers and red foxes are able to co-exist in these landscapes. The present favourable conservation status of both studied areas reflects the moderate human intervention that shaped these landscapes through centuries and potentiated the existence of a small and middle-sized carnivore community. However, in recent years human activities are also responsible for the main threats to these agro-systems of high conservation value and associated global biodiversity. Economic policies promote the increase in productivity through agricultural and grazing intensification and although refrained inside Natura 2000 Sites, these changes occur in its surroundings. The intensification of agricultural practices has increased habitat fragmentation during the last years in these landscapes and in a significant part of southern Portugal (VanDoom and Bakker, 2007). Habitat fragmentation reduces landscape functional connectivity limiting wildlife movements, particularly for species with larger home ranges, like carnivores, threatening the long-term persistence of populations of these species even inside protected areas. The increase in livestock production has lead to the degradation of vegetation and watercourses while the traditional moderate levels of grazing enhanced the occurrence of some carnivores like the genet. Domestic carnivores like dogs and cats, which compete for food items with wild carnivores are increasingly overspreading due to human presence (Gittleman et al., 2001). The illegal predator control actions that can occur in some hunting estates of Southern Iberia may compromise the stability of the overall carnivore community (Duarte and Vargas, 2001; Virgós and Travaini, 2005). Recent studies in this region also highlight the negative influence roads on several wild carnivores (Galantinho and Mira, 2009; Grilo et al., 2009; Carvalho and Mira, in press). Most of the carnivore species are often road-killed (e.g. stone marten, genet, polecat) and some show road vicinity avoidance behaviour. The complex relation between the crescent human activities and the community of carnivores must be understood if we wish to assure long term persistence and integrity of carnivore communities in the Mediterranean region. The continuous threat upon this biodiversity hotspot highlights our responsible conservation duty. The three studies presented in this chapter were made on Natura 2000 sites, which are extremely important areas throughout Europe. In Portugal, the proportion of Natura 2000 sites and protected areas, already reaches about 21% of the national territory (ICN, 2005). However, not all biological

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diversity can be preserved just inside protected areas while most of the world land is outside them (Maiorano et al., 2006; Dotta and Verdade, 2007). This is especially true for carnivores due to their large home ranges (Crooks, 2002). Enhancing connectivity measures between remnant native patches, where the links to survivorship could be attained especially by the young individuals while in dispersion seems to be fundamental for the conservation of several carnivore species. Mainly studies have to be made in human altered areas, especially in agroecosystems, once they are dominating the south Portugal landscapes (VanDoom and Bakker, 2007). Nevertheless, to be successful we need to understand deeply the effects of human intervention and landscapes changes in the wild carnivores and find new ways to cope with change. We must also inform and involve people responsible for land use change and management about the key importance of carnivores in regulating ecosystems and the implications of land use practices in their ecology if we wish to obtain an efficient way for carnivores to exist in human altered landscapes (Riley et al., 2003).

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Chapter 3

FRUITS AND MESOCARNIVORES IN MEDITERRANEAN EUROPE Luís M. Rosalino* and Margarida Santos-Reis Centro de Biologia Ambiental, Faculdade de Ciências de Lisboa, Portugal

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ABSTRACT Because of the importance of fruits as a food resource to mesocarnivores and the increasing perception of the role of those mammalian predators as effective seed dispersers, it‘s important to review the relation between fruits and carnivore in areas where these plant resources are diverse and important. Therefore, in this chapter we present a review on why fruits, available and accessible to carnivores in the Mediterranean, are important to carnivores. We present a detailed analysis of fruit consumption by mammalian carnivores in the Mediterranean region, pointing out that generalist predators, such as red fox, stone marten, Eurasian badger, and common genet are those which consumed a more diversified fruit diet. In our review, we also detected an increase importance of fruits in the diet of mesopredators in eastern Mediterranean areas. Moreover, we also present data from western Portugal corroborating the hypothesis that mesopredators act as effective fruits dispersers. Although, variation was detected among species, common genets apparently act as the most efficient disperser by significantly promoting germination of pears, olives, and grapes. Finally, we present some considerations concerning fruits, mesocarnivores, and conservation.

1. INTRODUCTION The relation between fruits and carnivores can assume different perspectives. On one hand fruits are a good source of carbohydrates (including fibers), minerals, and water *

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(especially flesh fruits) for mesocarnivores (Herrera, 1987), and are particularly important in areas such as the Mediterranean Basin, where environmental heterogeneity and climatic unpredictability constrains food availability in some seasons (e.g., during hot and dry summers). On the other hand, mesocarnivores are vehicles to disperse the seed contained within fruits, and often promote seed germination. For these reasons understanding this two-way relation is fundamental to untangling the complex relationships between community members and delineating effective conservation and management action focused on many endemic or threatened Mediterranean plant or mesocarnivore species.

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2. WHAT ARE FRUITS? ―Fruit‖ is an expression widely used in biology although not always properly utilized. Some have mention this term while considering gymnosperms cones or even multiple fruits, such as pineapple or figs. However, fruit is the ripened ovary of a flower (i.e., angiosperms) together with any accessory parts associated with it (Lewis, 2002). They are usually formed after pollination and fertilization, but sometimes can also arise without fertilization, such as in commercial bananas, a process called parthenocarpy. The fruit is composed by a pericarp (formed by the ovary walls), which is subdivided in endocarp, mesocarp and exocarp, enclosing the seeds. The seed consists of the embryo surrounded by the cotyledons (or cotyledon in monocotyledon plants), what is left of the endosperm, and a seed coat (Jensen and Salisbury, 1972). Although artificial, fruits are often divided into two categories, which represent different strategies to achieve their main evolutionary purposes, i.e., protection and dispersion of the seeds: 1) dry fruits – the pericarp is dry when fruits are mature (e.g., nuts such as hazel-nuts and acorns; or the follicle such as in peonys); and 2) fleshy fruits – the pericarp is partly or entirely fleshy or fibrous (e.g., drupes like olives; berries such as grapes; or pomes like apples or pears) (Jensen and Salisbury, 1972). Fruits may also vary widely in size, average number of seeds, and chemical components. For example, while the majority of Iberian fruits are relatively short in length (6-8cm e.g., Mastic tree Pistacia lentiscus), some can reach more than 18cm (e,g., pears Pyrus bourgaeana) (Herrera, 1987). An even wider distribution is found in the number of seeds per fruits, which range between one (e.g., olives Olea europaea) to more than 1700 (e.g., figs Ficus carica) (unpublished data). Some have higher water content (more than 85% of the fruit - Autumn Mandrakes Wandragora autumnalis), while other are lipid (59% - Pistacia lentiscus), protein (28% - European white bryony Bryonia dioica), fiber (46% - wild pears Pyrus bourgaeana), or nonstructural carbohydrate-rich fruits (94% - common yew Taxus baccata; Herrera, 1987). Chemical element concentrations also show wide variation, especially copper (Cu– 0.6 to 74.1 mg/g; Asparagus albus and Phillyrea angustifolia , respectively) and iron (Fe – 6.3 to 204.9 mg/g; fly honeysuckle Lonicera xylosteum and St Lucie cherry Prunus rnahaleb, respectively; Herrera, 1987).

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3. FRUITS IN MEDITERRANEAN EUROPE: NATIVE AND INTRODUCED BY MAN The Mediterranean basin is considered to be a crossroads for plant and animal communities, as well as for ancient civilizations. Due to its geographical position, it establishes the bridge between several biogeographical regions (Euro-Siberian, Sahelian, Irano-Turanian, Arabian and Atlantic - Blondel and Aronson, 1999). It is the northern distribution limit of typical African species (such as the Egyptian mongoose, herperstes ichneumon) and the southern limit of European species (e.g., Taxus baccata). These characteristics, together with the fact that many Mediterranean regions were considered to be plant and animal glacial refugia (e.g., Iberia, Balkans), which influence biodiversity patterns in the Mediterranean Basin (Médail and Diadema, 2009), resulted in the high biodiversity of the region, concurrent with a high number of endemic species. These characteristics led Myers et al. (2000) to consider the Mediterranean Basin as one of the 25 biodiversity hotspots in the world. Indeed, the Mediterranean region encompasses about 25.000 plant species (almost 8% of the world species and 50% of which are considered endemic - Blondel and Aronson, 1999), which represent a huge source of available fruits. Additionally, the Mediterranean Basin was the origin of many of the more influential world civilizations including the Sumerian, Akkadian, Egyptians, Phoenicians, Greeks, Romans, and Ottomans (Blondel and Aronson, 1999). All have transformed the Mediterranean landscape, being responsible for the selection of some species with economic values as well as for the introduction of others (e.g., Conyza spp. - Blondel and Aronson, 1999). A good example of the ancient shaping of the Mediterranean areas, is the fact that the Minoan culture in Crete (4000-3000 years BC) started to transform the landscapes around the important city of Kommos into a mosaic of cultivated fields and orchards (Blondel and Aronson, 1999). The presence of a high number of fruit producing plants in Mediterranean Europe, native and introduced by man, transform fruits into a important food resource available to mesocarnivores, especially generalist species that take advantage of resources most available during each season. As a result, the role of mammals as potential fruit dispersers has increased.

4. FRUITS IN THE DIET OF MESOCARNIVORES Several studies in Europe have showed that, for many mesocarnivores, diet composition changes gradually over a latitudinal trend (e.g., common genet Genetta genetta – Virgós et al., 1999; pine marten Martes martes - de Marinis and Massetti, 1995, Zalewski, 2004). Moreover, the importance of vegetable food items, and fruits in particular, seem to increase southward (e.g., Eurasian badger Meles meles - Goszczynski et al., 2000). Some authors have stated that this higher consumption of fruits could be influenced by Mediterranean climates (Rosalino et al., 2005) characterised by hot a dry summers, mild winters, and environmental heterogeneity and seasonal and temporal climate unpredictability (Virgós et al., 1999). However, the Mediterranean region is not climatically homogeneous, with north-south (EuroSiberia to Sahara) and east-west (Irano-Turanian/Arabian to Macaronesia/Atlantic) gradients

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(Blondel and Aronson, 1999) that will surely influence food resources distribution and availability. For fruits, in particular, this heterogeneity of environments within the Mediterranean region will affect not only the distribution of fruit producing plants, but also the fruit ripening season among populations of the same species. Beside its importance as food resources for mesocarnivores, in some areas the presence of fruit patches can also shape carnivores spatial structure (e.g., olive groves and Eurasian badgers – Rosalino et al., 2004). Therefore understanding the importance of fruit in mesocarnivore diets, the variability of consumed fruit, and its spatial variation is of great value in conservation planning. Due to this significant need to gather data on fruits importance to Mediterranean carnivores, Rosalino and Santos-Reis (2009) reviewed 65 studies focused on fruit consumption by 10 predator species (red fox Vulpes vulpes, weasel Mustela nivalis, stoat Mustela erminea, polecat Mustela putorius, stone marten Martes foina, pine marten, Eurasian badger, common genet, Egyptian mongoose and wildcat Felis silvestris) in areas around the Mediterranean basin (confined by the Euro-Siberian at the north, Sahelian at south, Irano-Turanian/Arabian at east and Macaronesia/Atlantic at west Blondel and Aronson, 1999; Figure 1). They identified a total of 79 different fruits (58 to species level) in the diet of 8 carnivores; no fruits were identified in the diets of stoats or wildcats (Table 1).

Figure 1. Area considered in Rosalino and Santos-Reis (2009) study

Although most of the mesopredators included in the Rosalino and Santos-Reis (2009) survey had a broad geographical distribution within the Mediterranean, with the exception of genets and mongooses (Mitchell-Jones et al., 1999), only four consumed more than 30 fruit species (red fox, stone marten, Eurasian badger and common genet).

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Taxaceae Pinaceae Cupressaceae

Berberidaceae Betulaceae Fagaceae

Amaranthaceae Cannabaceae Moraceae Ericaceae

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Ebenaceae Rosaceae

Taxus baccata Pinus pinea Cupressus sp. Juniperus macrocarpa J. oxycedrus J. phoenicea Berberis vulgaris Corylus avellana Castanea sativa Quercus suber Quercus sp. Amaranthus sp. Celtis australis Ficus carica Arbutus unedo Arctostaphylos uva-ursi Vaccinium myrtillus Diospyros sp. Amelanchier ovalis Crataegus monogyna Cydonia oblonga Eriobotrya japonica Malus domestica M. sylvatica M. communis Malus sp. Prunus avium P. cerasifera P. spinosa Prunus sp. Pyrus bourgaeana P. communis Pyrus sp. Rosa canina Rosa sp. Rubus sp. Sorbus aria S. aucuparia Sorbus sp.

● ●

Egyptian Mongoose

Genet

Badger

Beech marten

Stone marten

Polecat

Fruit species

Weasel

Red fox

Table 1. List of fruits described in the different reviewed carnivores diet studies from the Mediterranean region (adapted from Rosalino et al. 2009)

● ●

● ● ● ● ● ●

● ● ●

●

● ● ●

●

● ● ●

● ●

● ● ●

● ● ● ●

● ● ●

● ●

● ●

● ●

●

● ●

● ● ●

●

●

● ●

● ● ●

● ● ●

● ●

●

● ● ●

● ● ● ● ● ● ● ● ● ●

● ●

●

● ● ● ● ● ● ● ●

●

● ● ●

● ● ●

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●

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Luís M. Rosalino and Margarida Santos-Reis

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Grossulariaceae Fabaceae

Myrtaceae Cornaceae Rhamnaceae Vitaceae

Geraniaceae Anacardiaceae Rutaceae Juglandaceae Oleaceae

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Solanaceae

Rubiaceae Myoporaceae Plantaginaceae Adoxaceae Asteraceae Araceae Arecaceae Poaceae

Ribes rubrum Acacia longifolia Astragalus sp. Ceratonia siliqua Lathyrus sp. Medicago sp. Vicia sativa Myrtus communis Cornus mas Rhamnus alaternus Vitis labrusca V. vinifera Vitis sp. Geranium sp. Pistacia lentiscus Citrus sp. Cneorum tricoccon Juglans regia Ligustrum sp. Olea europaea Hyoscyamus niger Hyoscyamus sp. Solanum lycopersicum S. nigrum Rubia peregrina Myoporum acuminatum Plantago major Sambucus nigra Viburnum sp. Helianthus annuus Arum italicum Chamaerops humilis Phoenix dactylifera Avena sativa Hordeum vulgare Setaria italica Triticum aestivum Triticum sp. Zea mays Unidentified Poaceae

● ● ● ● ● ● ● ●

●

● ●

● ● ● ●

● ●

●

● ●

● ● ● ●

● ● ●

● ●

●

●

● ● ● ●

● ● ●

● ●

● ●

● ● ●

● ● ● ● ●

● ● ●

● ● ●

Middle-Sized Carnivores in Agricultural Landscapes, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

● ● ● ●

● ●

Egyptian Mongoose

Genet

Badger

Beech marten

Stone marten

Polecat

Fruit species

Weasel

Red fox

Table 1. (Continued)

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Fruits and Mesocarnivores in Mediterranean Europe

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Since fruit consumption involves foraging behavior and fruit ripening is highly variable (with some fruits, such as olives, mostly available in autumn, and other like pears in spring), fruit consumption is more diversified in food generalist species such as the red fox and stone marten (Serafini and Lovari, 1993), the Eurasian badgers (Rosalino et al., 2005), or the common genet (Rosalino and Santos-Reis, 2002). Moreover, Rosalino et al. (2009) corroborated what was previously stated by other studies regarding the relatively high importance of fruits to this predators in the Mediterranean, since the average frequency of consumption of fruits in the reviewed studies was higher than 30%, reaching values as high as 46% (Eurasian badgers). Rosalino and Santos-Reis (2009) detected that pine marten consumed a narrower assemblage of Mediterranean fruits. Although Mediterranean carnivores consume dry and fleshy fruits, more than 60% of all eaten fruits are fleshy (Rosalino and Santos-Reis, 2009). This preference appears to be related to the strategies of plants to enhance in seed dispersion. As Herrera (1989) mentioned, most plants whose seeds were generally dispersed by carnivores produce pulp-rich (high content of fibre and low proteins and minerals) fruits, with an attractive scent and sometime with bright colours that, after ripening, generally fell to the ground. This feature increases it conspicuousness and availability, thus biasing fruit selection by mammals. Good examples are blackberries (Rubus sp.), figs (Ficus carica, which is a multiple fruit), grapes (Vitis vinifera) or Phoenician junipers (Juniperus phoenicea) whose consumption has been described for at least five carnivores (Rosalino and Santos-Reis, 2009). Some studies have showed the existence of a latitudinal trend in the feeding behavior of carnivores (Virgós et al., 1999; de Marinis and Massetti, 1995, Zalewski, 2004), usually related with temperature and humidity trends. In those studies, southern populations were often more likely generalist feeders, focusing more on feeding upon non-animal items (such as fruits), while their northern counterparts preyed mostly on animals. However, within the Mediterranean region, Rosalino and Santos-Reis (2009) did not detect any latitudinal trend in fruit consumption. Nevertheless, a significant cline was identified regarding the frequency of fruit consumption along the Mediterranean longitudinal variation, with higher values towards eastern population (see Figure 2). Since the Iberian Peninsula has usually more wet days than the eastern region of the Mediterranean, such as Greece (Goodess, 2000), it‘s possible that the higher consumption of fruits in those regions might be related with higher temperatures and lower humidity that could constrains the abundance of other prey, such as rodents (since plant cover and productivity respond to precipitation, and influence rodents dynamics and density – Ernest et al., 2000). These factors may not be the only variables involved in the establishment of the pattern, since such a complex system (involving abiotic factors such as rainfall and elevaton, as well as plant dynamics, prey and predator distribution and dynamics) is difficult to untangle, especially in the absence of contemporary environmental and systematic data on fruit distribution and abundance. Nevertheless, it seems plausible to consider that fruits consumption will be highly influenced by diverse features that vary regionally across the Mediterranean area, especially fruit characteristics (e.g., pulp content), availability of wild and cultivated fruits (especially those commercially produced - e.g., oranges, olives, grapes) and availability of other food resources (Rosalino and Santos-Reis, 2009).

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F.O. (%) 100 75 50 25 0 -10

-5

0

5

Longitude

10

15

20

25

Figure 2. Variation in the fruit consumption (assessed as Frequency of occurrence - FO) by mesocarnivores along the Mediterranean longitude variation (negative values on the x-axis represent west and positive represent east - adapted from Rosalino et al. 2009).

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5. CARNIVORES AS EFFECTIVE FRUIT DISPERSERS IN PORTUGAL The relation between plants and seed dispersers is anchored on fruit consumption and understanding the nature of these animal-plant interactions is crucial for understanding ecosystem regeneration and evolution (Corlett, 1995). This association is based in the capability of plants to provide food to dispersers, as nutritious accessory structures closely associated with the seeds, as well as to capture dispersers attention by using attractive colours (e.g., for birds, monkeys or others diurnal vertebrates – Corlett, 1998) or scents (e.g., bats – Bianconi et al., 2007). Species involved in this biotic relation vary according with fruit characteristics, ecology of the dispersers, and species distribution ranges (Herrera, 1987). Because dispersers may deposit viable seeds into sites where seeds can germinate and seedlings can establish themselves, they can help shape plant communities diversity and genetic structure, expand population distribution ranges, influence demography (Jordano et al., 2007), and have an active role on the colonization of vacant habitats by plants (Nathan et al., 2008). Many animal groups have been identified as disperser, including invertebrates such as earthworms (Brown et al., 1994) or ants (Aronne and Wilcock, 1994). The type of disperser will determine the dispersion route and scale. For example, while medium /large size mammals and large birds might transport seeds over long distances and diverse landscapes, small and medium /size birds or invertebrates often deposit seeds in the vicinity of the seed producing plant (Jordano et al., 2007). In Mediterranean Europe, vertebrates assume a higher importance, especially birds and mammalian carnivores (Herrera 1987), although reptiles might also play a role in seed dispersion (Olesen and Valido, 2003). The importance of vertebrate in the dispersion of Mediterranean plants is highlighted by the fact that 30-65% of fruit producing plants in Southern Europe are dispersed by those animals (Herrera, 2001). In Iberia, several carnivores

Middle-Sized Carnivores in Agricultural Landscapes, Nova Science Publishers, Incorporated, 2011. ProQuest Ebook Central,

Fruits and Mesocarnivores in Mediterranean Europe

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are good candidates for being an effective disperser since fruits are an important resource for those species, at least seasonally (e.g., Eurasian badgers, common genets, red-foxes or stone martens - Rosalino and Santos-Reis, 2002; Rosalino et al., 2005; Santos et al., 2007). Some authors have used the criterion of fruit consumption to question the ability of carnivores to function has seed dispersers. For example, Herrera (1989; 1995) used the seeds found in scats (together with a visual inspection to determine if the seeds were destroyed) to determine that about 40% of the fleshy-fruited plants occurring in the region were dispersed by three carnivores species: red-fox, Eurasian badger, and stone marten. However, although various plant species are consumed by carnivores and many seeds are not destroyed during digestion, the successful dispersion of a seed involves more than just these two factors. An effective seed disperser must allow the ingested seed to be viable. For such a frugivorous species the seed‘s passage through its gut should enhance germination (i.e., increase the rate of germination), or in the worse case have a neutral effect on the seed's viability (Traveset, 1998). The seed germination improvement after passing through the gut may be related to the separation of the fruit from the fleshy pulp, the moistening and fertilizer effect of the non-digested material in scats and the softening and scarification of the seed coat through mastication or action of acids and enzymes in saliva and stomach (Razanamandranto et al., 2004). Table 2. Percentage of germinated seeds collected in faeces (% faeces) and seeds isolated in wild fruits (% fruits) (* indicates statistical differences - adapted from Rosalino et al., 2010)

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Plant species

% fruits

Martes foina

Meles meles

% scats Herpestes ichneumon

Genetta genetta

Vulpes vulpes

Arbutus unedo

9.48

1.45*

0.00*

0.00*

0.00*

0.00*

Ficus carica Olea europaea