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Reproductive PATTERNS and Human-INFLUENCED Z]`Ynagj Af Zjgof Z]Yjk& Aehda[Ylagfk ^Gj the Conservation of LARGE Carnivores

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Reproductive PATTERNS and Human-INFLUENCED Z]`Ynagj Af Zjgof Z]Yjk& Aehda[Ylagfk ^Gj the Conservation of LARGE Carnivores P Natural and Department Ecology of Resource Management Fgjo]_aYfMfan]jkalqg^Da^]K[a]f[]kMfan]jkal]l]l^gjeadb¬ hilosophiae Doctor <]hYjlYe]flg\]:ag\an]jka\Y\q?]klaf9eZa]flYd& Universidad de León E-24071 León, Spain. www.unileon.es Reproductive patterns and human-influenced ( P Z]`YnagjafZjgofZ]Yjk&Aehda[Ylagfk^gj h D) the conservation of large carnivores. Thesis 2010:01 Thesis J]hjg\mckbgfke¬fkl]jg_e]ff]kc]kcYhlYl^]j\k]f\jaf_`gkZjmfZb¬jf& Cgfk]cn]fk]j^gjZ]nYjaf_]fYnklgj]jgn\qj& Andrés Ordiz %g_Zagnal]fkcYh Reproductive patterns and human-influenced behavior in brown bears Implications for the conservation of large carnivores Reproduksjonsmønster og menneskeskapt atferdsendring hos brunbjørn Konsekvenser for bevaringen av store rovdyr Philosophiae Doctor (PhD) Thesis Andrés Ordiz Dept. of Ecology and Natural Resource Management Norwegian University of Life Sciences & Dept. de Biodiversidad y Gestión Ambiental Universidad de León Ås/León 2010 UMB Thesis number 2010: 01 ISSN 1503-1667 ISBN 978-82-575-0913-2 This thesis has been conducted as a PhD research co-supervision agreement between the Norwegian University of Life Sciences and the University of León (Spain). I acknowledge the effort of J. E. Swenson, E. de Luis Calabuig and E. Panero to succeed in establishing the agreement. PhD supervisors Prof. Jon E. Swenson (main supervisor) Department of Ecology and Natural Resource Management Norwegian University of Life Sciences Pb. 5003, 1432 Ås, Norway Dr. Ole-Gunnar Støen Department of Ecology and Natural Resource Management Norwegian University of Life Sciences Pb. 5003, 1432 Ås, Norway Prof. Miguel Delibes de Castro Estación Biológica de Doñana, Consejo Superior de Investigaciones Científicas Avenida Américo Vespucio s/n Isla de la Cartuja E-41092 Sevilla, Spain Adjudication committee Prof. Tim Caro Department of Wildlife, Fish and Conservation Biology University of California 1 Shields Avenue, Davis CA 95616, USA Prof. Ilse Storch Department of Wildlife Ecology and Management Albert-Ludwigs-University of Freiburg Tennenbacher Straße 4 79106 Freiburg, Germany Prof. Svein Dale Department of Ecology and Natural Resource Management Norwegian University of Life Sciences Pb. 5003, 1432 Ås, Norway © Andrés Ordiz 2010 © Ecological Society of America 2008 © International Bear Association 2007, 2009, 2010 Contents Abstract……………………………………………………………………...1 Sammendrag (in Norwegian)..........................................................................2 Resumen (in Spanish).....................................................................................3 List of papers………………...……………………………………………...4 Synopsis…………………………………………...………………………..5 Placing this study in the frame of Conservation Biology………………...5 Major objectives of this thesis………………………………………........5 Brown bear, the model species…………………………………………...6 Study areas, study populations…………………………………………...7 Rationale behind the interest on females with cubs-of-the-year …………9 Human “disturbance”; Rationale behind a predator-prey approach......12 Discussion…………………………………………………………….....17 References cited……………………………………………………........24 Acknowledgements……………………………………………………..... 34 Compilation of papers.......................... ...................................................... 36 Abstract Monitoring the status of wildlife populations and understanding the effects of human activities on wildlife is central for wildlife conservation. Both goals are especially challenging for rare, secretive large carnivores. The objectives of this thesis, which includes 7 papers and manuscripts, were 1) to improve a method to monitor brown bear (Ursus arctos) populations based on counting females with cubs-of-the-year, including a study on reproductive patterns, and 2) to explore some effects of human activities on the behavior of large carnivores, using brown bears as a model species. Six of the studies were descriptive, based on empirical data, and the main findings in relation to the aforementioned objectives are the following: 1) Regarding females with cubs, the study improved distance-based criteria to distinguish family groups, and highlighted the need for cautious interpretation of population trends based on counting females with cubs. In addition, the results showing the existence of a spatiotemporal reproductive pattern for Scandinavian female bears suggested that interactions among individuals and behavior in solitary carnivores may play a larger role in their population dynamics than what has been traditionally assumed. 2) Bears showed a fine-scale behavioral response, both on a seasonal and daily basis, to human activities, which resembled the sensitive responses of prey to their predators. Bears appeared to rely on cover and distance to human settlements to avoid encounters with people, and altered their movement patterns when the bear hunting seasons started. Finally, the last manuscript unites demographic and behavioral perspectives of human persecution on large carnivores to discuss that such persecution is likely influencing not only them, but the apex, regulatory role that large carnivores can play in the ecosystems. The importance of behavioral responses and single individuals should be taken into account for conservation-oriented management of large carnivores, which should not only be based on demographic sustainability. The results also stressed carnivores’ need for cover and areas with low accessibility for humans. Results from the papers and synopsis led me to propose that large carnivores should not be systematically hunted. 1 Sammendrag Bestandsovervåking og forståelsen av de påvirkningene den menneskelige aktiviteten har på viltbestander er sentrale i bevaringen av viltarter. Begge mål er spesielt utfordrende for sky og sjeldne store rovdyr. Målene for denne oppgaven som inneholder sju arbeider var: 1) å forbedre en overvåkningsmetode for brunbjørnbestander (Ursus arctos) basert på telling av binner med årsunger, som også inkluderer en studie av reproduksjonsmønster, og 2) å undersøke noen av påvirkningene menneskelig aktivitet har på store rovdyrs atferd med brunbjørn som en modellart. Seks av arbeidene er deskriptive basert på empiriske data og hovedresultatene i henhold til de nevnte målene er: 1) Med hensyn til binner med unger forbedret studien de distansebaserte kriteriene for å skille familiegrupper, og trekker frem nødvendigheten av en varsom tolkning av bestandstrender basert på tellinger av binner med årsunger. I tillegg, resultatene som viser forekomsten av et spatiotemporalt reproduksjonsmønster hos skandinaviske binner antyder at gjensidig påvirkning mellom individer og atferd blant solitære rovdyr kan spille en større rolle i deres bestandsdynamikk enn det man tidligere har antatt. 2) Bjørnene viste en finskala tilpassning i atferd, både på sesong- og døgnbasis, overfor menneskelig aktivitet, som likner på den atferd byttedyr har overfor rovdyr. Bjørner ser ut til å bruke skjul og avstand til bebyggelse for å unngå møte med folk, og endrer deres bevegelsesmønster når jaktsesongen startet. Det siste manuskriptet forener demografiske og atferdsmessige perspektiver på menneskelig forfølgelse av store rovdyr og diskuterer at slik forfølgelse trolig påvirker ikke bare rovdyrene, men også den regulerende rollen som topp- predator de store rovdyrene bør ha i et økosystem. Viktigheten av atferdmessig respons og enkeltindivider bør tas hensyn til i bevaringsorientert forvaltning av store rovdyr og ikke bare være basert på demografisk bærekraftighet. Resultatene viser betydningen av at rovdyr trenger skjul og områder som er lite tilgjengelig for mennesker. Resultatene av disse arbeidene har fått meg til å foreslå at store rovdyr bør ikke jaktes systematisk. 2 Resumen El seguimiento del estado poblacional y la evaluación de los efectos de las actividades humanas sobre la fauna son esenciales para su conservación. Ambas metas son un reto especial trabajando con grandes carnívoros, en su condición de animales elusivos y escasos. Los objetivos de esta tesis, que incluye siete artículos y manuscritos, fueron 1) mejorar un método de seguimiento de poblaciones de oso pardo (Ursus arctos) basado en el conteo de osas con crías, incluyendo un estudio sobre patrones de reproducción, y 2) explorar algunos efectos de actividades humanas sobre el comportamiento de los grandes carnívoros, con los osos como especie modelo. Seis de los trabajos fueron descriptivos, basados en datos empíricos, y sus resultados esenciales, en relación con los objetivos expuestos, son los siguientes: 1) En relación a las osas con crías, el estudio mejoró los criterios basados en distancias entre observaciones para distinguir grupos familiares, y resaltó la necesidad de interpretar con cautela tendencias de la población basadas en el conteo de osas con crías. Además, los resultados mostraron la existencia de un patrón espacial y temporal de la reproducción, sugiriendo que las interacciones entre individuos y el comportamiento en carnívoros solitarios pueden jugar un papel más importante en las dinámicas poblacionales de lo que se viene suponiendo. 2) Los osos mostraron una respuesta comportamental concreta a las actividades humanas, tanto a escala estacional como diaria, que recuerda a las sensibles respuestas de las presas respecto a sus predadores. Parece que los osos utilizan la cobertura de la vegetación y la distancia a asentamientos humanos como medios para evitar encuentros con gente, y modifican sus patrones de movimiento cuando comienza la temporada de caza. El último manuscrito liga perspectivas
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