Forestry, Faculty of
DSpace
UBCV
Kennedy, Tabitha
Master of Science - MSc
University of British Columbia
Climate change continues to alter ecological systems, fauna, and flora within Canada. Many species are predicted to undergo range shifts as they track changing environmental conditions. Such range shifts add complexity to protected area (PA) planning if coverage within PA networks is reduced. In my MSc thesis, I assessed potential future distributions for a climate-sensitive family, Trochilidae (Hummingbirds), and the effectiveness of protected areas in tracking their shifting distributions. Following a maximum entropy approach, I modelled current and future species distribution for four commonly occurring species across British Columbia (B.C.): Annaâs Hummingbird (Calypte anna), Black-chinned Hummingbird (Archilochus alexandri), Calliope Hummingbird (Selasphorus calliope), and Rufous Hummingbird (Selasphorus rufus). Models included climatic, ecological, and geographic variables and were built using citizen science observations (eBird program) from 1981 to 2020 to assess the impact of climate (direct) and climate-induced habitat change (indirect) on potential future distributions. I found that climate-related variables were the strongest predictors of distribution for all species. Under the Representative Concentration Pathway (RCP) +8.5 W/m² climate warming scenario, northward range expansions were projected for all four hummingbird species. Absolute protected area coverage was projected to increase for all species following range expansions (Annaâs: +6,378 km², Black-chinned: +20,952 km², Calliope: +23,829 km², Rufous: +31,619 km²), while relative protected area coverage was projected to increase for all species (Calliope: +1.6%, Black-chinned: +2%, Annaâs: +10.8%) except for Rufous Hummingbird (â 3.8%). Greater range overlap was also projected to occur as a result of range expansion within the province. My findings indicate that while some future range expansion is predicted to fall within the boundaries of protected areas, a higher degree of range expansion is predicted to occur outside these boundaries in non-protected areas. Hence, future conservation efforts for declining species such as the Rufous Hummingbird will benefit from strengthened habitat protection and conservation measures within, but especially outside the boundaries of protected areas. Future studies will be needed to address the implications of greater interspecific range overlap among breeding hummingbird species in B.C., and whether native vegetation that contributes key floral resources to hummingbirds will follow similar range shifts.
EXAMINING THE INFLUENCE OF CLIMATE CHANGE ON DISTRIBUTION SHIFTS AND THE EFFICACY OF PROTECTED AREAS FOR HUMMINGBIRDS IN BRITISH COLUMBIA by Tabitha Kennedy B.Sc., Carleton University, 2020 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE AND POSTDOCTORAL STUDIES (Forestry) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) August 2023 © Tabitha Kennedy, 2023 ii The following individuals certify that they have read, and recommend to the Faculty of Graduate and Postdoctoral Studies for acceptance, the thesis entitled: Examining the influence of climate change on distribution shifts and the efficacy of protected areas for hummingbirds in British Columbia submitted by Tabitha Kennedy in partial fulfilment of the requirements for the degree of Master of Science in Forestry Examining Committee: Dr. Kathy Martin, Professor, Forest and Conservation Sciences, UBC Co-supervisor Dr. Scott Wilson. Adjunct Professor, Forest and Conservation Sciences, UBC Co-supervisor Dr. Sarah Gergel, Professor and Associate Dean, Forest and Conservation Sciences, UBC Supervisory Committee Member Dr. Jill Jankowski, Associate Professor, Zoology, UBC Additional Examiner iii Abstract Climate change continues to alter ecological systems, fauna, and flora within Canada. Many species are predicted to undergo range shifts as they track changing environmental conditions. Such range shifts add complexity to protected area (PA) planning if coverage within PA networks is reduced. In my MSc thesis, I assessed potential future distributions for a climate-sensitive family, Trochilidae (Hummingbirds), and the effectiveness of protected areas in tracking their shifting distributions. Following a maximum entropy approach, I modelled current and future species distribution for four commonly occurring species across British Columbia (B.C.): Annaâs Hummingbird (Calypte anna), Black-chinned Hummingbird (Archilochus alexandri), Calliope Hummingbird (Selasphorus calliope), and Rufous Hummingbird (Selasphorus rufus). Models included climatic, ecological, and geographic variables and were built using citizen science observations (eBird program) from 1981 to 2020 to assess the impact of climate (direct) and climate-induced habitat change (indirect) on potential future distributions. I found that climate-related variables were the strongest predictors of distribution for all species. Under the Representative Concentration Pathway (RCP) +8.5 W/m2 climate warming scenario, northward range expansions were projected for all four hummingbird species. Absolute protected area coverage was projected to increase for all species following range expansions (Annaâs: +6,378 km2, Black-chinned: +20,952 km2, Calliope: +23,829 km2, Rufous: +31,619 km2), while relative protected area coverage was projected to increase for all species (Calliope: +1.6%, Black-chinned: +2%, Annaâs: +10.8%) except for Rufous Hummingbird (â 3.8%). Greater range overlap was also projected to occur as a result of range expansion within the province. My findings indicate that while some future range expansion is predicted to fall within the boundaries of protected areas, a higher degree of range expansion is predicted to occur outside iv these boundaries in non-protected areas. Hence, future conservation efforts for declining species such as the Rufous Hummingbird will benefit from strengthened habitat protection and conservation measures within, but especially outside the boundaries of protected areas. Future studies will be needed to address the implications of greater interspecific range overlap among breeding hummingbird species in B.C., and whether native vegetation that contributes key floral resources to hummingbirds will follow similar range shifts. v Lay Summary Many species are predicted to shift ranges under climate change as they adapt and search for suitable habitat. This movement will create new challenges for protecting species and their habitats, especially if coverage across protected area (PA) networks is reduced for species undergoing range shifts. I used eBird citizen science data and considered climate, ecology and geology to examine present and future distributions for four hummingbird species in British Columbia (B.C.). I found that variables related to climate were most important in influencing hummingbird breeding ranges, and that all species ranges were predicted to expand northward in B.C. This expansion will mean greater relative PA coverage, except for Rufous Hummingbird. My work highlights the importance of protecting habitat for hummingbirds both within and outside of PAs. Future conservation efforts for declining species like Rufous Hummingbird will benefit from strengthening habitat protection and conservation measures within, but especially outside PA boundaries. vi Preface This thesis and the methodology and analysis therein were written and conducted by Tabitha Kennedy, with supervision from the main committee members: Dr. Scott Wilson (UBC Forestry, ECCC), Dr. Kathy Martin (UBC Forestry), Dr. Sarah Gergel (UBC Forestry) and valuable methodology design and analysis contributions from Dr. Davide Scridel (National Research Council of Italy). The project used bird observation data available through the online citizen science database eBird (https://ebird.org/home). vii Table of Contents Abstract ......................................................................................................................................... iii Lay Summary ................................................................................................................................ v Preface ........................................................................................................................................... vi Table of Contents ........................................................................................................................ vii List of Tables ................................................................................................................................ ix List of Figures ................................................................................................................................ x Acknowledgements ..................................................................................................................... xii Chapter 1: Introduction .............................................................................................................. 1 1.1 Impacts of climate change and land use on biodiversity ..................................................... 1 1.2 Protected areas as a tool for conserving biodiversity ........................................................... 2 1.2.1 Protected areas: definition, importance, and benefits ...................................................... 2 1.2.2 Limitations to protected areas under future change ......................................................... 5 1.2.3 Protected areas and the importance of hummingbird conservation ................................. 7 1.2.4 B.C. hummingbirds distribution and conservation status ................................................ 9 1.3 Study goals, objectives, and hypotheses ............................................................................ 11 Chapter 2: Methods ................................................................................................................... 15 2.1 Study area........................................................................................................................... 15 2.2 Hummingbird data ............................................................................................................. 17 2.3 Predictor variables ............................................................................................................. 18 2.4 Protected areas data............................................................................................................ 20 2.5 Data analysis ...................................................................................................................... 20 Chapter 3: Results ..................................................................................................................... 24 3.1. Model performance ............................................................................................................ 24 3.2 Importance of predictor variables ...................................................................................... 24 3.3 Future range expansion ...................................................................................................... 26 3.4 Changes in protected area coverage ................................................................................... 28 3.5 Changes in range overlap ................................................................................................... 29 Chapter 4: Discussion ................................................................................................................ 41 4.1 Overview ............................................................................................................................ 41 viii 4.2 Northward range expansion ............................................................................................... 41 4.3 Protected area (PA) coverage and effectiveness ................................................................ 45 4.4 Implications of future range overlap .................................................................................. 47 4.5 Caveats ............................................................................................................................... 49 4.6 Conservation implications ................................................................................................. 52 4.7 Future opportunities and studies ........................................................................................ 53 Bibliography ................................................................................................................................ 55 Appendix ..................................................................................................................................... 71 ix List of Tables Table 1. Climate, topography, and habitat variables selected in the final models for each species (see Table A1 for full list of variables), with relative variable importance (%) and standard deviation (in brackets)................................................................................................................... 23 Table 2. Range size (km2) and proportion (%) of potential suitable habitat for hummingbirds across B.C. and within protected (PAs) and in unprotected areas (UPAs) under current and future scenarios (2080s) according to ACCESS 1.0 climatic projections. .............................................. 40 Table 3. Range overlap (km2) for hummingbirds across B.C. under current and future scenarios (2080s) according to ACCESS 1.0 climatic projections. .............................................................. 40 Table A1. List of climatic, habitat and topographic variables considered in the modelling procedureâ¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦â¦71 x List of Figures Figure 1. Terrestrial protected areas in British Columbia (UNEP-WCMC & IUCN, 2021). ...... 14 Figure 2. Biogeoclimatic Ecosystem Classification (BEC) zones of British Columbia (UBC Centre for Forest Conservation Genetics, 2023)........................................................................... 16 Figure 3. Ecoprovinces of British Columbia (Demarchi, 1996) .................................................. 16 Figure 4. Modelled potential breeding range distributions of Anna's Hummingbird in B.C. for current (1981-2020) and future (2080s) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2 A) Suitability according to the maximum training sensitivity plus specificity (MTSS) threshold is depicted in green. B) Suitability gradient ranging from 0.0 to 1.0. ............................................................................................................................................ 30 Figure 5. Modelled potential breeding range distributions of Black-chinned Hummingbird in B.C. for current (1981-2020) and future (2080s) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2 A) Suitability according to the maximum training sensitivity plus specificity (MTSS) threshold is depicted in green. B) Suitability gradient ranging from 0.0 to 1.0. .............................................................................................................................. 31 Figure 6. Modelled potential breeding range distributions of Calliope Hummingbird in B.C. for current (1981-2020) and future (2080s) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2 A) Suitability according to the maximum training sensitivity plus specificity (MTSS) threshold is depicted in green. B) Suitability gradient ranging from 0.0 to 1.0. ............................................................................................................................................ 32 Figure 7. Modelled potential breeding range distributions of Rufous Hummingbird in B.C. for current (1981-2020) and future (2080s) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2 A) Suitability according to the maximum training sensitivity plus specificity (MTSS) threshold is depicted in green. B) Suitability gradient ranging from 0.0 to 1.0. ............................................................................................................................................ 33 Figure 8. Anna's Hummingbird occupancy in B.C. protected areas under current (1981-2020) and future (2080s: 2071-2100) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2. .............................................................................................................. 34 Figure 9. Black-chinned Hummingbird occupancy in B.C. protected areas under current (1981-2020) and future (2080s: 2071-2100) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2. ............................................................................................. 35 Figure 10. Calliope Hummingbird occupancy in B.C. protected areas under current (1981-2020) and future (2080s: 2071-2100) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2. .............................................................................................................. 36 Figure 11. Rufous Hummingbird occupancy in B.C. protected areas under current (1981-2020) and future (2080s: 2071-2100) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2. .............................................................................................................. 37 Figure 12. Current (1981-2020) range overlap for Anna's, Black-chinned, Calliope, and Rufous Hummingbirds. ............................................................................................................................. 38 Figure 13. Future (2080s) range overlap for Anna's, Black-chinned, Calliope, and Rufous Hummingbirds. ............................................................................................................................. 39 xi Figure A1. Most supported variables defining Annaâs Hummingbird distribution (temperature variables are expressed in °C/10, precipitation in mm, slope in degrees, topographic position index values above zero show locations that are higher than then average of immediately surrounding terrain). Permutation importance (highest to lowest): Annual mean temperature (52.9%), Temperature annual range (29.0%), Slope (15.0%), Urban land cover (1.7%), Coastal Douglas fir (1.5%). ....................................................................................................................... 72 Figure A2. Most supported variables defining Black-chinned Hummingbird distribution (temperature variables are expressed in C°/10, precipitation in mm, slope in degrees, topographic position index values above zero show locations that are higher than then average of immediately surrounding terrain). Permutation importance (highest to lowest): Mean temperature of warmest quarter (54.6%), Temperature annual range (15.4%), Precipitation of warmest quarter (9.9%), Topographic position index (9.4), Solar radiation (6.8%), Slope (1.8%), Ponderosa Pine (1.1%), Montane Spruce (1.0%). ............................................................................................................... 73 Figure A3. Most supported variables defining Calliope Hummingbird distribution (temperature variables are expressed in C°/10, precipitation in mm, slope in degrees, topographic position index values above zero show locations that are higher than then average of immediately surrounding terrain). Permutation importance (highest to lowest): Temperature annual range (47.2%), Annual mean temperature (29.4%), Precipitation of warmest quarter (12.2%), Slope (5.7%), Coastal Western Hemlock (3.0%), Ponderosa Pine (2.1%). ............................................ 74 Figure A4. Most Most supported variables defining Rufous Hummingbird distribution (temperature variables are expressed in C°/10, precipitation in mm, slope in degrees, topographic position index values above zero show locations that are higher than then average of immediately surrounding terrain). Permutation importance (highest to lowest): Mean temperature of warmest quarter (37.8%), Temperature annual range (23.0%), Slope (20.6%), Topographic position index (15.6%), Coastal Douglas fir (1.9%), Coastal Western Hemlock (1.3%). ................................... 75 xii Acknowledgements This research project would not be possible without the encouragement and support of my supervisory committee, family, and friends. Many sincere thanks to Dr. Scott Wilson, Dr. Kathy Martin, and Dr. Sarah Gergel for your valued mentorship, expertise, and support in shaping this research project. Special thanks to Dr. Davide Scridel for sharing your wealth of knowledge on species distribution modelling and for being an excellent instructor in all things R and Maxent. To my grandparents, Pierre and Karen Cauchy, thank you for your love, support, and eagerness to hear updates on my research progress during our phone calls. To my parents, Douglas and Chantal Kennedy, thank you for loving and supporting me throughout the highs and lows of graduate school, and for always believing in me. To my very dear friends Jillian Auger, Kasuna Gamage, and Sarah Robinson, thank you for all the laughs and emotional support you brought me during long nights of lit review and writing, and for being the greatest hypewomen a STEM girl could ever have. To my team and community at the UBC Sustainability Hub, thank you for becoming my second home on campus and for being such wonderful, hilarious, and inspiring people to know and work with. 1 Chapter 1: Introduction 1.1 Impacts of climate change and land use on biodiversity Anthropogenic climate change and land-use activities pose major threats to biodiversity on a global scale (IPCC, 2021; Newbold et al., 2015; Lenoir & Svenning, 2015). Habitat degradation, fragmentation, and loss resulting from human activities have compromised ecological integrity and heightened threats of species extinction (Ceballos et al., 2020; Diaz et al., 2019). Climate change processes, which encompass a range of interconnected mechanisms influencing the Earthâs climate, have similarly been disrupted by anthropogenic activities (IPCC, 2022). Humans have inadvertently altered natural processes such as the greenhouse effect through the burning of fossil fuels and deforestation (IPCC, 2022). Such activities have led to increased concentrations of carbon dioxide (CO2), methane (CH4), nitrous oxide (N2O), and other greenhouse gases (GHG) in the atmosphere, causing global temperatures to rise in a phenomenon known as global warming (IPCC, 2022). Global warming is one of the greatest threats to maintaining life on Earth, as it disrupts all of the planetâs subsystems (lithosphere, hydrosphere, biosphere, atmosphere) in addition to triggering positive feedback loops that further amplify warming effects (IPCC, 2022). Global average precipitation has increased by approximately 2% over the past 100 years (IPCC, 2007), while global temperatures are estimated to have increased by 1.0°C compared to pre-industrial levels, with predicted increases of 0.2 ± 0.1 °C per decade if current GHG emission rates remain constant (IPCC, 2018). Certain geographical regions are experiencing warming at higher rates than the global average, such as mountain areas with an average warming rate of 0.3 ± 0.2 °C per decade (IPCC, 2018; IPCC, 2021). If we are to adequately conserve biodiversity amidst such changes, we need to understand 2 both how species are likely to respond to climate and land use change, and how best to protect them as ecosystems shift in response to the current and future levels of environmental change. Species affected by climate and land use change typically have three options available to them: adapt to their evolving environment, perish, or relocate (Rinawati et al., 2013). Numerous species across the globe have exhibited the latter, as displayed in both poleward and upslope shifts in their respective geographic distributions as they track shifting habitat conditions (Lenoir & Svenning, 2015; Chen et al., 2011). Such trends have resulted in the disaggregation of local communities and the introduction of warm-adapted species to habitats previously outside of their historic range (Heller & Zavaleta, 2009; Parmesan, 2006). Mountain ecosystems are of particular conservation concern, as they possess unique and diverse ecological communities (Rodríguez-Rodríguez et al., 2011; Pearce-Higgins & Martin, 2023; Chamberlain et al., 2023; Martin et al., 2023), and are especially susceptible to impacts of climate change (Boyle & Martin, 2015; Pearce-Higgins & Martin, 2023; Rangwala & Miller, 2012). Species which inhabit mountain ecosystems face limitations for upwards movement imposed by the topography of their environment (La Sorte & Jetz, 2010), highlighting the challenges that rapid distribution shifts pose for their future protection (Freeman et al., 2018). 1.2 Protected areas as a tool for conserving biodiversity 1.2.1 Protected areas: definition, importance, and benefits In the face of the ongoing global biodiversity crisis, protected areas are a cornerstone conservation tool for preventing and reversing current rates of extinction (Cazalis et al., 2020; 3 Geldmann et al., 2019; Wulder et al., 2018). By limiting harmful anthropogenic activities such as habitat degradation, loss, and wildlife overexploitation, protected areas can be an invaluable mechanism for safeguarding ecosystems and species (Scridel et al., 2021; Cazalis et al., 2020). Relative to unprotected areas, protected areas have been shown to retain more species with specialized habitat requirements, species of higher conservation concern on the IUCN Red List, and species with narrow habitat ranges (Cazalis et al., 2020). The Giant Panda is one example of a high priority species of global conservation concern that has benefitted greatly from the creation and continued management of protected areas (Huang et al., 2020). From the early 1960s until 2015, Chinaâs Giant Panda population received a protected area network expansion from 4 to 67 reserves, increasing Giant Panda habitat from 9,347 km2 to 33,118 km2 between the First and Fourth National Giant Panda Surveys (Wei et al., 2018a; Huang et al., 2020). This protected area network expansion helped to protect 54% of the Giant Pandaâs habitat range across China. These protected areas are considered to have played a key role in supporting a 17% population increase between the years 2004 to 2014 (Wei et al., 2018b; Swaisgood et al., 2016). Within Canada, protected areas are defined as âa clearly defined geographical space, recognised, dedicated and managed, through legal or other effective means, to achieve the long-term conservation of nature with associated ecosystem services and cultural values." (ECCC, 2020). These areas can include National, Provincial, and Territorial parks, National Wildlife Areas, Indigenous Protected Areas, Migratory Bird Sanctuaries, and Marine Protected Areas (ECCC, 2020). Canadaâs protected area system began in 1885 with the creation of Banff National Park. The Federal park system has since expanded to 47 National Parks and park reserves, 7 of which are located in British Columbia (Parks Canada, 2021). Other protected areas within British Columbia include National Wildlife Areas, Provincially Protected Areas managed 4 by BC Parks, and Wildlife Management Areas managed by the British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development (Government of Canada, 2021). As of 2020, roughly 11.7% of Canadaâs terrestrial area and 15.5% of British Columbia terrestrial area are designated as Protected Areas (ECCC, 2020; Figure 1). As with the conservation success of the Giant Panda in China, protected areas have played an important role in the recovery of endangered species in Canada. Wood Buffalo National Park, currently the largest national park in Canada, was established in northeastern Alberta in 1922 to conserve the last remaining bison herds in the country (Parks Canada, 2022). Historically, Canadaâs Wood Bison population was estimated to have been near 168,000 individuals in the early 1800âs. However, threats such as habitat change, overhunting, and severe winter weather likely contributed to population declines, resulting in only a few hundred individuals remaining by the late 1800s (Government of Canada, 2020). Critical habitat protection and disease management programs in Wood Buffalo National Park have helped this population recover to approximately 8,587 free-ranging individuals by 2015 (ECCC, 2018; Government of Canada, 2020). Wood Buffalo National Park is also home to one of the rarest and most endangered bird species in North America, the Whooping Crane (Nature Conservancy of Canada, 2022). The Aransas-Wood Buffalo Whooping Crane population was on the verge of extinction in the mid-1940s with only 21 individuals remaining. Through captive breeding programs and protection of key breeding habitat in Wood Buffalo National Park, the Whooping Crane population rose to 800 individuals in 2018, with 98 nests recorded in a 2017 survey of the park (Nature Conservancy of Canada, 2022). 5 1.2.2 Limitations to protected areas under future change While protected areas are a crucial tool for biodiversity conservation, there are also challenges to their implementation and effectiveness. Protected areas are generally regarded positively by the public due to their perceived benefits for conservation and ecosystem services, including forest and wildlife conservation, reduced pollution and protection of soil and watershed integrity (Allendorf, 2022). Protected areas also present social, cultural, economic, and health benefits to local communities through supporting community development programs, ecotourism, and opportunities for collaboration between local residents and managers of protected areas to achieve conservation goals (Allendorf, 2022). Despite this generally positive attitude towards protected areas, there are often negative perceptions which can include both public and private sector frustration with lack of legal means for natural resource extraction, conflicts with wildlife, disappointment resulting from unmet project goals in protected areas, and negative interactions between the public and park management (Allendorf et al., 2022). The establishment of protected areas in recent past have negatively impacted Indigenous communities through forced displacement from their ancestral lands, disruption of cultural practices, and loss of access to traditional resources (Redford & Fearn, 2007). On Canadaâs path towards reconciliation, Indigenous Protected Areas (IPAs) such as Gwaii Haanas National Park have been established to strengthen Indigenous autonomy and governance and prioritize conservation of both natural ecosystems and areas of cultural heritage (Redford & Fearn, 2007). A particularly common criticism for protected areas has been concern over protected area establishment in areas of weak biodiversity importance (Maxwell, 2020). Such situations may arise unintentionally due to poor planning and lack of understanding for the ecological interactions and biodiversity of a particular area (Pressey et al., 2015), or in some cases, 6 intentionally via selection informed by political and economic bias (Jantke et al., 2018; Lemieux et al., 2019; Joppa & Pfaff, 2009). Indeed, protected areas established under static conservation concepts often do not account for long-term environmental, climactic, and species distribution changes, which may result in failure to support species and habitat persistence for which an area was designed to protect (Sieck et al., 2011). Speciesâ representation in protected area systems can be highly variable, in part due to many protected areas having been created prior to the 1980s when terms such as biological diversity and biodiversity were coined (Hannah et al., 2007). Range shifts induced by climate change may drive species out of protected area boundaries, thereby reducing the effectiveness of static protected areas and their relevance in future conservation plans (Hannah et al., 2007). Over the next century, ecosystems, flora, and fauna that exist within protected areas are predicted to face significant impacts resulting from climate and land use change (Normand et al., 2007; Rannow et al., 2014). However, as impacts will likely vary depending on species, ecosystem type, and location, there is often no simple solution for creating a climate resilience strategy that will be applicable across all protected areas (Rannow et al., 2014). Rannow et al. (2014) argues that climate resiliency strategies for protected areas should be developed on a case-by-case basis, taking into consideration the unique sensitivities and challenges posed by climate change, along with the options available for action within each area. As anthropogenic climate change and land use change are expected to disrupt spatial and temporal scales within protected areas, policy makers and managers of protected areas will need to amend conservation objectives and guiding principles within protected areas legislation to reflect these changes (Dietz et al., 2021; Rannow et al., 2014). This could include expanding protected area boundaries 7 to create buffers from external pressures, or the acquisition of neighboring lands to accommodate future range shifts in species of conservation concern (Hannah et al., 2007; Rannow et al., 2014). 1.2.3 Protected areas and the importance of hummingbird conservation Decisions on protected area designations are often made to protect species and communities under current climate and environmental conditions (Elsen et al., 2018). However, recent range shifts observed in species occupying elevational and latitudinal gradients will present new challenges in protected area planning, if or when the ranges of species targeted for protection shift outside of current protected area boundaries (Elsen et al., 2018). Therefore, research is needed to identify how faunal and floral communities are expected to change under future climate and environmental conditions, and how this will influence their coverage in protected area networks. The goal of my Masters thesis will hence be to assess range shifts in birds of conservation concern in British Columbia resulting from both direct (i.e. climate) and indirect (i.e. climate-induced habitat alterations) effects of climate change in order to provide insight to inform and support protected area planning and policy within B.C. Birds have proven to be an excellent taxa for understanding and monitoring environmental integrity, as their interactions and connectivity with the environment at varying spatial and temporal scales make them excellent indicators of environmental change (Carignan & Villard, 2002; Morrison, 1986). Since 1970, North America has observed rapid declines in avian abundance, with an estimated net loss of three billion birds across the continent (Rosenberg et al., 2019). Concern for the Trochilidae (Hummingbird) family have escalated as 60% of species populations are in decline (IUCN, 2020). Hummingbirds share an integral connection to 8 ecological systems, as many co-dependent and mutualistic relationships exist between various species of hummingbirds and plants (Leimberger et al., 2022). It is estimated that over 7,000 plant species depend on hummingbirds for pollination, while hummingbirds depend on plants as sources of nectar (Abrahamczyk et al., 2015). Despite their small body size, hummingbirds can be fiercely aggressive and territorial, often partaking in interspecific and intraspecific competition (LópezâSegoviano et al., 2018). Niche partitioning among hummingbird species is often determined by aggressive dominance, body size, and success in defending quality food sources (LópezâSegoviano et al., 2018). As climate and land use change continue to influence species range shifts, hummingbirds may experience new overlap in habitat occupancy with other hummingbird species, raising interest and concern in the potential implications of interspecific competition among species that share similar diets and foraging habits. My thesis will focus on 4 species of hummingbirds that currently overlap to differing extents in British Columbia, the Annaâs Hummingbird (Calypte anna), Black-chinned Hummingbird (Archilochus alexandri), Calliope Hummingbird (Selasphorus calliope), and Rufous Hummingbird (Selasphorus rufus). All hummingbird species selected are common breeding species in British Columbia (Clark & Russell, 2020; Baltosser & Russell, 2020; Calder & Calder, 2020; Healy & Calder, 2020). Annaâs Hummingbird is currently expanding its range in British Columbia, while the Rufous Hummingbird is a species of conservation concern in Canada owning to long-term population declines across their range in North America (English et al., 2021). 9 1.2.4 B.C. hummingbirds distribution and conservation status Annaâs Hummingbirds have an S5 (Yellow list) B.C. Provincial Conservation Status as of 2022, indicating that the species is demonstrably widespread, abundant, and secure (B.C. Conservation Data Centre, 2022; Government of B.C., 2022). Annaâs Hummingbirds are a resident species in British Columbia, where the population core is primarily restricted to the southern coast of the province within the Georgia Depression ecoregion (Moran & Fraser, 2015a; Government of Canada, 2015). The breeding range for Annaâs Hummingbird has been expanding in coastal B.C. to encompass the entire lower Fraser Valley, eastern and southern Vancouver Island, as well as the southern Gulf Islands (Moran & Fraser, 2015a). Occasional breeding records have been documented as far north as the Stikine River (Moran & Fraser, 2015a). Some breeding occurs in the southern interior of the province near the Similkameen watershed (Moran & Fraser, 2015a). Annaâs Hummingbirds breed at elevations ranging from sea level to approximately 500 meters, with a preference for open habitats associated with humans and urban areas (Moran & Fraser, 2015a). Recent trends indicate that Annaâs Hummingbird populations and breeding range have been increasing, likely in part due to the proliferation of hummingbird feeders year-round in urban areas (Government of Canada, 2015). The Black-chinned Hummingbird exhibits long-term range expansion within British Columbia (Moran & Fraser, 2015b). The B.C. Provincial Conservation Status as of 2022 is S5B (Yellow list), indicating the species is locally secure and not of conservation concern (B.C. Conservation Data Centre, 2022; Government of B.C., 2022). The northern limit of this speciesâ breeding range occurs in southern British Columbia (Moran & Fraser, 2015b). The population core is found in the Southern Interior within the Okanagan and Similkameen valleys, with other breeding areas including Shuswap, Thompson-Nicola, Creston Valley, and the Southern Rocky 10 Mountain Trench (Moran & Fraser, 2015b). Nesting habitat preferences are deciduous trees and shrubs occurring in riparian woodlands and human-inhabited areas, typically at elevations ranging from 250 to 1,000 meters above sea level (Moran & Fraser, 2015b). The Calliope Hummingbird has an S5B (Yellow list) B.C. Provincial Conservation Status as of 2022 due to a moderate and long-term population increase in the province (B.C. Conservation Data Centre, 2022; Government of B.C., 2022). The Calliope Hummingbirdâs breeding range occurs throughout a large portion of the Interior, with species abundance most concentrated at elevations below 500 meters in Douglas-fir and Ponderosa Pine biogeoclimatic zones (Moran & Fraser, 2015c). As well, Calliope Hummingbirds display moderate-abundance in Sub-boreal Spruce habitat of the northern Central Interior, occupying elevations ranging 670 to 1,250 meters above sea level (Moran & Fraser, 2015c). Habitat preference for this species is warmer valleys within the Southern Interior containing shrubland and forest openings associated with human activity, such as fire or timber harvest (Moran & Fraser, 2015c). Breeding Bird Survey (BBS) trends indicate that Rufous Hummingbird populations have been in decline across North America since the 1970s, with rates of decline having increased from 2009 to 2019, possibly resulting from poor juvenile recruitment (English et al., 2021; Drake et al., 2022; Government of Canada, 2022). The B.C. Provincial Conservation Status for Rufous Hummingbirds was last updated as apparently secure (S4B) in 2015 with a Yellow List Status (B.C. Conservation Data Centre, 2022; Government of B.C., 2022). However, more recent national surveys have identified Rufous Hummingbirds as priority species for conservation within Canadaâs Bird Conservation Region Strategies (BCRs) in the Northern Pacific Rainforest, Northern Rockies, and Northern Interior Forest (Government of Canada, 2022). Similarly, the Rufous Hummingbird has been listed as a High Priority Candidate by the Committee on the 11 Status of Endangered Wildlife in Canada (COSEWIC) as of November 2021 (COSEWIC, 2021). Of the four species selected for my study, the Rufous Hummingbird occupies the highest breeding elevations, ranging from sea level to 1,830 meters with a preference for mountain meadows and forests within southwestern Canada (Cornell Lab of Ornithology, 2022; Nature Conservancy of Canada, 2020). As British Columbia holds the most widespread breeding population of Rufous Hummingbirds in Canada, mountain ranges within the province will likely act as strongholds for this internationally threatened species. 1.3 Study goals, objectives, and hypotheses My research will integrate long-term citizen science data with databases on protected areas and projected climate change, to address the following questions: 1) how are the distribution and community composition of breeding hummingbird communities in British Columbia (B.C.), Canada expected to change within the next decades due to changes in climate, 2) which hummingbird communities are currently under protection in B.C. and how will protection change as communities shift due to changes in climate, 3) how can protected areas planning in B.C. accommodate range shifts within hummingbird communities. I produced models of correlative species distribution using the maximum entropy approach (Maxent, e.g. Scridel et al. 2021) to test the hypothesis that climate change will result in distribution shifts in breeding hummingbirds across British Columbia. Specifically, I hypothesize a northward shift in the future breeding ranges of all 4 study species (Rufous Hummingbird, Annaâs Hummingbird, Black-chinned Hummingbird, and Calliope Hummingbird) as birds track poleward shifts in habitat ranges. 12 Current habitat occupancy for Annaâs Hummingbird includes South-eastern Vancouver Island, the southern Gulf Islands, and the Lower Fraser Valley, with some evidence of breeding in the Similkameen watershed in Interior B.C (Moran & Fraser, 2015a). As a primarily coastal breeding species with a preference for low-elevations and urban environments (Clark & Russell, 2012a; Moran & Fraser, 2015a), I hypothesize that Annaâs Hummingbird breeding ranges will expand in the following areas: northward following the East coast of Vancouver Island, and northward along the Mainland coast into low-elevation valleys within the Coast and Mountains Ecoprovince. Breeding range contractions are hypothesized to occur in Interior B.C. due to this region likely becoming increasingly dry with climate change, which is supported by limited evidence for breeding potential in this area (Moran & Fraser, 2015a). Black-chinned Hummingbird populations are currently centred in the southern Interior B.C., in regions such as the Thompson-Okanagan, Similkameen Valley, Creston Valley, Shuswap Lake and Southern Rocky Mountain Trench (Moran & Fraser, 2015b). As climate conditions warm across B.C., I hypothesize that Black-chinned Hummingbirds will experience northward range expansions into the Central Interior Ecoprovince, following northward movement of the Interior dry-zone and its associated vegetation. Furthermore, as this species exhibits a wider range of habitat occupancy in its southwestern breeding range in the United States, I hypothesize that warming climatic conditions in B.C. will allow for the Black-chinned Hummingbird to occupy a wider array of low-elevation habitats in its northern breeding limit, such as riparian areas, open woodlands, ponderosa-pine forests, and urban areas (Clark & Russell, 2012b; Moran & Fraser, 2015b). However, as this species has historically displayed preference for dry habitat conditions in their northern range limit (Clark & Russell, 2012b), 13 Black-chinned Hummingbirds are unlikely to occupy coastal areas of B.C. in future predictive models. Current habitat occupancy in B.C. for the Calliope Hummingbird includes much of the Southern Interior Ecoprovince, along with the Kootenay Interior Valley, Columbia, and a restricted breeding range in the Bulkley Valley and northern Central Interior Ecoprovince (Moran & Fraser, 2015c). Similar to the Black-chinned Hummingbird, Calliope Hummingbirds are very rarely found on Vancouver Island or the coastal lowlands of the Coast and Mountain Ecoprovince (Moran & Fraser, 2015c). As this species frequently occupies shrubland and open mixed conifer forest habitat (Clark & Russell, 2012c), I hypothesize that Calliope Hummingbird populations will continue to expand within the Southern Interior and expand northward to occupy the Central Interior. Based on possible and probable breeding evidence records (2008-2012), the breeding range of this species may move as far north as the Boreal Plains Ecoprovince (Moran & Fraser, 2015c). Rufous Hummingbirds occupy a wide variety of habitats, as displayed in their broad geographical range across Central and Southern B.C (Clark & Russell, 2012d). This species is also known to exhibit complex movement patterns, whereby local population distributions are in constant flux during the breeding season (Moran & Fraser, 2015d). This movement can present challenges in predicting range shifts and habitat occupancy. Population abundance for Rufous Hummingbirds is currently most concentrated in the Georgia Depression and Coast and Mountains Ecoprovince (Moran & Fraser, 2015d). In accordance with northern range expansions observed since 1990 (Moran & Fraser, 2015d), I hypothesize that this species will continue to expand northward into areas such as Prince Rupert, Mount Edziza Provincial Park, and Great Glacier Provincial Park, areas that had been identified for possible and probable breeding habitat 14 based on data collected in a 2008-2012 survey (Moran & Fraser, 2015d). Similar to Annaâs Hummingbird, I hypothesize that Rufous Hummingbirds will exhibit breeding range contraction in dry, warmer areas of Southern B.C., such as the Thompson-Okanagan in Interior B.C. Regarding interspecific habitat occupancy, I hypothesize a greater degree of overlap among Annaâs Hummingbird and Rufous Hummingbird breeding ranges in B.C. under future climate change scenarios. Annaâs Hummingbirds are likely to continue to expand northward and encroach further into habitat occupied by Rufous Hummingbirds, such as in the northeastern Vancouver Island and along the Mainland coast. Black-chinned and Calliope Hummingbirds may also share more habitat with Rufous Hummingbirds in the Central Interior, presenting new concerns for how these species will interact and establish interspecific dominance hierarchies on their breeding grounds (Cotton, 1998). Figure 1. Terrestrial protected areas in British Columbia (UNEP-WCMC & IUCN, 2021). 15 Chapter 2: Methods 2.1 Study area The study took place within the province of British Columbia (B.C.), Canada. British Columbia is one of the largest provinces in Canada, with a total land area of approximately 945,000 km2, and most mountainous, with 75% of land measuring over 1,000 m above sea level (Martin et al., 2021). The province is diverse in both climate and topography, dominated by four characteristic mountain regions: insular (Vancouver Island, Haida Gwaii), eastern (Canadian Rockies), interior (Cassiar Mountains, Omineca Mountains), and western (Cascade Mountains, Coast Mountains) (Scridel et al., 2021). British Columbia is composed of 14 unique Biogeoclimatic Ecosystem Classification (BEC) zones (Figure 2) and 10 terrestrial Ecoprovinces (Figure 3), owing to its large latitudinal (48°â60° N) and longitudinal (120°â140° W) range, maritime influence, and mountainous topography (Cannings & Cannings, 2015). British Columbiaâs climate has undergone significant change over the past century, with average annual precipitation having increased by 12% (1900-2013) and average annual temperature having increased by 1.4°C (1900-2013) (B.C. Ministry of Environment, 2016). Average annual maximum (+0.7°C) and minimum (+2.0°C) temperatures have also increased over the past century (B.C. Ministry of Environment, 2016). 16 Figure 2. Biogeoclimatic Ecosystem Classification (BEC) zones of British Columbia (UBC Centre for Forest Conservation Genetics, 2023). Figure 3. Ecoprovinces of British Columbia (Demarchi, 1996). 17 2.2 Hummingbird data Annaâs, Black-chinned, Calliope, and Rufous Hummingbirds are monitored across B.C. by the citizen science program eBird. Citizen science (CS) programs have gained popularity for use in research due to their ability to enable large-scale data collection across wide geographic areas and time periods (Conrad & Hilchey, 2011). While CS programs rely on the contributions of volunteers, who may possess varying levels of expertise and training in data collection, typically CS programs are capable of maintaining a high degree of data quality through employing vigorous automated data filters and teams of experts to review and validate data submissions (Conrad & Hilchey, 2011). The eBird database, maintained by the Cornell Lab of Ornithology, is a citizen science program containing weekly estimates of relative abundance throughout the year for all species. Due to its ease of use, volume of data and high degree of data integrity, eBird is one of the largest and most widely used citizen science databases for analyzing avian distributions and population trends (Horns et al., 2018; Callaghan & Gawlik, 2015). To model potential current and future distribution of the four hummingbird species, I used eBird presence-only data compiled from georeferenced observations in B.C. between 1981 and 2020 during the peak breeding period for all species. These dates were May 1-June 30 for the three migratory species (Rufous, Black-chinned, and Calliope Hummingbirds) and March 1 â April 30 for the resident Annaâs Hummingbird. I considered stationary and travelling checklist observations with a spatial accuracy of <2km to capture a resolution that provided a balanced compromise among the need for high accuracy, sample size limitations, a large spatial extent and the climatic and habitat data resolution. To control for sample selection bias, duplicate records were manually removed from the dataset where multiple observations occurred within the same spatial unit, even if they occurred at different times (1km2 grid resolution). 18 The final dataset included observations for 1,596 unique locations for Annaâs, 339 for Black-chinned, 837 for Calliope, and 4,030 for Rufous Hummingbirds. 2.3 Predictor variables To assess current habitat suitability for hummingbirds, I obtained a set of variables describing bioclimatic conditions, ecosystem type, topography, and urban land cover (Table 1). Bioclimatic variables were obtained from the CHELSA database (Karger et al., 2017, Table A1) representing mean temperature and precipitation patterns from 1979 to 2013. To reduce potential multicollinearity, I selected a priori three temperature and one precipitation variable considered to be ecologically and biologically meaningful for hummingbirds. Annual mean temperature (BIO1), temperature annual range (BIO7), and precipitation of warmest quarter (BIO18) were considered representative of general climate patterns and likely to influence the structure of pollinator networks and availability of vegetation on the breeding grounds (Leimberger et al., 2022; Brambilla et al., 2020; Abrahamczyk & Kessler, 2015). Mean temperature of warmest quarter (BIO10) was hypothesized to influence specialized relationships between hummingbirds and their food plants (Leimberger et al., 2022). For habitat predictors, data were available between 2001 and 2009 for the Biogeoclimatic Ecosystem Classification (hereinafter âBEC zonesâ, Table A1), which classes B.C. into 14 zones representing natural vegetation in the province based on local climate and geology (Pojar et al., 1987). Topographic variables including slope, topographic position index, and average summer solar radiation for B.C. were obtained from a previous species distribution modelling study (Scridel et al., 2021) which used a 50m digital elevation model (U.S. Geological Survey; https://earthexplorer.usgs. gov/) via Grass 19 7.04 (Neteler et al., 2012, Tables A1). Urban land cover data were obtained from the Copernicus Land Monitoring Service using PROBA-V satellite observations of global built-up land cover at 100m spatial resolution (Buchhorn et al., 2020). To model future habitat suitability, I used projections of future BEC zones into the 2080s (2071-2100), as well as the climate scenario model ACCESS1.0 (available from the CHELSA data base) which emulates the representative concentration pathway (RCP) +8.5 W/m2 scenario averaged across the years 2061â2081. RCP +8.5 W/m2 represents a high baseline emissions scenario, operating under the assumption that global emissions will continue to rise throughout the twenty-first century (Riahi et al., 2011). I selected a pessimistic (i.e. extreme) scenario for future distribution modelling as: (a) it best matches the future projections of the target BEC zones (2080s: 2071â2100), which average projections across 20 different emission scenarios producing an overall emission scenario between RCP +6.0 and +8.5 W/m2 (Wang et al., 2012), (b) observed emission from 2005 to 2020 matches the RCP +8.5 W/m2 (Peters et al., 2013) and the probability of realizing this scenario during the remainder of the century is great enough to warrant its continued use (Schwalm et al., 2020), and (c) for conservation purposes, using intermediate scenarios would lead to intermediate changes. Thus, applying a pessimistic emissions scenario within a set of realistic predictions enables us to test for the potential effectiveness of protected areas under more extreme conditions. Therefore, I attempted to accounted for both direct (i.e. climate) and indirect (i.e. climate-induced habitat alterations) impacts of climate change on the potential future distribution of Annaâs, Black-chinned, Calliope, and Rufous hummingbirds (hereinafter defined as the 2080s). 20 2.4 Protected areas data I used a standardized layer provided by the World Database on Protected Areas (UNEP-WCMC & IUCN, 2023) to estimate the extent of current and future protected area coverage for hummingbirds in B.C. I considered all terrestrial PAs in B.C. (n = 1,035) holding a national (IUCN category I to VI; n = 1,029 PAs) or international designation (n = 6). These terrestrial PAs cover a total of 146,069.3 km2 (approx. 15% of terrestrial land base of B.C.). 2.5 Data analysis I produced correlative species distribution models using a maximum entropy approach (Maxent version 3.4.4; Phillips & Dudik, 2008) through R package âSDMtuneâ (version: 1.1.5; Vignali et al., 2020) and dependent packages (i.e. âdismoâ version 1.3-8, Hijmans et al., 2020; âENMevalâ version: 2.0.3, Muscarella et al., 2014). Maxent is widely used for broad-scale species distribution modelling (Bradie & Leung, 2017; Elith et al., 2011) and is relatively insensitive to small sample sizes (Phillips et al., 2006). Environmental suitability was estimated for each species as a function of climate, habitat, and topography variables in a gridded domain at ~1 km resolution, which would likely cover the territories of hummingbirds detected in eBird surveys (Wilson et al., 2017). For Maxent models, the method of background point selection is critical to reducing potential effects of sampling effort bias. Background points were produced for each species to characterize environments within the study region that could be of potential suitability to hummingbirds (Phillips et al., 2009). Limited sample locations for Calliope Hummingbird (837 presence locations) and Black-chinned Hummingbird (339 presence locations) resulted in insufficient numbers of background points (<50,000). Thus, presence 21 locations across all study species were unified to produce a total of 5,131 presence locations (after removing duplicate points). From here, I applied a buffer radius of 4km between hummingbird presence points which reflected a balance between selecting enough points for this process and ensuring placement of background points were within reasonable proximity to the observations (Scridel et al., 2021). This increases the certainty that background points were likely accessible to observers and within the area of occurrence for each species. This process resulted in the creation of 50,000 background points for each of the four hummingbird species. For each species, occurrence data were partitioned into two groups (testing and training) using the âcheckerboard1â function in the package âENMevalâ (version: 2.0.3; Muscarella et al., 2014) with each of the checkerboard units aggregating four original 1 km2 cells. In respect to B.C.âs broad environmental gradients and high heterogeneity of landscapes, topography, and climate, such a checkerboard subdivision at a rather fine scale presents a compromise between each data partition being representative, and spatial independence between the testing and training data. Only linear and quadratic features were considered when producing SDMs. I reduced model complexity and collinearity by applying two functions (âvarSelâ and âreduceVarâ) within âSDMtuneâ (Vignali et al., 2020) to remove highly correlated variables (Pearsonâs R > 0.7) and variables with low importance (contribution <1%). After this, I tested different values of the regularization multiplier, training each model with 50 different values between 0.1 and 5 at 0.1 incremental steps. I selected the regularization multiplier resulting in the model with the highest test True Skill Statistic (TSS; Allouche et al., 2006) among those with a TSS difference (training â testing) lower than ±0.05. A leave-one-out jackknife procedure was applied to assess relative importance of predictor variables in the final model (Scridel et al., 2021). This was done by omitting each 22 variable in turn and calculating the difference in TSS (Scridel et al., 2021). Model accuracy was assessed by comparing the area under the curve (AUC) of a receiver operating characteristic (ROC) and via TSS values estimated from the evaluation datasets. Models with AUC <0.60 or TSS<0.10 were considered no better than random. The final logistic (cloglog; complementary log-log) output of the model was then reclassified into a binary occurrence map using the maximum training sensitivity plus specificity (MTSS) threshold (Liu et al., 2013; Scridel et al., 2021). To assess efficacy of protected areas (PAs) in conserving hummingbird species under a changing climate, I compared the absolute and proportional range sizes of suitable hummingbird habitat inside and outside PAs for both current and future scenarios. Species range overlap was assessed through overlaying binary occurrence rasters. 23 Table 1. Climate, topography, and habitat variables selected in the final models for each species (see Table A1 for full list of variables), with relative variable importance (%) and standard deviation (in brackets). Variable type Variable Annaâs Hummingbird Black-chinned Hummingbird Calliope Hummingbird Rufous Hummingbird Climate Mean temperature of warmest quarter (BIO10) 54.6 (0.015) + 37.8 (0.003) + Climate Temperature annual range (BIO7) 29.0 (0.003) +/- 15.4 (0.004) + 47.2 (0.005) + 23.0 (0.004) - Climate Annual mean temperature (BIO1) 52.9 (0.004) + 29.4 (0.003) + Climate Precipitation of warmest quarter (BIO18) 9.9 (0.008) - 12.2 (0.006) - Topographic Solar radiation 6.8 (0.005) + Topography Slope 15.0 (0.004) +/- 1.8 (0.003) +/- 5.7 (0.003) +/- 20.6 (0.004) - Topography Topographic position index 9.4 (0.009) +/- 15.6 (0.003) +/- Topography Urban land cover 1.7 + Habitat Coastal Douglas fir (CDF) 1.5 + 1.9 + Habitat Coastal Western Hemlock (CWH) 3.0 (0.003) - 1.3 (0.001) - Habitat Ponderosa Pine (PP) 1.1 (0.001) + 2.1 (0.001) + Habitat Montane Spruce (MS) 1.0 (0.001) + Sum of all Climate variables 81.9 (0.007) 79.9 (0.027) 88.8 (0.014) 60.8 (0.007) Sum of all Topographic variables 15.0 (0.004) 18.0 (0.017) 5.7 (0.003) 36.2 (0.007) Sum of all Habitat variables 3.2 2.1 (0.002) 5.1 (0.004) 3.2 (0.001) Note: The shape of the relationship is reported as: + (positive), - (negative), +/- (downward quadratic). Dominant variables selected in the model are bolded. Plots of species climate, environment, and topographic relationships are shown in Figures A1-A4 (Appendix). 24 Chapter 3: Results 3.1. Model performance For all four species, values of model performance over the test dataset ranged from 0.77 to 0.96 AUC and 0.42 to 0.85 TSS. This confirmed that models were capable of predict species distributions with high accuracy, even for species containing smaller sample sizes (e.g. Black-chinned Hummingbird: n = 339). Omission rates over the test dataset were close to expected values, indicating good prediction accuracy (Maximum training sensitivity threshold omission rates = 0.05-0.19). Projected current distributions (1981-2020) matched well with known species distributions and were in agreement with known ecology of hummingbirds (Clark and Russell, 2020; Baltosser and Russell, 2020; Calder and Calder, 2020; Healy and Calder, 2020). 3.2 Importance of predictor variables Temperature related variables were the strongest predictors for all species followed by precipitation and topography, while ecosystem variables were relatively weak predictors. My results showed that climate-related variables were the strongest predictors of distributions for all species with total permutation importance ranging from 60 to 89% across the four species (Table 1). Topographic variables contained variable importance ranging from approximately 6% permutation importance for Calliope Hummingbird, to 36% for Rufous Hummingbird (Table 1). Variables related to urban land cover and habitat type were of relatively little importance for all species (<5% permutation importance; Table 1). 25 For Annaâs Hummingbird, two temperature-related climate variables comprised a total 81.9% of permutation importance, with annual mean temperature and temperature annual range having the highest importance (Table 1). Annaâs Hummingbird probability of occurrence had a positive relationship with annual mean temperature, urban land cover, and Coastal Douglas fir, and a downward quadratic relationship with both slope and temperature annual range (Table 1). For Black-chinned Hummingbird, two temperature-related climate variables comprised a total of 70% of permutation importance, with mean temperature of warmest quarter and temperature annual range having the highest importance (Table 1). Black-chinned Hummingbird probability of occurrence had a positive relationship with mean temperature of warmest quarter, temperature annual range, solar radiation, Ponderosa Pine and Montane Spruce, and a downward quadratic relationship with slope and topographic position index (Table 1). Probability of occurrence had a negative relationship with precipitation of warmest quarter (Table 1). For Calliope Hummingbird, two temperature-related climate variables comprised a total of 75.1% permutation importance, with temperature annual range and annual mean temperature having the highest importance (Table 1). Calliope Hummingbird probability of occurrence had a positive relationship with temperature annual range, annual mean temperature, and Ponderosa Pine, and a downward quadratic relationship with slope (Table 1). Probability of occurrence had a negative relationship with precipitation of warmest quarter and Coastal Western Hemlock (Table 1). For Rufous Hummingbird, two temperature-related climate variables comprised a total of 60.8% permutation importance. Mean temperature of warmest quarter was the most important variable similar to the Black-chinned Hummingbird, while temperature annual range had the second highest importance (comparable importance to Annaâs Hummingbird; (Table 1). Rufous 26 Hummingbird probability of occurrence had a positive relationship with mean temperature of warmest quarter, Coastal Douglas fir and urban, and a downward quadratic relationship with topographic position index (Table 1). Probability of occurrence had a negative relationship with both temperature annual range, slope, and Coastal Western Hemlock (Table 1). 3.3 Future range expansion Across all four species, northward range expansions were projected under the RCP +8.5 W/m2 climate scenario (Figures 4-7; Table 2). Range expansions generally followed speciesâ topographic preferences, with coastal species such as Annaâs Hummingbird projected to expand further north in the Coast and Mountains Ecoprovince, while Interior species such as Black-chinned and Calliope Hummingbirds were projected to expand further north into Central Interior B.C. Generalist species such as Rufous Hummingbird were projected to expand northward across a broad geographical range (Figures 4-7). Range expansion into new breeding areas such as the Boreal Plains and Taiga Plains Ecoprovinces in northeastern B.C. was projected for Black-chinned, Calliope and Rufous Hummingbirds. Under an RCP +8.5 W/m2 climate scenario, Annaâs Hummingbird ranges were projected to expand northward along the mainland coast and the East coast of Vancouver Island, encroaching further into the Coast and Mountains Ecoprovince. Future occupancy was projected to expand into currently unoccupied areas such as along the northeastern coast of the Haida Gwaii archipelago and northeastern coast of Vancouver Island. Higher occupancy was projected within the current breeding range such as in Southeastern Vancouver Island, the southern Gulf Islands, and sections of the Lower Fraser Valley. Other areas such as Interior B.C. remained 27 unoccupied under the RCP +8.5 W/m2 climate scenario, suggesting this area would likely continue to have low value as breeding habitat to Annaâs Hummingbird. Black-chinned Hummingbird ranges were projected to expand northward under an RCP +8.5 W/m2 climate scenario into Central Interior B.C. and northeast into areas such as the Sub-Boreal Interior and Boreal Plains Ecoprovinces. Black-chinned Hummingbird future occupancy was seen to further expand in current breeding areas such as Southern Interior B.C. and the Ponderosa Pine BEC zone. As hypothesized, this species did not display range expansions in coastal areas such as the Coast and Mountains Ecoprovince, likely due to the probability of occurrence exhibiting a negative relationship with precipitation of warmest quarter (millimeters), a negative relationship with slope, and a downward quadratic relationship with topographic position index. Similar to Black-chinned Hummingbirds, Calliope Hummingbirds were projected to expand northward under an RCP +8.5 W/m2 climate scenario into Central Interior B.C. and northeastern areas of the province, such as the Sub-Boreal Interior and Boreal Plains Ecoprovinces. Range expansion was also projected to occur in current breeding areas such as the Southern Interior and the Ponderosa Pine BEC zone. As hypothesized, this species did not display range expansions in coastal areas such as the Coast and Mountains Ecoprovince, likely due to a negative relationship with precipitation of warmest quarter (millimeters), a negative relationship with slope, and a negative relationship with Coastal Western Hemlock. Rufous Hummingbird ranges were projected to expand northward into the Central Interior and Sub-Boreal Interior Ecoprovinces under an RCP +8.5 W/m2 climate scenario. As well, future range expansion was projected to occur in northeastern areas of B.C. such as the Boreal Plains and Taiga Plains Ecoprovinces, along with a few smaller, scattered sections of the 28 Northern Boreal Mountains Ecoprovince (northwest). Future occupancy on Vancouver Island and the Haida Gwaii archipelago remained relatively unchanged from current range occupancy. Future occupancy was projected to further expand in current breeding areas such as the Southern Interior. 3.4 Changes in protected area coverage Absolute protected area coverage was projected to increase for all four hummingbird species following range expansions, with absolute protected area coverage ranging from +6,378 km2 for Annaâs Hummingbird to +31,619 km2 for Rufous Hummingbird (Table 2). Relative protected area coverage was projected to increase for three species of hummingbirds (Calliope: +1.6%, Black-chinned: +2%, Annaâs: +10.8%) following range expansion. The Rufous Hummingbird was the only species that showed projected declines for relative protected area coverage (-3.8%) following range expansion (Table 2). The current breeding range of Annaâs Hummingbird was estimated at 7,599 km2, of which 372 km2 (4.9%) fell within the boundaries of protected areas (Figure 8). The projected range expansion by 2080 was predicted to cover 43,074 km2 of which 6,750 km2 (15.7%) fell within the boundaries of protected areas (Figure 8). For the Black-chinned Hummingbird, current breeding range was estimated at 24,319 km2, of which 1,283 km2 (5.3%) fell within the boundaries of protected areas (Figure 9). The projected range expansion by 2080 was predicted to cover 304,421 km2, of which 22,235 km2 (7.3%) fell within the boundaries of protected areas (Figure 9). 29 The current breeding range for Calliope Hummingbird was estimated at 39,789 km2, of which 2,364 km2 (5.9%) fell within the boundaries of protected areas (Figure 10). The projected range expansion by 2080 was predicted to cover 351,528 km2, of which 26,193 km2 (7.5%) fell within the boundaries of protected areas (Figure 10). Finally, Rufous Hummingbirdâs current breeding range was estimated at 86,457 km2, of which 11,897 km2 (13.7%) fell within the boundaries of protected areas (Figure 11). The projected range expansion by 2080 was predicted to cover 440,999 km2, of which 43,516 km2 (9.9%) fell within the boundaries of protected areas (Figure 11). 3.5 Changes in range overlap Range overlap was projected to increase across all species combinations following range expansions, with gains in overlapping territory ranging from +604 km2 between Annaâs and Black-chinned Hummingbirds to +267,478 km2 between Calliope and Rufous Hummingbirds (Table 3). Increases in future range overlap between coastal species (Annaâs Hummingbird) and interior species (Black-chinned and Calliope Hummingbirds) were moderate in comparison to projected overlap increases for generalist (Rufous Hummingbird) and interior species (Black-chinned and Calliope Hummingbirds) (Table 3). 30 Figure 4. Modelled potential breeding range distributions of Anna's Hummingbird in B.C. for current (1981-2020) and future (2080s) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2 A) Suitability according to the maximum training sensitivity plus specificity (MTSS) threshold is depicted in green. B) Suitability gradient ranging from 0.0 to 1.0. 31 Figure 5. Modelled potential breeding range distributions of Black-chinned Hummingbird in B.C. for current (1981-2020) and future (2080s) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2 A) Suitability according to the maximum training sensitivity plus specificity (MTSS) threshold is depicted in green. B) Suitability gradient ranging from 0.0 to 1.0. 32 Figure 6. Modelled potential breeding range distributions of Calliope Hummingbird in B.C. for current (1981-2020) and future (2080s) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2 A) Suitability according to the maximum training sensitivity plus specificity (MTSS) threshold is depicted in green. B) Suitability gradient ranging from 0.0 to 1.0. 33 Figure 7. Modelled potential breeding range distributions of Rufous Hummingbird in B.C. for current (1981-2020) and future (2080s) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2 A) Suitability according to the maximum training sensitivity plus specificity (MTSS) threshold is depicted in green. B) Suitability gradient ranging from 0.0 to 1.0. 34 Figure 8. Anna's Hummingbird occupancy in B.C. protected areas under current (1981-2020) and future (2080s: 2071-2100) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2. 35 Figure 9. Black-chinned Hummingbird occupancy in B.C. protected areas under current (1981-2020) and future (2080s: 2071-2100) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2. 36 Figure 10. Calliope Hummingbird occupancy in B.C. protected areas under current (1981-2020) and future (2080s: 2071-2100) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2. 37 Figure 11. Rufous Hummingbird occupancy in B.C. protected areas under current (1981-2020) and future (2080s: 2071-2100) scenarios according to habitat and climate projections (ACCESS 1.0) for RCP +8.5 W/m2. 38 Figure 12. Current (1981-2020) range overlap for Anna's, Black-chinned, Calliope, and Rufous Hummingbirds. 39 Figure 13. Future (2080s) range overlap for Anna's, Black-chinned, Calliope, and Rufous Hummingbirds. 40 Table 2. Range size (km2) and proportion (%) of potential suitable habitat for hummingbirds across B.C. and within protected (PAs) and in unprotected areas (UPAs) under current and future scenarios (2080s) according to ACCESS 1.0 climatic projections. Note: Bold values represent proportional changes (%) of suitable habitat inside and outside PAs. Table 3. Range overlap (km2) for hummingbirds across B.C. under current and future scenarios (2080s) according to ACCESS 1.0 climatic projections. Species Current Range Overlap 2080s Range Overlap Change in Range Overlap Annaâs & Black-chinned Hummingbirds 4 km2 604 km2 +600 km2 Annaâs & Calliope Hummingbirds 10 km2 1,635 km2 +1,625 km2 Annaâs & Rufous Hummingbirds 7,594 km2 41,616 km2 +34,022 km2 Black-chinned & Calliope Hummingbirds 22,433 km2 286,628 km2 +264,195 km2 Black-chinned & Rufous Hummingbird 18,016 km2 263,513 km2 +245,497 km2 Calliope & Rufous Hummingbirds 24,265 km2 291,743 km2 +267,478 km2 Species Current Suitable range across province Current Suitable range in PAs Current Suitable range outside PAs 2080s Suitable range across province 2080s Suitable range in PAs (ACCESS 1.0) 2080s Suitable range outside PAs (ACCESS 1.0) Change in suitable range across province Absolute change in range size in PAs Relative change in range size in PAs Annaâs Hummingbird 7,599 km2 372 km2 (4.9%) 7,227 km2 (95.1%) 43,074 km2 6,750 km2 (15.7%) 36,324 km2 (84.3%) +35,475 km2 +6,378 km2 +10.8% Black-chinned Hummingbird 24,319 km2 1,283 km2 (5.3%) 23,036 km2 (94.7%) 304,421 km2 22,235 km2 (7.3%) 282,186 km2 (92.7%) +280,102 km2 +20,952 km2 +2% Calliope Hummingbird 39,789 km2 2,364 km2 (5.9%) 37,425 km2 (94.1%) 351,528 km2 26,193 km2 (7.5%) 325,335 km2 (92.5%) +311,739 km2 +23,829 km2 +1.6% Rufous Hummingbird 86,457 km2 11,897 km2 (13.7%) 74,560 km2 (86.3%) 440,999 km2 43,516 km2 (9.9%) 397,483 km2 (90.1%) +354,542 km2 +31,619 km2 -3.8% 41 Chapter 4: Discussion 4.1 Overview My study provides the first comparison of current and future hummingbird breeding range distributions under an RCP +8.5 W/m2 climate change scenario in British Columbia. My most important findings were (1) Future hummingbird ranges were predicted to expand northward under a pessimistic climate scenario (RCP +8.5 W/m2) (2) Protected area coverage was predicted to increase for Annaâs, Black-chinned, and Calliope Hummingbirds relative to total suitable habitat, except for Rufous Hummingbirds (relative decrease -3.8%), and (3) Greater species distribution overlap is predicted to occur as a result of range expansion. 4.2 Northward range expansion As climate change continues to alter temperature and precipitation regimes across the globe, range shifts may become increasingly necessary to species survival (Chen et al., 2011). By the 2080s, under an RCP +8.5 W/m2 climate scenario, British Columbiaâs annual mean temperature is predicted to increase by 3.96°C (SE = 0.21), while annual mean precipitation is predicted to increase modestly by 60.63mm (SE = 20.05) (Scridel et al., 2021). My findings showed that climate variables were stronger predictors of distribution than topographic and habitat variables, supporting a wider belief across ecological studies that climatic tolerance is the primary factor determining the constraints to speciesâ geographic ranges (Coristine & Kerr, 2015; Parmesan, 1996). More specifically, this theory suggests that speciesâ ranges will shift under climate change to remain within similar climate envelopes, and that ranges will expand 42 into new areas as they become climatically suitable (Coristine & Kerr, 2015). In my study, temperature variables contained higher modelling importance in comparison to precipitation variables, suggesting that temperature may act as the primary force influencing future range shifts for hummingbirds. In British Columbia, hummingbird breeding ranges are predicted to expand northward as species track shifting habitats induced by climate warming and environmental change. Northward range expansions were observed in all four species of hummingbird (Figure 4-7) as predicted. My findings are consistent with trends observed in other North American avifauna whose breeding ranges are shifting poleward in response to warming temperatures (Hovick et al., 2016, Melles et al., 2011; Zuckerberg et al., 2009; Bateman et al., 2016), such as the Hooded Warbler (Setophaga citrina) and Blackpoll Warbler (Setophaga striata) in Eastern Canada and the U.S (Melles et al., 2011; Ralston & Kirchman, 2013). Other taxa in North America such as butterflies (Lemoine, 2015) small mammals (Roy-Dufresne et al., 2013; True et al., 2021) and plants (Iverson & McKenzie, 2013) are also reported to be expanding their ranges poleward. Annaâs Hummingbirds have undergone rapid range expansion at their northern range limit over the 20th century, which has further intensified within the past decade (Battey, 2019; English et al., 2021). At the edge of their northern range in Canada, population growth increased at a rate of 20% per year (English et al., 2021). Annaâs Hummingbirds are commonly associated with urban areas (Clark & Russell, 2020), where supplementary feeding provided by introduced plants and nectar feeders have facilitated ecological release (Zimmerman, 1973; Adams et al., 2023). This has allowed Annaâs Hummingbird to expand their breeding range northward and persist in human-dominated landscapes over colder months, allowing for overwintering in areas such as the lower Fraser Valley, southern Vancouver Island, and the South Okanagan (Greig et 43 al., 2017). I predict that the species will expand its range further north as climates continue to warm. It is important to note that my models displayed a downward quadratic relationship for Annaâs predicted occurrence with temperature annual range. As Annaâs Hummingbirds are non-migrant residents in B.C., this relationship indicates that extreme temperatures beyond the speciesâ optimal annual range of 20°C likely represent a limit to their range expansion, especially eastward towards the colder interior portions of the province (Jeong et al., 2016a,b). As expected, my models predicted breeding ranges for Black-chinned and Calliope Hummingbird to expand northward into the Central Interior under an RCP +8.5 W/m2 climate scenario. Black-chinned and Calliope Hummingbirds are associated with hot, dry riparian habitats and ponderosa pine (Pinus ponderosa) stands, an ecosystem type that may become more widespread within Central Interior B.C. and along northern sections of the Fraser River as drought events increase in frequency and severity within the province (Baltosser & Russell, 2020; Calder & Calder, 2020). British Columbia is currently at the northern periphery of the range for these two species, but my results indicate how the province is likely to host a greater proportion of the breeding distributions for both species in the future. My results also suggest that other species that inhabit similar dry, interior ecosystems may similarly expand northward, although further studies would be needed to confirm that they respond to similar variables. For example, while hummingbirds may be capable of expanding their range further north into Central Interior B.C. under a warming climate, other Southern Interior species may face constraints that prevent northward range shifts. Such species may include the Western Rattlesnake (Crotalus oreganus), which is unlikely to disperse northwards at the same rate as Black-chinned and Calliope Hummingbirds due to mobility restrictions (Didiuk et al., 2004), and the Pallid bat (Antrozous pallidus) that experience topographic range constraints (being restricted 44 to elevations under 500m in B.C.) (Willis & Bast, 2000). While my study found climate variables to be the strongest predictors of hummingbird distribution and range expansion in B.C., there were differences in the importance of the climate variables among the study species (Table 1). Annaâs Hummingbird responded most strongly to annual mean temperature, whereas mean temperature of warmest quarter was most important for Black-chinned and Rufous Hummingbirds, and temperature annual range was most important for Calliope (Table 1). If these three climate variables exert different responses among the study species, it is possible that population responses could vary among hummingbird species. Rufous Hummingbirds are capable of tolerating cooler environmental conditions, reaching the northernmost latitude of any hummingbird in North America at 61°N (Healy & Calder, 2020). In their B.C. range, Rufous Hummingbirds occupy elevations ranging from sea level to 1,830m (Healy & Calder, 2020). While my model suggests a negative relationship between temperature annual range and predicted occupancy for Rufous Hummingbirds, a positive relationship with mean temperature of warmest quarter contained greater modelling importance. This result suggests that in their most northern breeding range, Rufous Hummingbirds may prefer more moderate climates which experience fewer annual fluctuations in temperature, such as the coastal regions of B.C warmed by Pacific Ocean currents. However, this does not limit the species from occupying drier, warmer regions of the province in its future range, such as the Interior and Central Interior, which are predicted to become increasingly suitable for Rufous Hummingbirds as average temperatures during the breeding season rise under climate change (Figure 11). One possible by-product of hummingbird range expansions could be rapid population growth (Battey, 2019). This may occur if limits to growth rates imposed by climate or 45 interspecific interactions are reduced in novel habitats (Battey, 2019). Although acclimation may be possible for hummingbird populations in the most northern limits of their predicted breeding range, some species may not occupy the full extent of predicted ranges in reality, due to other factors which pose limitations to geographic range extent such as migratory distance and seasonality of floral resources (Somveille et al., 2019). It is also important to consider that while my models predict significant breeding range expansions for all four species under climate change, rising temperatures may negatively impact hummingbirds if some species are unable to shift their ranges poleward fast enough to avoid incurring climate debts (Gaüzère & Devictor, 2021). The increase in frequency and intensity of extreme events associated with climate warming may limit the realized increases in predicted breeding range expansions (Pearse-Higgins & Martin, 2023). Further research is needed to fully understand the implications of range expansions for both hummingbirds and the ecosystems in which they inhabit. 4.3 Protected area (PA) coverage and effectiveness 86.3â95.1% of the predicted current breeding ranges of hummingbirds in B.C. occur outside of protected areas. Being unprotected, hummingbirds in these areas are more likely to be exposed to stressors such as unsustainable forestry, agriculture, and urbanization. This shortage in protected area coverage for hummingbirds in B.C. is in part due to an uneven geographical distribution of protected areas across the province. Protected areas are often situated in places undesirable for human settlement and cultivation, such as areas at high elevations or containing low ecological productivity (Wiens et al., 2011). This trend is seen in B.C., where many large networks of protected areas have been established along the Northern Boreal Mountains and Coast and Mountains Ecoprovinces in West and Northwestern B.C. (Figure 1,2). Gaps in 46 protected area coverage across the province are greatest in the Southern Interior, Central Interior, Taiga Plains, Boreal Plains, and urban areas (Figure 1,2). Within protected areas, breeding range coverage for hummingbirds was predicted to increase for three species (Annaâs, Black-chinned, Calliope) under the RCP +8.5 W/m2 climate scenario, with Annaâs Hummingbirds showing the largest increase in protected area coverage relative to their predicted suitable habitat across the province, from 4.9% (1981-2020) to 15.7% (2080s) (Table 2). Rufous Hummingbirds were the only species with predicted declines for protected area coverage relative to suitable habitat across the province (Table 2). However, this factor alone is unlikely to pose concern for this speciesâ conservation, as total breeding area within protected areas for Rufous Hummingbirds is predicted to increase by about 4-fold from the current amount (Table 2). As B.C. undergoes climate change, ecological conditions within protected areas will likely be altered due to rising temperatures and shifting precipitation regimes (Wiens et al., 2011). Such changes may prove to be beneficial for hummingbird conservation, as my models predict that under RCP +8.5 W/m2, suitable breeding habitat will not only expand into previously unoccupied PAs, but will also be maintained in currently occupied PAs. The hummingbirdâs ability to physically tolerate temperature extremes through passive and evaporative heat dissipation, as well as their ability to utilize thermal refugia and cooler microclimates through adaptive behaviour (Powers et al., 2017) may be examples of mechanisms allowing hummingbirds to persist and expand in B.C. PAs under future climate warming. 47 4.4 Implications of future range overlap Hummingbirds are highly territorial as they must secure and defend food sources and nesting territories from competing hummingbirds as well as other birds and floral competitors such as butterflies and bees. Due to their small size and high metabolism, hummingbirds consume frequent small meals of nectar and insects throughout the day to maximize net energy intake (Chen et al., 2014). Male hummingbirds establish feeding territories during the first weeks of the breeding season (Powers, 1987), although this process does not entirely eliminate the possibility of territory intrusion and overlap with competing species. Interspecific competition can have a strong influence on hummingbird community structure and pose limitations on species distribution and abundance (LópezâSegoviano et al., 2018). For example, species that rank lower in aggressive dominance hierarchies are placed at a disadvantage when interspecific competition forces birds to feed from lower-quality food sources or expend greater time and energy on foraging (Martin, 1988). Black-chinned, Calliope and Rufous Hummingbird breeding ranges overlap extensively in areas of South Interior and Central Interior B.C. (see Figure 12, Calder & Calder, 2020). By the 2080s, range overlap is predicted to further increase as these species expand northward to occupy new suitable habitat (Figure 13). Black-chinned and Calliope are some of the smallest species of breeding hummingbirds in North America, and as such are inferior competitors to the larger and more aggressive Rufous Hummingbird (Calder & Calder, 2020; Martin, 1988). Despite this, Black-chinned and Calliope Hummingbirds are capable of sharing feeding boundaries with limited apparent conflict (Calder & Calder, 2020). Calliope Hummingbirds have been shown to feed inconspicuously within Rufous Hummingbird territories when both species impart different feeding strategies. In a study by Martin (1988), Rufous Hummingbirds were 48 shown to select flowers at heights â¥ 30 cm above ground for feeding, while Calliope Hummingbirds selected lower hanging flowers (â¤ 20 cm above ground). Indeed, Calliope Hummingbirds are often thought to be sympatric with Rufous Hummingbirds over wide areas. Although my models predict extensive range expansion by the 2080s for Rufous Hummingbird populations in B.C., these previous findings on how they segregate in areas of overlap suggest that further overlap is unlikely to significantly impact the dynamics of their interspecific relations with Calliope and Black-chinned Hummingbirds. Due to Rufous Hummingbirds experiencing long-term population declines in Canada (English et al., 2021), their future breeding density may also become further reduced and dispersed across B.C., limiting the likelihood of increased interspecific competition with other species in the Interior (Government of Canada, 2022). Under the RCP +8.5 W/m2 climate scenario, Rufous Hummingbirds may see increased competition for food resources and nesting sites with Anna's Hummingbirds as my models predict their breeding range to encroach into Rufous breeding habitat along the mainland coast (Figure 13). Unlike Black-chinned and Calliope Hummingbirds, Annaâs Hummingbirds are non-migrant residents of B.C., and as such have the advantage to establish breeding territory before Rufous Hummingbirds arrive on the breeding grounds (Clark & Russell, 2020). Male Annaâs Hummingbirds are known to spend more time and energy defending breeding territory than they do defending feeding territory (Stiles, 1971). Rufous Hummingbirds have declined considerably in areas where Annaâs Hummingbirds are now common but it is not clear whether the expansion of Annaâs Hummingbird has led to reductions in Rufous Hummingbird populations or whether Annaâs are only replacing Rufous after the latter decline due to other factors, such as threats on the nonbreeding grounds (English et al. 2022). Nevertheless, increased competition with Annaâs 49 Hummingbirds may be an additional stressor for already declining populations of Rufous Hummingbirds. Closely related species can experience hybridization when secondary contact is facilitated by range expansion (Godwin et al., 2020). Hybridization has been observed between Annaâs, Black-chinned, Calliope, and Rufous Hummingbirds during the breeding season (eBird Library, 2023a-c). Although rare, hybridization may pose a threat to the long-term persistence of vulnerable species like Rufous Hummingbirds via outbreeding depression and assimilation (Levin, 2002). Hybrid species remain unprotected under the Canadian Species at Risk Act (SARA), regardless of the conservation status of their parent taxa (Government of Canada, 2023). 4.5 Caveats Fire regimes have an important role in shaping and maintaining habitat suitability for species by influencing landscape connectivity and composition (De Cáceres et al., 2013; Swan et al., 2021; Hoecker & Turner, 2022). Due to limitations imposed by modelling capacity, time constraints, and available data sources, I was unable to include fire regimes and forest disturbance as long-term land cover change variables in my models. Fire regimes has proven difficult to model over large time scales due to the complex relationship of fire with varying factors such as humidity, temperature, surrounding vegetation, and terrain (Parks et al., 2016). It is likely that climate warming already has and will continue to increase the severity, frequency, and duration of forest fires in B.C. (KirchmeierâYoung et al., 2019; Metsaranta et al., 2023). Recent studies of future fire risk in B.C. project annual burned area by 2070 to double those averaged between the period of 1950 to 2018 (Metsaranta et al., 2023). Such trends may 50 negatively impact hummingbirds by increasing habitat loss, degradation, and disruption to food resource availability. However, some species such as Black-chinned and Calliope Hummingbirds may benefit from altered fire regimes if it increases the availability of dry habitat that is of preference to these Interior species (Baltosser & Russell, 2020; Calder & Calder, 2020). Similarly, habitat generalists such as Rufous Hummingbirds may benefit from increased availability of secondary succession forest communities and openings created by fires (Healy and Calder, 2020). I acknowledge that forest disturbance and fire regimes could further have an effect on the predictive accuracy of my models, and suggest that these variables be included in future studies of hummingbird distribution and range expansion in B.C. To best match the time period of the climatic and habitat datasets, I modelled future hummingbird suitable habitat based on only one pessimistic climatic scenario. It is important to acknowledge the inherent uncertainty with downscaling regional climate models (RCMs) to produce future climate projections, such as ACCESS 1.0 RCP+8.5 W/m2, due to the complex and varying nature of Earthâs climate systems (Ahmadalipour et al., 2018). My predictions may overestimate range expansion for hummingbirds if current greenhouse gas emissions experience significant reductions below the RCP +8.5 W/m2 scenario. I predict that using a moderate climate scenario such as RCP +4.5 W/m2 would result in an intermediate distribution pattern between the current and future distribution forecast, with less pronounced range expansions in northern B.C compared to the RCP +8.5 W/m2 climate scenario. Developing predictive models of Rufous Hummingbird distribution proved challenging, as this species is known to nest in a wide array of habitats over broad geographical ranges, including coniferous forests, deciduous forests, riparian thickets, meadows, swamps, agricultural areas, and urban areas containing gardens and parks (Healy & Calder, 2020). Rufous 51 Hummingbirds are also considered to be generalists in their feeding strategy, making use of a wide array of flowering plants, insects (Healy & Calder, 2020), and even tree sap (Sutherland et al., 1982). Many aspects of this speciesâ life history remain poorly understood, such as their population dynamics, breeding ecology, and foraging patterns (Calder & Healy, 2020). Although not considered in this analysis, variables representing key vegetation for breeding hummingbirds, such as projected current and future distributions for Salmonberry (Rubus spectabilis) and Red-Flowering Currant (Ribes sanguineum) across B.C., may further improve the predictive accuracy of my models. Hummingbirds share close relationships with flowering plants, and migration timing for Rufous Hummingbird often coincides with the blooming period of important food resources (Moran & Fraser, 2015). As such, identifying changes in both spatial and temporal patterns for flowering plants in B.C. could provide valuable insights into future hummingbird distributions (McKinney et al., 2012). The Biogeoclimatic Ecosystem Classification (BEC) framework operates under the assumption that ecosystem types and composition will not undergo change (Pojar et al., 1987). However, it is possible that some of the current BEC zones may not possess future analogues in the 2080s, if climate change or other non-climate related processes cause ecosystems to change in composition, merge to produce novel ecosystem types, or disappear altogether. Finally, there are likely to be other underlying variables that affect hummingbird range expansion. A study on northward range shifts in a North American warbler (Melles et al., 2010) suggested that biotic processes, such as immigration and establishment, should be considered alongside abiotic factors when assessing species distribution under global change. However, the authors acknowledge that biotic processes are often difficult, if not impossible, to investigate in 52 broad multispecies studies, and are more suited to studies at the regional scale (Melles et al., 2010). 4.6 Conservation implications British Columbia will have a greater responsibility to conserving its populations of Annaâs, Black-chinned, Calliope and Rufous Hummingbirds as these species experience range expansion across the province. Although my predictions indicate some of their future range expansion will fall within the boundaries of protected areas, a larger degree of range expansion is predicted to occur outside these boundaries in non-protected areas. Therefore, I encourage conservation measures to be strengthened for hummingbird breeding and foraging habitat within, but especially outside, the boundaries of protected areas. One way this could be achieved would be through implementing habitat restoration programs and management regimes in non-protected areas, and encouraging communities and private landowners to partake in sustainable land use practices that promote biodiversity conservation (Monzón et al., 2011). Although the ranges for all four hummingbirds are predicted to expand under climate change, it is unclear whether food resources such as flowering plants will undergo similar shifts. This potential mismatch in breeding habitat and resource availability may deter hummingbirds from occupying novel suitable habitat at their northern breeding range limit, or present consequences such as increased interspecific competition for resources (Pearse-Higgins & Martin, 2023). The long-term impacts of increased interspecific range overlap and competition among hummingbird species on the breeding grounds similarly remain uncertain. For the Rufous Hummingbird, a species currently facing declines in Canada, heightened interspecific competition for breeding and feeding territory may negatively impact future reproductive output 53 and population viability for this species. Furthering our understanding of the interspecific dynamics between hummingbirds and potential climate-induced changes in vegetation important to these birds would help to better inform species conservation under a changing environment. 4.7 Future opportunities and studies I suggest that future studies of B.C. hummingbird distributions and range expansion incorporate variables of land use and land cover (LULC) change and provincial fire regimes in species distribution models to increase their predictive accuracy. This will be of particular importance as anthropogenic land use change and climate change alter natural disturbance regimes, increasing the frequency and severity of forest fires which exacerbate habitat loss (Park et al., 2016). Future changes to landcover in B.C. such as decreasing trends in total agricultural area (-7.9% from 2006-2011; Statistics Canada, 2016) may be advantageous to hummingbirds if agricultural lands are to be replaced by forest area. Similarly, changes to forest management practices and activities within B.C. are worthy of exploring in future studies, as they may affect the breeding range of species such as Rufous Hummingbird which are known to breed along forest edges and clearings (Healy & Calder, 2020). To reduce the likelihood of inaccurate climate change predictions within species distribution models (SDMs), hindcasting â which involves using historical records of species distribution to project past climate conditions â could be useful as a tool to validate and improve climate models (Crickenberger & Wethey, 2018). For example, species distribution data from the period 1980-1990 could be used to produce SDM projections for the year 2020 in order to assess how closely model projections match reality. 54 Other research avenues of ecological merit could include modelling range shifts in flowering plants and other food resources closely associated with hummingbirds (e.g. Rubus spectabilis and Ribes sanguineum), in addition to demographic studies of Annaâs, Black-chinned, Calliope and Rufous Hummingbirds that investigate factors affecting population growth, dispersal, and dynamics. This in turn could lead to the production of mechanistic models which incorporate both biological and ecological processes influencing species distribution (Mathewson et al., 2017). Such work would help to further inform effective conservation across the B.C. breeding range for these valued species. 55 Bibliography Abrahamczyk, S., Souto-Vilaros, D., McGuire, J. A., & Renner, S. S. (2015). 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(2009). Poleward shifts in breeding bird distributions in New York State. Global Change Biology, 15(8), 1866-1883. https://doi.org/10.1111/j.1365-2486.2009.01878.x 71 Appendix Table A1. List of climatic, habitat and topographic variables considered in the modelling procedure. Italicised* variables were removed prior to modelling as they showed correlation with final selected variables. Variable type Dataset source Variable name Climate CHELSA (V1.2) Annual mean temperature (Bio1) Climate CHELSA (V1.2) Temperature annual range (Bio7) Climate CHELSA (V1.2) Mean temperature of warmest quarter (Bio10) Climate CHELSA (V1.2) Precipitation of warmest quarter (Bio18) Climate* CHELSA (V1.2) Temperature seasonality Climate* CHELSA (V1.2) Max temperature of warmest month Climate* CHELSA (V1.2) Annual precipitation Climate* CHELSA (V1.2) Precipitation seasonality Habitat Pojar et al. 1987 Alpine Tundra (Boreal Altai Fescue Alpine, Coastal Mountain heather Alpine, & Interior Mountain heather Alpine) Habitat Pojar et al. 1987 Bunchgrass (BG) Habitat Pojar et al. 1987 Boreal White and Black Spruce (BWBS) Habitat Pojar et al. 1987 Coastal Douglas fir (CDF) Habitat Pojar et al. 1987 Coastal Western Hemlock (CWH) Habitat Pojar et al. 1987 Engelmann Spruce Subalpine Fir (ESSF) Habitat Pojar et al. 1987 Interior Cedar Hemlock (ICH) Habitat Pojar et al. 1987 Interior Douglas fir (IDF) Habitat Pojar et al. 1987 Mountain Hemlock (MH) Habitat Pojar et al. 1987 Montane Spruce (MS) Habitat Pojar et al. 1987 Ponderosa Pine (PP) Habitat Pojar et al. 1987 Sub-Boreal Pine Spruce (SBPS) Habitat Pojar et al. 1987 Sub-Boreal Spruce (SBS) Habitat Pojar et al. 1987 Spruce-Willow-Birch (SWB) Topography Computed in GRASS (Scridel et al., 2021). Slope (angle of inclination to the horizontal) Topography Computed in GRASS (Scridel et al., 2021). Solar radiation (summer) Calculated as global radiation on 21st June account the shadowing effect of the relief (slope and aspect). Topography Computed in GRASS (Scridel et al., 2021). Topographic position Index (difference in elevation between a central pixel and the mean of its surrounding 8 cells) Topography* Computed in GRASS (Scridel et al., 2021). Roughness 72 Figure A1. Most supported variables defining Annaâs Hummingbird distribution (temperature variables are expressed in °C/10, precipitation in mm, slope in degrees, topographic position index values above zero show locations that are higher than then average of immediately surrounding terrain). Permutation importance (highest to lowest): Annual mean temperature (52.9%), Temperature annual range (29.0%), Slope (15.0%), Urban land cover (1.7%), Coastal Douglas fir (1.5%). 73 Figure A2. Most supported variables defining Black-chinned Hummingbird distribution (temperature variables are expressed in C°/10, precipitation in mm, slope in degrees, topographic position index values above zero show locations that are higher than then average of immediately surrounding terrain). Permutation importance (highest to lowest): Mean temperature of warmest quarter (54.6%), Temperature annual range (15.4%), Precipitation of warmest quarter (9.9%), Topographic position index (9.4), Solar radiation (6.8%), Slope (1.8%), Ponderosa Pine (1.1%), Montane Spruce (1.0%). 74 Figure A3. Most supported variables defining Calliope Hummingbird distribution (temperature variables are expressed in C°/10, precipitation in mm, slope in degrees, topographic position index values above zero show locations that are higher than then average of immediately surrounding terrain). Permutation importance (highest to lowest): Temperature annual range (47.2%), Annual mean temperature (29.4%), Precipitation of warmest quarter (12.2%), Slope (5.7%), Coastal Western Hemlock (3.0%), Ponderosa Pine (2.1%). 75 Figure A4. Most Most supported variables defining Rufous Hummingbird distribution (temperature variables are expressed in C°/10, precipitation in mm, slope in degrees, topographic position index values above zero show locations that are higher than then average of immediately surrounding terrain). Permutation importance (highest to lowest): Mean temperature of warmest quarter (37.8%), Temperature annual range (23.0%), Slope (20.6%), Topographic position index (15.6%), Coastal Douglas fir (1.9%), Coastal Western Hemlock (1.3%).
Thesis/Dissertation
eng
Forestry
Vancouver : University of British Columbia Library
Attribution-NonCommercial-NoDerivatives 4.0 International
Graduate
Martin, Kathy (Katherine), 1949-
Wilson, Scott D.
Examining the influence of climate change on distribution shifts and the efficacy of protected areas for hummingbirds in British Columbia
Text