Ecological assessment of the red deer population in the Lousã Mountain

Texto

(1)Universidade de Aveiro Departamento de Biologia 2013. Joana Alexandra da Silva Alves. Avaliação Eco-cinegética da População de Veado da Serra da Lousã Ecological Assessment of the Red Deer Population in the Lousã Mountain.

(2) Universidade de Aveiro Departamento de Biologia 2013. Joana Alexandra da Silva Alves. Avaliação Eco-cinegética da População de Veado da Serra da Lousã Ecological Assessment of the Red Deer Population in the Lousã Mountain Dissertação apresentada à Universidade de Aveiro para cumprimento dos requisitos necessários à obtenção do grau de Doutor em Biologia, realizada sob a orientação científica do Prof. Doutor Carlos M. M. S. Fonseca, Professor Auxiliar com Agregação do Departamento de Biologia da Universidade de Aveiro e co-orientação do Prof. Doutor Amadeu M. V. M. Soares, Professor Catedrático do Departamento de Biologia da Universidade de Aveiro.. Apoio financeiro da FCT e do FSE no âmbito do III Quadro Comunitário de Apoio (Bolsa de Doutoramento SFRH/BD/22599/2005)..

(3) À minha família, em particular aos meus avós, pela sua fé inabalável em mim e nas minhas capacidades… Ao Toni, pela esperança e amor com que sempre me apoiou….

(4) o júri presidente. Prof. Doutor José Carlos da Silva Neves professor catedrático da Universidade de Aveiro. Prof. Doutor Roderick Putman professor emérito da Universidade Metropolitana de Manchester, Reino Unido. Prof. Doutor Amadeu Mortágua Velho da Maia Soares professor catedrático Universidade de Aveiro. Prof. Doutor Francisco Manuel Cardoso de Castro Rego professor associado com agregação da Universidade Técnica de Lisboa, Instituto Superior de Agronomia, Centro de Ecologia Aplicada. Prof. Doutor António José Arsénia Nogueira professor associado com agregação da Universidade de Aveiro. Prof. Doutor Carlos Manuel Martins Santos Fonseca professor auxiliar com agregação da Universidade de Aveiro. Prof. Doutor José Vítor Sousa Vingada professor auxiliar da Universidade do Minho. Doutor Miguel Nuno Sacramento Monteiro Bugalho investigador auxiliar da Universidade Técnica de Lisboa, Instituto Superior de Agronomia, Centro de Ecologia Aplicada. Doutora Rita Maria Tinoco da Silva Torres investigadora de pós-doutoramento do Departamento de Biologia da Universidade de Aveiro.

(5) agradecimentos. Chegada a hora dos agradecimentos… haverá certamente muito que ficará por dizer! Mas vou tentar… pelo menos tentar… fazer jus ao que foi esta caminhada! Quero começar por agradecer ao Prof. Doutor Amadeu Soares e em particular ao Prof. Doutor Carlos Fonseca, meus orientadores, o convite que me fizeram para vir para esta Instituição e aqui realizar o meu doutoramento… este foi sem dúvida o começo de uma grande aventura! O meu obrigada vai também para todas as instituições que permitiram que este projeto chegasse a bom porto: à FCT pelo suporte financeiro, à AFN, e em particular ao Eng. Jorge Cancela, pelo apoio financeiro e logístico, às entidades gestoras das zonas de caça presentes na Serra da Lousã por me terem recebido, pelos momentos inesquecíveis e pelos ensinamentos que em muito excedem o aspeto profissional… hoje sinto por vós verdadeira amizade! Este fantástico projeto não teria sido o mesmo se não fosse a colaboração de pessoas que aos poucos foram tornando o meu dia-a-dia mais fácil, mais produtivo e mesmo mais divertido… agradeço por isso ao Rui Figueiredo, ao Manuel Bento e a todos os que de algum modo tiveram um papel ativo neste trabalho. Ao Disco, por me levar sempre onde foi preciso, permitindo-me chegar aqui… Ao Mestre António Silva, o mais brilhante e competente jovem investigador que alguma vez conheci… obrigada por tudo o que fizeste desde o primeiro dia, por me fazeres acreditar em mim, pelo incentivo e acima de tudo por me teres permitido aprender contigo… esta Tese é também tua! Aos amigos que comigo percorreram este longo caminho… que me abraçaram nos momentos difíceis, que comigo riram, que tornaram esta viagem inesquecível… este é o meu obrigada ao Nuno, à Inês, ao Eduardo, ao Pascoal, à Constança, à Fernanda, ao Vítor, ao Filipe, ao Pi e a todos os que fazem parte da minha vida! Quero ainda agradecer à minha Mii (e ao telefone mágico…), sem ti esta batalha seria muito menos sucedida! Quero ainda agradecer o apoio de um grupo muito especial de amigos, pessoas que me apoiaram e me incentivaram a continuar numa luta que parecia interminável… não há palavras, mas este é o meu agradecimento ao Tiago, à Cátia, ao Henrique, à Sónia, à Dalila, à Sara, à Carla, à Cristina, à Cláudia, ao Filipe Martinho e à Cláudia! Deixo ainda um Obrigada por tudo… ao Prof. Paulo Sousa e ao Prof. Jaime Ramos! Às crianças da minha vida, a Filipa, o Francisco e a Inês, por terem sido o meu escape, a minha ligação à realidade, por me recordarem que a vida é mais do que uma tese… À minha família… aos presentes e aos ausentes, obrigada por tudo! Aos meus pais, pelo apoio incondicional e pelas palavras de incentivo… obrigada por serem quem são e por me ajudarem a tornar-me a pessoa que sou! À minha irmã, amiga de todas as horas… espero poder ser para ti tudo o que representas para mim! Ao meu namorado, por ter estado sempre presente… Por último, e como disse a minha mãe: a mim, por ter sempre tentado fazer melhor, por não ter sucumbido às dúvidas e por ter lutado sempre… custasse o que custasse! Sejam Felizes….

(6) palavras-chave. Cervus elaphus, condição física, estimativa de densidade, imunocompetência, segregação sexual, Serra da Lousã, uso de habitat.. resumo. Durante o último século, as populações de cervídeos têm aumentado substancialmente um pouco por toda a Europa. O veado Cervus elaphus não foi exceção, apresentando atualmente uma ampla distribuição geográfica. Após a quase extinção de todas as populações selvagens desta espécie em Portugal, o número de efetivos de veado e a área de distribuição da espécie têm aumentado significativamente nas últimas décadas. Esta proliferação deveu-se fundamentalmente aos processos de reintrodução implementados em algumas zonas do país, como foi o caso da Serra da Lousã. Dez anos após a reintrodução, a expansão geográfica e demográfica é já uma realidade e a espécie assume um papel relevante tanto do ponto de vista cinegético como turístico. Com o aumento das populações de ungulados, surge a necessidade de aumentar o conhecimento existente sobre estas espécies, particularmente ao nível das estimativas populacionais, do uso e seleção de habitat, do comportamento e da fisiologia das populações. Neste sentido, um dos objetivos foi comparar quatro métodos de contagem de excrementos em termos de estimativa populacional, aplicabilidade e eficiência. Face aos resultados obtidos, o método de transectos lineares com distance sampling revelou-se o mais eficiente, providenciando uma boa precisão num menor tempo. Relativamente ao uso e seleção do habitat, recorrendo a transectos de contagem de excrementos e a observações diretas, verificou-se que o veado seleciona positivamente habitats que lhe proporcionam simultaneamente alimento e refúgio, como é o caso das áreas de ecótono. A preferência por áreas de ecótono evidencia um comportamento adaptativo de maximização de energia, no qual os animais optam por procurar refúgio na proximidade das áreas de alimentação de forma a minimizarem os dispêndios energéticos. Em termos de dinâmica populacional, verificou-se uma relação entre a densidade de veado e o tamanho dos seus grupos, bem como o efeito nos mesmos do coberto vegetal. Contudo, dado o seu alto dimorfismo sexual em termos de tamanho corporal, as preferências de habitat podem também ser em parte responsáveis pela segregação sexual fora da época de reprodução. Face aos resultados obtidos através do método de observação direta de animais, a segregação sexual surge como consequência da interação entre fatores sociais e de habitat, que resultam de diferentes estratégias reprodutivas por parte de ambos os sexos. As diferentes estratégias reprodutivas são também notórias em termos de condição física e imunológica, com os machos a aumentarem a sua condição durante as épocas de primavera e verão de forma a melhorarem a sua performance durante o cio, e as fêmeas a viverem grande parte do ano no limiar da condição física devido ao seu investimento na sobrevivência da sua descendência. Globalmente, o veado apresenta uma boa adaptação à Serra da Lousã e representa um importante recurso natural para esta região. Contudo, para que a proliferação desta população selvagem continue, é necessária a implementação de medidas de gestão adequadas a este ecossistema mediterrâneo..

(7)

(8) keywords. Cervus elaphus, density estimates, habitat use, immunocompetence, Lousã Mountain, physical condition, sexual segregation.. abstract. Over the last century, the populations of deer species have increased substantially throughout Europe. Red deer Cervus elaphus was not an exception to this trend, having now a wide distribution. After having been close to extinction, the red deer populations in Portugal are increasing in both abundance and geographical range. This expansion results fundamentally from reintroduction processes, which was the case of the Lousã Mountain. Ten years after, geographic and demographic proliferations are now a reality, and this species assumes an important role for hunting and tourism. With the increase of wild populations of ungulates, many efforts have been made to improve the knowledge about this species, particularly in terms of populations’ estimates, habitat use, behaviour and physiology of the populations. As so, one of the goals was to compare four pellet count methods in terms of density estimates, applicability and efficiency. The results indicated that the standing crop line transects counts, was the most efficient method, providing great precision in less time. Regarding habitat use and selection, using pellet group counts and direct observations was verified that the red deer selects positively habitats that provide simultaneously food and cover, like the ecotone areas. Such preference highlights an adaptive behaviour for increasing fitness, in which the animals choose to take refuge near to food areas as a way of minimize the energetic losses. In terms of population dynamics, the results showed a positive relation between red deer density and group size, and also an effect of land cover in the size, numbers and composition of groups. However, due to the high body size dimorphism, the habitat preferences may be in part responsible by the sexual segregation outside the rut season. The results obtained by direct observation of animals showed that the sexual segregation arises because of the interaction between social and habitat factors, which result from different reproductive strategies of the sexes. The reproductive strategies were also noticed in the analysis of the physical and immunological conditions. Males increased their condition during spring and summer as a way of improve the performance during the rut time, while females live at the threshold of physical condition throughout great part of the year, due to its investment on the survival of their offspring. Globally, the red deer seems well adapted to Lousã Mountain and represents an important resource for this region. Although, for the proliferation of this wild population continues, it is necessary to implement management practices proper to this Mediterranean ecosystem..

(9) Contents . vii . . . Contents. . List of illustrations ............................................................................................................................ xiii List of tables .................................................................................................................................... xvii Status of the manuscripts ................................................................................................................ xix Chapter 1 ‐ General introduction ....................................................................................................... 1 1.1. Red deer Cervus elaphus in Europe ......................................................................................... 3 1.2. Red deer: ecology and behaviour ........................................................................................... 4 1.3. Red deer management ............................................................................................................ 6 1.4. Aims ......................................................................................................................................... 7 1.5. Thesis framework .................................................................................................................... 9 1.6. References ............................................................................................................................. 11 Chapter 2 ‐ Study area ...................................................................................................................... 17 2.1. Study Area ............................................................................................................................. 19 2.1.1 Location ........................................................................................................................... 19 2.1.2. Bio‐physical characterization ......................................................................................... 20 2.1.2.1. Climate ........................................................................................................................ 20 2.1.2.2. Topography ................................................................................................................. 21 2.1.2.3. Land Cover ................................................................................................................... 22 2.1.2.4. Fauna ........................................................................................................................... 24 2.1.3. Natura 2000 Site – PTCON0060 Serra da Lousã ............................................................. 25 2.1.4. Deer management and hunting areas ........................................................................... 26 2.2. References ............................................................................................................................. 27 Chapter 3 ‐ Red deer distribution and population parameters ....................................................... 29 .

(10) viii . Contents . 3.1 Abstract .................................................................................................................................. 31 3.2 Introduction ............................................................................................................................ 31 3.3 Methods ................................................................................................................................. 32 3.3.1 Study area ........................................................................................................................ 32 3.3.2 Data collection ................................................................................................................. 32 3.3.3 Data analysis .................................................................................................................... 33 3.4 Results and Discussion ........................................................................................................... 34 3.4.1 Distribution area .............................................................................................................. 34 3.4.2 Red deer density and demographic parameters ............................................................. 36 3.5 References .............................................................................................................................. 38 Chapter 4 ‐ Pellet count methods to estimate red deer: precision, potential accuracy and efficiency .......................................................................................................................................... 41 4.1. Abstract ................................................................................................................................. 43 4.2. Introduction ........................................................................................................................... 43 4.3. Methods ................................................................................................................................ 46 4.3.1. Study area ....................................................................................................................... 46 4.3.2. Field work ....................................................................................................................... 46 4.3.3. Simulations ..................................................................................................................... 47 4.3.4. Data analysis ................................................................................................................... 48 4.4. Results ................................................................................................................................... 50 4.4.1. Field data ........................................................................................................................ 50 4.4.2. Simulated data ............................................................................................................... 52 4.5. Discussion .............................................................................................................................. 56 4.6. References ............................................................................................................................. 60 Chapter 5 ‐ Spatial and temporal habitat use and selection by red deer in a Mediterranean area 65 5.1. Abstract ................................................................................................................................. 67 5.2. Introduction ........................................................................................................................... 67 .

(11) Contents . ix . 5.3. Methods ................................................................................................................................ 70 5.3.1. Study area ...................................................................................................................... 70 5.3.2. Data collection ............................................................................................................... 70 5.3.3. Habitat variables ............................................................................................................ 71 5.3.4. Data analysis ................................................................................................................... 72 5.4. Results ................................................................................................................................... 74 5.4.1. Habitat use ..................................................................................................................... 74 5.4.2. Habitat selection ............................................................................................................ 77 5.5. Discussion .............................................................................................................................. 79 5.6. References ............................................................................................................................. 83 Chapter 6 ‐ Group dynamics of red deer at the rutting season: the influence of local population density and land cover ..................................................................................................................... 89 6.1. Abstract ................................................................................................................................. 91 6.2. Introduction........................................................................................................................... 91 6.3. Methods ................................................................................................................................ 93 6.3.1. Study area ...................................................................................................................... 93 6.3.2. Red deer population ....................................................................................................... 94 6.3.3. Data collection ............................................................................................................... 94 6.3.4. Data analysis ................................................................................................................... 95 6.4. Results ................................................................................................................................... 97 6.4.1. Group dynamics ............................................................................................................. 97 6.4.2. Effect of local population density .................................................................................. 99 6.5. Discussion ............................................................................................................................ 102 6.5.1. Group dynamics ........................................................................................................... 102 6.5.2. Effect of local population density ................................................................................ 103 6.5.3. Effect of land cover ...................................................................................................... 104 6.6. Conclusions.......................................................................................................................... 105 .

(12) x . Contents . 6.7. References ........................................................................................................................... 106 Chapter 7 ‐ Sexual segregation in red deer: is social behaviour more important than habitat preferences? ................................................................................................................................... 111 7.1. Abstract ............................................................................................................................... 113 7.2. Introduction ......................................................................................................................... 113 7.3. Methods .............................................................................................................................. 118 7.3.1. Study area ..................................................................................................................... 118 7.3.2. Red deer population ..................................................................................................... 118 7.3.3. Data collection .............................................................................................................. 119 7.3.4. Data analysis ................................................................................................................. 120 7.4. Results ................................................................................................................................. 122 7.4.1. Patterns of Sexual Segregation .................................................................................... 122 7.4.2. Effect of Presence of Young ......................................................................................... 125 7.4.3. Age‐Related Segregation .............................................................................................. 127 7.5. Discussion ............................................................................................................................ 128 7.5.1. Patterns of Sexual Segregation .................................................................................... 128 7.5.2. Evaluating sexual segregation hypotheses ................................................................... 129 7.5.3. Age‐related segregation ............................................................................................... 131 7.6. Conclusions .......................................................................................................................... 132 7.7. References ........................................................................................................................... 132 Chapter 8 ‐ Physical condition and immunocompetence of red deer in a Mediterranean ecosystem ........................................................................................................................................................ 139 8.1. Abstract ............................................................................................................................... 141 8.2. Introduction ......................................................................................................................... 142 8.3. Methods .............................................................................................................................. 145 8.3.1. Study area ..................................................................................................................... 145 8.3.2. Data collection and analysis ......................................................................................... 145 .

(13) Contents . xi . 8.3.3. Data analysis ................................................................................................................. 147 8.4. Results ................................................................................................................................. 148 8.4.1. Physical condition ......................................................................................................... 148 8.4.2. Immunocompetence .................................................................................................... 150 8.4.3. Biological relationships ................................................................................................ 150 8.5. Discussion ............................................................................................................................ 152 8.6. References ........................................................................................................................... 156 Chapter 9 ‐ General discussion and conclusions ............................................................................ 163 9.1. Estimating deer populations ............................................................................................... 165 9.2. Red deer at Lousã Mountain: habitat use, population dynamics and behaviour ............... 166 9.3. Evaluation of the red deer reintroduction in the Lousã Mountain: geographical range, density and physiological condition ........................................................................................... 167 9.4. Management guidelines and implications .......................................................................... 168 9.5. Future research ................................................................................................................... 169 9.6. References ........................................................................................................................... 170 .

(14)

(15) List of illustrations Fig. 1.1. Map of current distribution of red deer across Europe by a 50km2 grid (adapted from ©Societas Europaea Mammalogica). ................................................................................................ 3 Fig. 2.1. Location of the study area, the Lousã Mountain, in Portugal. ........................................... 19 Fig. 2.2. Climate diagram for the studied period (2005‐2009) in Lousã Mountain (data from SNIRH). The solid line represents the precipitation (mm) and the dashed line the temperature (ᵒC). .......................................................................................................................................................... 20 Fig. 2.3. Topographic map of the study area. .................................................................................. 21 Fig. 2.4. Hydrologic network in the study area. ............................................................................... 22 Fig. 2.5. Land cover of the study area, based on aerial photographs interpretation. ..................... 23 Fig. 2.6. Natura 2000 Site – PTCON0060 Serra da Lousã. ................................................................ 25 Fig. 2.7. Map of hunting areas in the study area. ............................................................................ 26 Fig. 3.1. Limits of the distribution area of red deer in the study area and its natural and artificial barriers. ............................................................................................................................................ 34 Fig. 3.2. Map of current distribution of red deer in Lousã Mountain and surrounding areas......... 35 Fig. 3.3. Deer density estimates along the studied years. The dashed line represents the global deer density for the whole period (2005‐2009) and the error bars are 95% confidence intervals. 36 Fig. 4.1. Deer density estimates from faecal standing crop plot counts (FSCP), faecal clearance plot counts (FAR), faecal standing crop strip transect counts (ST) and faecal standing crop line transect counts (LT) for three habitats of Lousã. The error bars are bootstrapped 95% confidence intervals. .......................................................................................................................................................... 51 Fig. 4.2. Deer density estimates from a) faecal standing crop plot counts, b) faecal clearance plot counts, c) faecal standing crop strip transect counts and d) faecal standing crop line transect counts for a simulated deer population at 5.5 deer km‐2 (dashed line) for different survey efforts. The error bars are bootstrapped 95% confidence intervals. ........................................................... 53 Fig. 4.3. Mean coefficient of variation (CV) for a) faecal standing crop plot counts, b) faecal clearance plot counts, c) faecal standing crop strip transect counts and d) faecal standing crop line transect counts for a simulated deer population at 5.5 deer km‐2 for different survey efforts. The error bars are bootstrapped 95% confidence intervals. .................................................................. 54 .

(16) xiv . List of illustrations . Fig. 4.4. Mean potential accuracy (SMΛE) for a) faecal standing crop plot counts, b) faecal clearance plot counts, c) faecal standing crop strip transect counts and d) faecal standing crop line transect counts for a simulated deer population at 5.5 deer km‐2 for different survey efforts. The error bars are bootstrapped 95% confidence intervals. .................................................................. 55 Fig. 4.5. Relative net precision (RNP) for faecal standing crop line transect counts (LT), faecal standing crop strip transect counts (ST), faecal standing crop plot counts (FSCP) and faecal clearance plot counts (FAR) for a simulated deer population at 5.5 deer km‐2 for different survey efforts. The higher the RNP (eqn. 4.6) the higher the efficiency of the pellet group count method. .......................................................................................................................................................... 56 Fig. 5.1. Proportion of availability and use of land cover units by males and females of red deer. 76 Fig. 5.2. Selection ratios for a) land cover, b) distance to water, c) distance to active roads, d) distance to closed areas, and e) distance to open areas by red deer during birth time, rut season and winter dispersal. Land covers was divided into coniferous stage I (Conif I), coniferous stage II (Conif II), coniferous stage III (Conif III), mixed forest (Mixed), shrubland (Shrub) and grasslands (Grass).Distances were pooled into 25 m intervals. If selection ratio is significantly higher than 1 (dashed line) it indicates preference, if significantly lower than 1 it indicates avoidance. The error bars are Bonferroni‐adjusted 95% confidence intervals. *P <0.05, **P <0.01, ***P <0.001. ......... 78 Fig. 6.1. (a) Proportion of red deer groups observed in each size category at pre‐rut, rut and post‐ rut. b) Proportion of red deer individuals observed in each size category at pre‐rut, rut and post‐ rut. .................................................................................................................................................... 97 Fig. 6.2. Occurrence of group typologies of red deer observed in the sampling periods of the reproductive season. ........................................................................................................................ 99 Fig. 6.3. (a) Mean group size and (b) number of groups as a function of local population density of red deer during the rutting season. ............................................................................................... 100 Fig. 6.4. Histograms of perpendicular distances and fitted detection functions for (a) shrublands (hazard‐rate key without adjustment terms, N=280) and (b) forests (hazard‐rate key with simple polynomial adjustment, N=545), using observations of red deer groups during the reproductive season. ............................................................................................................................................ 101 Fig. 6.5. Occurrence of group typologies of red deer observed in the land cover units during the reproductive season. ...................................................................................................................... 102 Fig. 7.1. (a) Seasonal patterns of sexual segregation and aggregation for adult males and females of red deer (NWinter=37; NSpring=28; NSummer=104; NRut=697; NAutumn=58). (b) Patterns of sexual segregation and aggregation for adult males and females of red deer at prerut (N16‐31 Aug=36; N1‐15 . .

(17) List of illustrations . xv . Sep=63), rut (N16‐30 Sep=316; N1‐15 Oct=222; N16‐31 Oct=96) and postrut (N1‐15 Nov=41; N16‐30 Nov=17). The . SSAS indicates significant sexual segregation or aggregation when the observed value (black point) falls above or below the SSAS expected interval (shaded area), respectively. .............................. 123 Fig. 7.2. (a) Determination of the best scale to measure spatial segregation (N=104). The dashed grey line is the observed spatial segregation, while the continuous black line is the simulated segregation. The best scale is found when the simulated value falls into the SSAS expected interval (shaded area). (b) Seasonal patterns of spatial segregation (1 km2 scale) for adult males and females of red deer (NWinter=37; NSpring=28; NSummer=104; NRut=697; NAutumn=58). (c) Patterns of sexual segregation and aggregation in red deer for adult males and females with young at prerut (N16‐31 Aug=26; N1‐15 Sep=49), rut (N16‐30 Sep=271; N1‐15 Oct=207; N16‐31 Oct=84) and postrut (N1‐15 Nov=35; N16‐30 Nov=16). (d) Patterns of sexual segregation and aggregation in red deer for adult males and females without young at prerut (N16‐31 Aug=21; N1‐15 Sep=40), rut (N16‐30 Sep=246; N1‐15 Oct=143; N16‐31 Oct=59) and postrut (N1‐15 Nov=25; N16‐30 Nov=8). The SSAS indicates significant sexual segregation or . aggregation when the observed value (black point) falls above or below the SSAS expected interval (shaded area), respectively. .............................................................................................. 125 Fig. 8.1. Relationship between bone marrow fat (BMF) and kidney fat index (KFI). The dashed lines represent the breakpoints of the patterns of mobilization from both fat depots. ....................... 146 Fig. 8.2. Annual patterns of physical condition for red deer males and females. a) kidney fat index (KFI), b) bone marrow fat (BMF) and c) a combination of KFI and BMF (Conindex). The values (squares and circles) represent the estimated marginal means using body length as covariate and the error bars are bootstrapped 95% confidence intervals. .......................................................... 149 Fig. 8.3. Annual patterns of immunocompetence (spleen weight) for red deer males and females. The values (squares and circles) represent the estimated marginal means using body length as covariate and the error bars are bootstrapped 95% confidence intervals. ................................... 150 Fig. 8.4. Annual patterns of testosterone levels (combined testis weight) for red deer males. The values (squares) represent the estimated marginal means using body length as covariate and the error bars are bootstrapped 95% confidence intervals. ................................................................ 151 .

(18)

(19) . . List of tables. Table 3.1. Population structure, sex ratio and reproductive rate of the observed red deer population of the Lousã Mountain between 2005 and 2009. ......................................................... 37 Table 4.1. Number of sampling units and effort used in faecal standing crop plot counts (FSCP), faecal clearance plot counts (FAR), faecal standing crop strip transect counts (ST) and faecal standing crop line transect counts (LT) per habitat type. ................................................................ 47 Table 4.2. Red deer estimates in Lousã from faecal standing crop plot counts (FSCP), faecal clearance plot counts (FAR), faecal standing crop strip transect counts (ST) and faecal standing crop line transect counts (LT). .......................................................................................................... 51 Table 5.1. Best models of habitat use by red deer, using pellet group counts and direct observations, and best models of habitat selection by red deer using direct observations. .......... 74 Table 5.2. Regression coefficients (β) for the best model of habitat use by red deer, using pellet group counts and direct observations. ............................................................................................ 75 Table 5.3. Proportions of availability and use of different environmental variables by males and females of red deer. ......................................................................................................................... 76 Table 5.4. Seasonal selection ratios for topographic variables (altitude, aspect and slope) by red deer in Lousã, Central Portugal. ....................................................................................................... 77 Table 5.5. Regression coefficients (β) for the best model of habitat selection by red deer, using direct observations. .......................................................................................................................... 79 Table 6.1. Group size statistics of red deer among the sampling periods of the reproductive season. ............................................................................................................................................. 98 Table 6.2. Estimation of the parameters of red deer groups for shrubland and forest areas. ....... 98 Table 7.1. Hypotheses and predictions to explain sexual segregation .......................................... 117 Table 7.2. Percentage of red deer sexual segregation explained by habitat (land cover and environmental variables) at different seasons .............................................................................. 124 Table 7.3. Seasonal patterns of sexual segregation and aggregation for males and females with and without young of red deer and respective social and habitat components (in percentage). 126 Table 7.4. Seasonal patterns of age‐related segregation and aggregation for red deer and respective social and habitat components (in percentage). .......................................................... 127 .

(20) xviii . List of illustrations . Table 8.1. Partial correlations between physical condition index (KFI, BMF and Conindex), immunocompetence (Spleen weight) and testosterone (Combined testis weight), controlled for body length..................................................................................................................................... 151 . .

(21) . Status of the manuscripts Chapter 4 Pellet count methods to estimate red deer: precision, potential accuracy and efficiency Joana Alves*, António Alves da Silva, Amadeu M.V.M. Soares & Carlos Fonseca . In press ‐ Mammalian Biology . Chapter 5 Spatial and temporal habitat use and selection by red deer in a Mediterranean area Joana Alves*, António Alves da Silva, Amadeu M.V.M. Soares & Carlos Fonseca . In prep. . Chapter 6 Group dynamics of red deer at the rutting season: the influence of local population density and land cover Joana Alves*, António Alves da Silva, Amadeu M.V.M. Soares & Carlos Fonseca . . In prep. Chapter 7 Sexual segregation in red deer: is social behaviour more important than habitat preferences? Joana Alves*, António Alves da Silva, Amadeu M.V.M. Soares & Carlos Fonseca . In press ‐ Animal Behaviour .

(22) xx . Status of the manuscripts . . Chapter 6 Physical condition and immunocompetence of red deer in a Mediterranean ecosystem Joana Alves*, António Alves da Silva, Amadeu M.V.M. Soares & Carlos Fonseca . Paper submitted to Acta Oecologica . .

(23) . . . Chapter 1. General introduction .

(24)

(25) General Introduction . 3 . . 1.1. Red deer Cervus elaphus in Europe Over the last century, the populations of many deer species have substantially increased throughout Europe (e.g. Gill 1990; Apollonio, Andersen & Putman 2011) both in numbers and geographical range, as a consequence of habitat changes (alterations in land cover and agricultural and forestry practices) and cervids management (protection from over hunting, supplementary feeding and reintroduction programmes) (Milner et al. 2006). Red deer Cervus elaphus (Linnaeaus, 1758) was not an exception to this trend and currently has a widely distribution across Europe (Koubek & Zima 2009) (Fig. 1.1). . Fig. 1.1. Map of current distribution of red deer across Europe by a 50km2 grid (adapted from ©Societas Europaea Mammalogica). The evolution of the genus Cervus over the time results in distinct species (C. elaphus, C. nippon, C. eldi and C. canadensis) and several subspecies (Ludt et al. 2004; Pitra et al. 2004). In Europe, the division of Cervus elaphus into different subspecies (e.g. C. e. atlanticus C. e. brauneri, C. e. elaphus, C. e. hippelaphus, C. e. hispanicus, and C. e. scoticus) is widely accepted (see Ludt et al. 2004; Pitra et al. 2004). Nevertheless, such subdivision has been questioned and recent works showed that some of the Cervus elaphus subspecies could not be differentiated by the mtDNA .

(26) 4 . Chapter 1 . cytochrome b sequence data (Ludt et al. 2004). However, the absence of a taxonomic revision of the subspecies of Cervus elaphus so far, the morphological characteristics and the results of Skog et al. (2009), still allow the classification of Portuguese red deer as belonging to C. e. hispanicus. In Portugal, the overexploitation and the destruction/fragmentation of habitat led wild populations of red deer almost to extinction at the end of XIX century. After the sixties, the abandon of agriculture and the reintroduction programmes together with natural dispersion of the species allow the growth in both abundance and geographical range (Salazar 2009). As in many other regions of Portugal, the presence of red deer in the Lousã Mountain is an outcome of a reintroduction process that occurs from 1995 to 1999, with the released of 96 animals (32 males and 64 females). This programme aimed to improve the natural patrimony of this Mediterranean mountainous area, enabling at the same time the viability of hunting and tourism. . 1.2. Red deer: ecology and behaviour The habitat selection by ungulates, like by other animals, reflects their resource requirements (Manly et al. 2002), which can be aggregated into two major components, food and cover (Borkowski 2004). Red deer is considered a species with a high plasticity justified by its ability to utilize a wide variety of habitats, from woodlands at temperate climates (Bobek, Boyce & Kosobucka 1984; Theuerkauf & Rouys 2008) to open areas (shrublands and grasslands) in Mediterranean zones (Soriguer et al. 1994; Garín 2000). In Mediterranean environments, the red deer uses a high variety of habitats, but seems to prefer shrublands and pastures with Quercus trees (“dehesas”) (Carranza et al. 1991; Soriguer et al. 1994; Garín 2000; Lovari et al. 2007; Acevedo et al. 2008). Given the red deer life cycle, where reproduction and births seasons are well temporally established, is expected that its requirements change between seasons, resulting in a temporal effect on habitat selection. Another important aspect is the seasonal dispersion and movements of animals, a fact so often associated with deer (Clutton‐Brock, Guinness & Albon 1982; Bonnet & Klein 1991; Soriguer et al. 1994) and that may constitute a spatial gradient of habitat selection. On the other hand, habitat use by red deer tends to be influenced by behavioural and physiological responses to environmental changes, interspecific competition, predation risk and anthropogenic factors (e.g. Jeppesen 1987; Bartos et al. 2002; Lovari et al. 2007; Barja & Rosellini 2008; Sánchez‐ Prieto et al. 2010). . .

(27) General Introduction . 5 . Classified as an intermediate feeder (Hofmann 1985; Hofmann 1989), the red deer diet is composed by a wide variety of plant species. The feeding behaviour of this cervid switch between grazing (grass and sedges) and browsing (leaves of broadleaf, coniferous and shrubs) according to food availability, using fall fruits and seeds when available (Gebert & Verheyden‐Tixier 2001). Contrarily, to what happens at northern latitudes, the summer at Mediterranean climates appears as the most limiting period of the year and may lead to nutritional constrain caused by the hot and dry weather. During this season, the red deer may use a higher proportion of browse in its diet (e.g. leaves of broadleaf trees and blackberry Rubus spp.). These changes in diet composition result from the senescence of the herbage layer and are more pronounced in areas rich in grassland habitats (Bugalho & Milne 2003). In the Lousã Mountain, the red deer makes use of shrublands (e.g. Erica sp.; Pterospartum tridentatum) as its main food resource during all year. Some Quercus trees are also an important item in red deer diet, mainly in summer (Ferreira 1998). Red deer, as many others’ sexual body size dimorphic mammals, is a polygamous species (Clutton‐ Brock 1989). Their slightly gregarious behaviour is characterized by a sexual segregation outside the reproductive time, with males and females live apart during much of the year. At pre‐rut, males become progressively intolerant of each other and move to rutting areas in order to gather and defend groups of adult females. This time marks the change in groups’ composition, with an association between adult males and females and the start of matting activities (Clutton‐Brock, Guinness & Albon 1982). Usually described as a non‐territorial ungulate, red deer may present a highly territorial behaviour depending on the resources available (Carranza, Alvarez & Redondo 1990). This exceptional behaviour exhibit by red deer in Mediterranean environments is a consequence of the spatial heterogeneity of food patches, becoming more advantageous to defend the territory itself than the female groups (Carranza, Alvarez & Redondo 1990; Carranza, Fernandez‐Llario & Gomendio 1996). Outside the rut, the red deer shows a matriarchal society, where adult females live aggregate with sub‐adult ones but segregated from adult and sub‐adult males (Clutton‐Brock, Guinness & Albon 1982). This type of social interactions reflects the choice that the offspring has to face: remain at natal areas or leave? At this stage, females adopt a more philopatric behaviour and males a dispersal tactic (Loison et al. 2008). The dispersal tactic of sub‐adult males leads to the occupation of new territories and to the increase of the population range (Clutton‐Brock, Guinness & Albon 1982; Bonnet & Klein 1991; Soriguer et al. 1994). .

(28) 6 . Chapter 1 . Red deer reproductive cycle is considered highly synchronized, with a concentration of all the conceptions in a short period in order to increase the offspring survival (Festa‐Bianchet 1988; Coulson et al. 2003). Both sexes show a differential reproductive strategy, where the reproductive success of males lies in its physical condition at rutting period and the females’ success depends on the survival of the offspring (Clutton‐Brock, Guinness & Albon 1982; Main & du Toit 2005). The maternal investment is energetically very costly, resulting in a lower physical condition exhibit by lactating females than by barren ones (Clutton‐Brock, Guinness & Albon 1982; Cook et al. 2001). . 1.3. Red deer management Similar to what happened with other ungulates (e.g. wild boar Sus scrofa L.), the increase in abundance and geographical range of deer populations throughout Europe leads to conflicts with human activities, such as agriculture, forestry and road traffic (e.g. Putman 1997; Caudullo et al. 2003; Joys, Fuller & Dolman 2004; MacMillan 2004; Putman, Apollonio & Anderson 2011). Historically, the deer densities were controlled by large predators. However, their disappearance in great part of Europe results in uncontrolled deer populations. In order to reduce these wildlife populations the hunting pressure has increased, but sometimes with opposite results (Latham 1999). Currently, red deer represents both an important natural and economic resource as a source of considerable damage. In an agricultural perspective, the damages caused by this species affect irrigated (i.e. horticultural) and non‐irrigated crops (i.e. cereals and pastures), but also orchards and nursery crops (Putman & Moore 1998; Trdan & Vidrih 2008). Nevertheless, is on forest ecosystems that its impact is more evident, mainly as a consequence of browsing, bark‐stripping and rubbing (Putman & Moore 1998; Verheyden et al. 2006). In general, red deer selective browsing promotes changes in forest composition by retarding natural regeneration and modifying plant species diversity (Putman 1996; Kuiters & Slim 2002; Stewart et al. 2009). Besides the impact on vegetation, red deer also influences soil nutrients, soil biota and animal communities. At the soil level, red deer seems to increase the organic matter content and the microbial activity in the soil by the deposition of faeces (Mohr, Cohnstaedt & Topp 2005). However, it is possible that at certain environmental conditions, deer may exert a negative effect on some communities of invertebrate or vertebrate through its grazing activities to ground vegetation (Baines, Sage & Baines 1994; Mohr, Cohnstaedt & Topp 2005). Furthermore, the niche . .