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Of 1 46 a Trophic Cascade and the Potential !1 of !46 A Trophic Cascade and the Potential Consequences of Trophic Downgrading: The Cougar-Elk-Aspen System A Thesis Presented to The Faculty of the Environmental Studies Program The Colorado College In Partial Fulfillment of the requirements for the degree Bachelor of Science By: Bryson Sawyer Camp May/2019 ____________________________________ Miroslav Kummel Associate Professor of Environmental Science _____________________________________ Brian Linkhart Professor of Organismal Biology and Ecology !2 of !46 Table of Contents • Page 3: Abstract • Page 4: Introduction/Literature Review • Page 15: Methods • Page 23: Results • Page 31: Discussion • Page 41: Conclusions • Page 43: Works Cited !3 of !46 Abstract A trophic cascade can be defined as a specific food web or trophic structure where a predator preys on a herbivorous consumer which forages on a local vegetative resource; therefore, in this type of trophic structure top-down processes allow carnivorous apex predators to have indirect effects on local vegetative resources through their effects on the density or behavior/traits of the herbivores (M. Kummel, personal communication, April 8, 2019 and Ford et. al. 2015). In the case of predation on herbivorous consumers, both density and trait mediation can indirectly effect the density and growth pattern of vegetation that correlate directly to the alteration of prey populations via density and trait mediation (Ford et. al. 2015). Cougars have been identified as one of the seven apex predators that have been specifically associated with trophic cascades based on other empirical studies (Ripple et. al. 2014). Trophic downgrading follows the same pattern as alterations to trophic cascade structures: Trophic downgrading can have numerous direct and indirect ramifications on the local ecology. Trophic downgrading has similar consequences and is defined as, “the consequences of removing large apex consumers from nature (Estes et. al. 2011, 301).” Due to the unique characteristics that define the 6th mass extinction, one species has been the cause of most of the extinctions and the period has been characterized by the extinction of various large bodied animals, addressing trophic downgrading has become a prominent issue in the management of a wide array of ecological contexts globally (Estes et. al. 2011). In addition, apex predators, like cougars, facilitate ecosystem services such as carbon storage to buffer climate change, biodiversity enhancement, the reestablishment of native plant diversity, riparian restoration, and even the regulation of diseases (Estes et. al. 2011 and Ripple et. al. 2014). Thorough analyses of cougar habitat selection are rare, and have yet to be conducted in relation to the movement of elk and the growth of aspen saplings in the Pikes Peak region of Colorado U.S.A. until now. Through this study, it was observed and statistically shown that the number of aspen saplings tends to increase in areas that correspond with preferential habitat usage of cougars; whereas, the number of aspen saplings decreases in areas that correspond with a high prominence of observed elk herbivory. Therefore, in the Cougar-Elk-Aspen system within the Pikes Peak region cougars, carnivorous apex predators, are having indirect effects on local plants through top-down processes: This is a trophic cascade scenario. !4 of !46 Introduction/Literature Review Earth’s 6th Mass Extinction: The ongoing process of what is considered to be Earth’s 6th mass extinction has been unique in two distinct ways: one species has been the cause of most of the extinctions, and the period has been characterized by the extinction of various large bodied animals (Estes et. al. 2011). Large bodied animals are particularly at risk due to their tendency to have a low reproductive rate and large home ranges due to low density populations these qualities minimize the chance for adapting to changing environmental conditions and to pass on favorable traits for the new conditions to their offspring; furthermore, since climatic conditions are generally warming species are adapting by becoming smaller because larger surface-to-volume ratios are generally favorable under warmer conditions, due to standard metabolic principles (Scheffers et. al. 2016). To survive climate change, and avoid being affected by the 6th mass extinction or local extirpations species will have to adapt in at least one of three ways: adjust the range they use, alter themselves physiologically, or change how they operate in time which entails changing their phenology (Bellard et. al. 2012). Such adaptations are much more difficult when elongated life histories are taken into account: For this reason, smaller bodied species, those with shorter life histories, and/or those with a large capacity for phenotypic plasticity tend to be more able to adapt to changing environmental conditions which allows these species to pass on favorable traits for the new conditions to their offspring (Scheffers et. al. 2016). Trophic Cascades and Trophic Downgrading: !5 of !46 Trophic cascades have numerous direct and indirect ramifications on the local ecology. Trophic cascades are a well known concept within ecology that have striking patterns across ecological contexts (Estes et. al 2011). A trophic cascade can be defined as a specific food web or trophic structure where a predator preys on a herbivorous consumer which forages on a local vegetative resource; therefore, in this type of trophic structure top-down processes allow carnivorous apex predators to have indirect effects on local vegetative resources through their effects on the density or behavior/traits of the herbivores (M. Kummel, personal communication, April 8, 2019 and Ford et. al. 2015). It is also well known that the consequences of altering trophic cascade structures are far- reaching, often leading to regime shifts and alternative states of ecosystems, and the strength of these impacts will likely differ among species and ecosystems (Estes et. al 2011). Trophic downgrading has similar consequences and is defined as, “the consequences of removing large apex consumers from nature (Estes et. al. 2011, 301).” Trophic downgrading follows the same pattern as alterations to trophic cascade structures: Trophic downgrading can have numerous direct and indirect ramifications on the local ecology. For this reason, along with the impending prominence of trophic downgrading ecological triggering events, due to the 6th mass extinction, Estes et. al 2011 encouraged a shift in the ecological paradigm to focus more management efforts and resources on maintaining the integrity of top-down trophic interactions, on top of maintaining the status quo of bottom-up oriented management. Although, as a relatively new focus within ecology, there is numerous gaps in our scientific understanding of the top-down control aspects of trophic interactions since the ensuing consequences are unique to their ecological context. The time for action, research and conservation to limit !6 of !46 local extirpations and entire extinctions for some species is limited due to rapid and radical ecological changes associated with climate change and human development. The Role of Apex Predators: Notably, large bodied animals include numerous apex predators (Estes et. al. 2011). Due to the unique characteristics that define the 6th mass extinction, addressing trophic downgrading has become a prominent issue in the management of a wide array of ecological contexts globally. Thus, the production of novel and more effective management solutions to address the upcoming and unidentified consequences of trophic downgrading will be reliant on our ability to continually extract ecologically contextualized knowledge of specific events, encroachments, or local extirpations that cause trophic downgrading, alterations to trophic cascade structures, and/or ecological collapse. In an effort to elucidate ecological events that trigger alterations to trophic cascade structures and trophic downgrading, such as extinctions and local extirpations, ecologists must continue to study the intricacies of food webs, with a specific focus on the behaviors and ecological impacts of apex predators and large bodied animals. This must be done extensively in a variety of ecological contexts globally in order to understand the potential consequences of impending alterations to trophic cascades structures and trophic downgrading. In the case of specifically attempting to elucidate trophic downgrading ecological triggering events, wide ranging apex predators may be ideal to focus on for uniformity. Hopefully, such scientific endeavors will elucidate !7 of !46 potential management strategies to protect apex predators and large bodied animals in general, within unique ecological contexts around the world. Historically, ecological studies have tended to lack focus on apex predators and large bodied animals alike due to the strong focus on bottom-up processes, since bottom- up processes has been viewed as fundamental to the function of ecosystems (Estes et. al. 2011). However, apex predators, are now commonly considered to be keystone species within their ecosystems who uphold overlooked roles in regulating natural occurrences of disease, fire, carbon sequestration, invasive species along with biogeochemical exchanges amongst Earth’s soil, water, and air (Estes et. al. 2011). Consequently, trophic downgrading has been associated with biodiversity loss (Estes et. al. 2011). Thus, the new found importance of apex predators and the rise of extinctions
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