Detectability, Habitat Relationships and Reliability As Biodiversity Surrogates
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ANDEAN TEMPERATE FOREST OWLS: DETECTABILITY, HABITAT RELATIONSHIPS AND RELIABILITY AS BIODIVERSITY SURROGATES by José Tomás Ibarra Eliessetch B.Sc., Pontificia Universidad Católica de Chile (Agricultural Engineering), 2005 M.Sc., Pontificia Universidad Católica de Chile (Conservation and Wildlife Management), 2007 M.Sc., University of Kent (Environmental Anthropology), 2010 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE AND POSTDOCTORAL STUDIES (Forestry) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) December 2014 © José Tomás Ibarra Eliessetch, 2014 Abstract South American temperate forests are globally exceptional for their high concentration of endemic species. These forests are among the most endangered ecosystems on Earth because nearly 70% of them have been lost. Current knowledge of most Neotropical forest owls is limited. I studied how environmental and habitat conditions might influence the ecology of sympatric forest owls, and evaluated whether owls can be used as surrogates for temperate forest biodiversity. Specifically, I examined (i) factors associated with the detectability, (ii) occurrence rates and habitat-resource utilization across spatial scales, and (iii) surrogacy reliability of the habitat-specialist rufous-legged owl (Strix rufipes) and the habitat-generalist austral pygmy-owl (Glaucidium nana) in southern Chile. During 2011- 2013, I conducted 1,145 owl surveys, 505 vegetation surveys and 505 avian point-transects across 101 sites comprising a range of conditions from degraded habitat to structurally complex old-growth forest stands. I recorded 292 detections of S. rufipes and 334 detections of G. nana. Detectability for both owls increased with greater moonlight and decreased with environmental noise, and greater wind speed decreased detectability for G. nana. Detection of both species was positively correlated with the detection of the other species. For S. rufipes, occurrence probability ranged from 0.05-1 across sites, and was positively associated with bamboo density and the variability in diameter at breast height of trees (multi-aged forests). For G. nana, occurrence ranged from 0.67-0.98, but no habitat characteristic was related to this species occurrence. Relative to G. nana, S. rufipes had lower total resource utilization, but achieved similar peak occurrence for resources related to stand-level forest complexity and forest homogeneity at the landscape scale. I found that only S. rufipes was a reliable surrogate for all avian biodiversity measures, including endemism and functional diversity. With increasing occurrence of habitat-specialist owls, the density of target specialized biodiversity (guilds and communities) increased non- linearly and peaked at the least degraded sites. This “specialist aggregation” was driven by forest-stand structural complexity. Forest management practices that maintain multi-aged stands with large trees and high bamboo cover will benefit both owl species, and likely will benefit vulnerable endemic species and specialized avian communities in temperate forests. ii Preface My thesis is written in a manuscript-based format. Chapters 2 through 4 represent independent chapters that have been or will be submitted in a similar format, except that I moved the general material on study area, study species and methods, common to the data sections, to Chapter 1. I took the lead in developing the research framework, conducting the research, data analysis and manuscript preparation for Chapters 2 to 4. A version of Chapter 2 has been published: Ibarra, J.T., Martin, K., Altamirano, T.A., Vargas, F.H., Bonacic, C., 2014. Factors associated with the detectability of owls in South American temperate forests: implications for nocturnal raptor monitoring. J. Wildl. Manage. 78, 1078-1086. A version of Chapter 3 has also been published: Ibarra, J.T., Martin, K., Drever, M.C., Vergara, G., 2014. Occurrence patterns and niche relationships of sympatric owls in South American temperate forests: a multi-scale approach. Forest Ecol. Manag. 331, 281- 291. iii Table of contents Abstract ............................................................................................................................ ii Preface ............................................................................................................................. iii Table of contents ............................................................................................................. iv List of tables .................................................................................................................... vi List of figures ................................................................................................................ viii Acknowledgements .......................................................................................................... x Dedication....................................................................................................................... xii Chapter 1. General introduction and thesis overview .................................................... 1 1.1. Background ........................................................................................................... 1 1.1.1. Endangered ecosystems and habitat-specialist owls .......................................... 1 1.1.2. Detectability of forest owls ............................................................................... 2 1.1.3. Owl niches and habitat suitability across spatial scales ..................................... 3 1.1.4. Forest owls as reliable biodiversity surrogates .................................................. 4 1.2. Thesis objectives .................................................................................................... 5 1.3. Study area .............................................................................................................. 5 1.4. Study species .......................................................................................................... 6 1.5. General field methods ........................................................................................... 7 1.5.1. Study design and allocation of survey effort ..................................................... 7 1.5.2. Nocturnal raptor surveys .................................................................................. 7 1.6. Thesis overview...................................................................................................... 8 Chapter 2. Detectability of owls in South American temperate forests ....................... 10 2.1. Introduction ............................................................................................................. 10 2.2. Methods ................................................................................................................... 11 2.2.1. Field methods ..................................................................................................... 11 2.2.2. Data analysis ...................................................................................................... 12 2.3. Results ...................................................................................................................... 13 2.3.1. Occurrence and detectability of rufous-legged owls ............................................ 13 2.3.2. Occurrence and detectability of austral pygmy-owls ........................................... 14 2.4. Discussion ................................................................................................................ 14 2.4.1. Recommendations for owl monitoring ................................................................ 17 2.5. Conclusions .............................................................................................................. 18 Chapter 3. Occurrence patterns of specialist and generalist owls across spatial scales in South American temperate forests ............................................................................ 26 3.1. Introduction ............................................................................................................. 26 3.2. Methods ................................................................................................................... 28 3.2.1. Field methods ..................................................................................................... 28 iv 3.2.2. Data analysis ...................................................................................................... 30 3.3. Results ...................................................................................................................... 31 3.3.1. Habitat suitability for owls.................................................................................. 32 3.3.2. Resource utilization and peak performance by owls ............................................ 33 3.4. Discussion ................................................................................................................ 33 3.4.1. Habitat suitability across spatial scales ............................................................... 34 3.4.2. Niche width of forest owls .................................................................................. 36 3.4.3. Recommendations for management .................................................................... 37 3.5. Conclusions .............................................................................................................