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• 1. [Article] Independent effects of habitat loss and fragmentation on pollination : tropical forest fragmentation alters hummingbird movements and pollination dynamics

  A growing body of work reveals that animal-mediated pollination is negatively affected by anthropogenic disturbance. Landscape-scale disturbance results in two often inter-related processes: (1) habitat ...

  Citation × Citation Citation Abstract Copy Title: Independent effects of habitat loss and fragmentation on pollination : tropical forest fragmentation alters hummingbird movements and pollination dynamics Author: Hadley, Adam S. A growing body of work reveals that animal-mediated pollination is negatively affected by anthropogenic disturbance. Landscape-scale disturbance results in two often inter-related processes: (1) habitat loss, and (2) disruptions of habitat configuration (i.e. fragmentation). Understanding the relative effects of such processes is critical in designing effective management strategies to limit pollination and pollinator decline. I reviewed existing published work from 1989 to 2009 and found that only six of 303 studies separated the effects of habitat loss from fragmentation. I provide a synthesis of the current landscape, behavioral, and pollination ecology literature in order to present preliminary multiple working hypotheses to explain how these two landscape processes might independently influence pollination dynamics (Chapter 2). Despite the potential importance of independent effects of habitat fragmentation, effects on pollination remain largely untested. Studies designed to disentangle the independent effects of habitat loss and fragmentation are essential for gaining insight into landscape-mediated pollination declines. I also found that the field of landscape pollination ecology could benefit from quantification of the matrix, landscape functional connectivity, and pollinator movement behavior. To test the hypothesis that pollinator movement can be influenced by landscape configuration, I translocated radio-tagged hummingbirds across agricultural and forested landscapes near Las Cruces, Costa Rica (Chapter 3). I found return paths were on average more direct in forested than in agricultural landscapes. In addition, movement paths chosen in agricultural landscapes were more forested than the most direct route suggesting that hummingbirds avoided crossing open areas when possible. To determine if differences in pollinator movement translated to differences in plant reproduction, I tested the relative importance of landscape composition versus configuration on the reproductive success of Heliconia tortuosa, a hummingbird-pollinated forest herb (Chapter 4). I used a stratified random sampling design to select sites across orthogonal gradients in patch size, amount of forest, and elevation. I tested four landscape change hypotheses (i.e., local, landscape composition, landscape fragmentation, and fragmentation threshold). I found that Heliconia reproduction supported both the local site and landscape fragmentation hypotheses. Seed set increased with increasing forest patch size independent of amount of forest in the surrounding landscape. I also found that increasing patch size positively influenced the relative abundance of pollinators. The observed differences in seed set likely resulted from differences in hummingbird movements (Chapter 3) and/or abundance under different landscape configurations. Title: Independent effects of habitat loss and fragmentation on pollination : tropical forest fragmentation alters hummingbird movements and pollination dynamics Author: Hadley, Adam S. A growing body of work reveals that animal-mediated pollination is negatively affected by anthropogenic disturbance. Landscape-scale disturbance results in two often inter-related processes: (1) habitat loss, and (2) disruptions of habitat configuration (i.e. fragmentation). Understanding the relative effects of such processes is critical in designing effective management strategies to limit pollination and pollinator decline. I reviewed existing published work from 1989 to 2009 and found that only six of 303 studies separated the effects of habitat loss from fragmentation. I provide a synthesis of the current landscape, behavioral, and pollination ecology literature in order to present preliminary multiple working hypotheses to explain how these two landscape processes might independently influence pollination dynamics (Chapter 2). Despite the potential importance of independent effects of habitat fragmentation, effects on pollination remain largely untested. Studies designed to disentangle the independent effects of habitat loss and fragmentation are essential for gaining insight into landscape-mediated pollination declines. I also found that the field of landscape pollination ecology could benefit from quantification of the matrix, landscape functional connectivity, and pollinator movement behavior. To test the hypothesis that pollinator movement can be influenced by landscape configuration, I translocated radio-tagged hummingbirds across agricultural and forested landscapes near Las Cruces, Costa Rica (Chapter 3). I found return paths were on average more direct in forested than in agricultural landscapes. In addition, movement paths chosen in agricultural landscapes were more forested than the most direct route suggesting that hummingbirds avoided crossing open areas when possible. To determine if differences in pollinator movement translated to differences in plant reproduction, I tested the relative importance of landscape composition versus configuration on the reproductive success of Heliconia tortuosa, a hummingbird-pollinated forest herb (Chapter 4). I used a stratified random sampling design to select sites across orthogonal gradients in patch size, amount of forest, and elevation. I tested four landscape change hypotheses (i.e., local, landscape composition, landscape fragmentation, and fragmentation threshold). I found that Heliconia reproduction supported both the local site and landscape fragmentation hypotheses. Seed set increased with increasing forest patch size independent of amount of forest in the surrounding landscape. I also found that increasing patch size positively influenced the relative abundance of pollinators. The observed differences in seed set likely resulted from differences in hummingbird movements (Chapter 3) and/or abundance under different landscape configurations. Close

• 2. [Article] Movement and space use by the green hermit (Phaethornis guy) in a fragmented landscape in Costa Rica

  Human activities have altered Earth’s ecosystems. Most biomes have experienced a 20-50% conversion to human use. Loss of habitat has obvious effects on the persistence of species. Fragmentation, however, ...

  Citation × Citation Citation Abstract Copy Title: Movement and space use by the green hermit (Phaethornis guy) in a fragmented landscape in Costa Rica Author: Volpe, Noelia L. Human activities have altered Earth’s ecosystems. Most biomes have experienced a 20-50% conversion to human use. Loss of habitat has obvious effects on the persistence of species. Fragmentation, however, may also negatively affect biodiversity for those species that exhibit behavioral responses to changes in habitat configuration. Such behavioral changes include movement, which is influenced by subdividing and isolating habitats through which animals prefer to travel. Ecosystem services that depend on the ability of animals to move through the landscape could be affected by changes in habitat configuration. Pollination is one such process because access of pollinators to flowers is often the key determinant of plant reproductive success. Yet, relatively little is known about how forest fragmentation may influence the flow of pollen carried by forest-dwelling pollinators, such as hummingbirds. I evaluate how changes in the distribution of pollinators produced by habitat fragmentation in southern Costa Rica affects a pollination web occupied by a generalist tropical pollinator, the green hermit hummingbird (Phaethornis guy). I used radio-telemetry to measure patterns of space use by green hermits living in forested areas with different levels of fragmentation. I first characterize patterns of space use by green hermits at three scales: point, path and home range. I found that green hermits have marked preference for locations in forested areas with high density of Heliconia plants and canopy cover; prefer paths close to streams that minimize crossing large stretches of non-forested matrix; and establish their home ranges in areas with high forest amount. In addition, home range area was negatively related to the amount of forest within it and positively related to fragmentation per se. Average resource density within the home range appeared to be unrelated to the area covered by the green hermits. My observations indicate that while the presence of green hermits in an area is mainly determined by the amount of forest available, the way they move through a landscape will be strongly affected by habitat configuration. In this context, pollen transfer by green hermits will be limited by the presence of open, non-forested areas owing to matrix avoidance. In landscapes with small discontinuous patches, the total area covered by an individual green hermit is likely to be small so I hypothesize that the genetic variability of pollen it transports may be diminished. I also assess the effectiveness of a tool commonly used to study the effect of habitat fragmentation on animal movement: translocation experiments. Such experiments are purported to have important advantages of being time efficient and standardizing across individuals the ‘motivation’ to move. Yet, we lack tests of whether movement behavior of translocated birds reflects natural behavior of unmanipulated birds. I compared the routine movement behavior of green hermits to that of experimentally translocated individuals. Behaviors documented during translocation experiments reflected those observed during routine movements. Both translocated and non-translocated birds showed similar levels of preference for mature tropical forest at the point level and avoided moving across non-forested matrix while selecting streams as movement corridors at the path level. Movement rates (distance covered per unit time) were generally higher during translocation experiments. However, the negative influence of forest cover on movement rates was proportionately similar in translocation and routine movement treatments. My results show that movement behavior of birds during translocation experiments is similar to natural movement behavior of birds. Therefore, translocation experiments may be reliable tools to address effects of landscape structure on animal movement. Title: Movement and space use by the green hermit (Phaethornis guy) in a fragmented landscape in Costa Rica Author: Volpe, Noelia L. Human activities have altered Earth’s ecosystems. Most biomes have experienced a 20-50% conversion to human use. Loss of habitat has obvious effects on the persistence of species. Fragmentation, however, may also negatively affect biodiversity for those species that exhibit behavioral responses to changes in habitat configuration. Such behavioral changes include movement, which is influenced by subdividing and isolating habitats through which animals prefer to travel. Ecosystem services that depend on the ability of animals to move through the landscape could be affected by changes in habitat configuration. Pollination is one such process because access of pollinators to flowers is often the key determinant of plant reproductive success. Yet, relatively little is known about how forest fragmentation may influence the flow of pollen carried by forest-dwelling pollinators, such as hummingbirds. I evaluate how changes in the distribution of pollinators produced by habitat fragmentation in southern Costa Rica affects a pollination web occupied by a generalist tropical pollinator, the green hermit hummingbird (Phaethornis guy). I used radio-telemetry to measure patterns of space use by green hermits living in forested areas with different levels of fragmentation. I first characterize patterns of space use by green hermits at three scales: point, path and home range. I found that green hermits have marked preference for locations in forested areas with high density of Heliconia plants and canopy cover; prefer paths close to streams that minimize crossing large stretches of non-forested matrix; and establish their home ranges in areas with high forest amount. In addition, home range area was negatively related to the amount of forest within it and positively related to fragmentation per se. Average resource density within the home range appeared to be unrelated to the area covered by the green hermits. My observations indicate that while the presence of green hermits in an area is mainly determined by the amount of forest available, the way they move through a landscape will be strongly affected by habitat configuration. In this context, pollen transfer by green hermits will be limited by the presence of open, non-forested areas owing to matrix avoidance. In landscapes with small discontinuous patches, the total area covered by an individual green hermit is likely to be small so I hypothesize that the genetic variability of pollen it transports may be diminished. I also assess the effectiveness of a tool commonly used to study the effect of habitat fragmentation on animal movement: translocation experiments. Such experiments are purported to have important advantages of being time efficient and standardizing across individuals the ‘motivation’ to move. Yet, we lack tests of whether movement behavior of translocated birds reflects natural behavior of unmanipulated birds. I compared the routine movement behavior of green hermits to that of experimentally translocated individuals. Behaviors documented during translocation experiments reflected those observed during routine movements. Both translocated and non-translocated birds showed similar levels of preference for mature tropical forest at the point level and avoided moving across non-forested matrix while selecting streams as movement corridors at the path level. Movement rates (distance covered per unit time) were generally higher during translocation experiments. However, the negative influence of forest cover on movement rates was proportionately similar in translocation and routine movement treatments. My results show that movement behavior of birds during translocation experiments is similar to natural movement behavior of birds. Therefore, translocation experiments may be reliable tools to address effects of landscape structure on animal movement. Close