DOCSLIB.ORG

Population Dynamics of Reintroduced Elk (Cervus Elaphus) in Eastern North America by Jesse N. Popp

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Population Dynamics of Reintroduced Elk (Cervus Elaphus) in Eastern North America by Jesse N. Popp Population Dynamics of Reintroduced Elk (Cervus elaphus) in Eastern North America by Jesse N. Popp Thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy (Ph.D.) in Boreal Ecology Faculty of Graduate Studies Laurentian University Sudbury, Ontario © Jesse N. Popp 2017 THESIS DEFENCE COMMITTEE/COMITÉ DE SOUTENANCE DE THÈSE Laurentian Université/Université Laurentienne Faculty of Graduate Studies/Faculté des études supérieures Title of Thesis Titre de la thèse Population Dynamics of Reintroduced Elk (Cervus elaphus) in Eastern North America Name of Candidate Nom du candidat Popp, Jesse Degree Diplôme Doctor of Philosophy Department/Program Date of Defence Département/Programme PhD Boreal Ecology Date de la soutenance April 20, 2017 APPROVED/APPROUVÉ Thesis Examiners/Examinateurs de thèse: Dr. Frank F. Mallory (Supervisor/Directeur(trice) de thèse) Dr. Joseph Hamr (Co-supervisor/Co-directeur(trice) de thèse) Dr. Michael Persinger (Committee member/Membre du comité) Dr. Mark Boyce (Committee member/Membre du comité) Approved for the Faculty of Graduate Studies Dr. Jeff Larkin Approuvé pour la Faculté des études supérieures (Committee member/Membre du comité) Dr. David Lesbarrères Monsieur David Lesbarrères Dr. Arthur Rodgers Dean, Faculty of Graduate Studies (External Examiner/Examinateur externe) Doyen, Faculté des études supérieures Dr. Robert Lafrenie (Internal Examiner/Examinateur interne) ACCESSIBILITY CLAUSE AND PERMISSION TO USE I, Jesse Popp, hereby grant to Laurentian University and/or its agents the non-exclusive license to archive and make accessible my thesis, dissertation, or project report in whole or in part in all forms of media, now or for the duration of my copyright ownership. I retain all other ownership rights to the copyright of the thesis, dissertation or project report. I also reserve the right to use in future works (such as articles or books) all or part of this thesis, dissertation, or project report. I further agree that permission for copying of this thesis in any manner, in whole or in part, for scholarly purposes may be granted by the professor or professors who supervised my thesis work or, in their absence, by the Head of the Department in which my thesis work was done. It is understood that any copying or publication or use of this thesis or parts thereof for financial gain shall not be allowed without my written permission. It is also understood that this copy is being made available in this form by the authority of the copyright owner solely for the purpose of private study and research and may not be copied or reproduced except as permitted by the copyright laws without written authority from the copyright owner. ii Abstract Studies that focus on identifying factors that influence reintroduction success have often taken an individual population approach; however, investigating multiple populations can provide additional insight. The overall objective of this research was to emphasize the value of using within- and among-population approaches to identifying factors that influence the population dynamics of a reintroduced species. Elk (Cervus elaphus), a species that was extirpated from eastern North America during the late 1800s, has been reintroduced to portions of its former range over the past century through several initiatives. Today, there are several established populations across eastern regions of the USA and Canada, for which extensive monitoring data are available, creating an opportunity to investigate reintroduction success. I aimed to use these data to identify factors associated with changes in the survival and population growth rates of 10 reintroduced elk populations across eastern North America. More specifically, I: (1) performed a literature review detailing the history of elk reintroduction in eastern North America over the past century, (2) identified factors associated with the variation in population growth rates (reintroduction success) for 10 reintroduced elk populations using an among-population approach, (3) identified and assessed how climate affected the population growth rates of 7 reintroduced elk populations, and (4) investigated direct causes of mortality (predation and train collisions) associated with a single elk population experiencing low population growth. Although the number of successful elk restoration attempts has increased over the past century, there has been substantial variation in population growth rates among reintroductions. Major iii causes of elk mortality in restored populations differed between the pre- to post-acclimation phases of reintroduction. Population growth rates were negatively related to the percentage of coniferous forest within elk population range, suggesting that expansive areas of coniferous forests in eastern North America may represent sub-optimal elk habitat. The Burwash elk population in Ontario had low growth rate compared to most other populations reintroduced into eastern North America. Predation and train collisions were the most important source of mortality for this population. The number of annual elk-train collisions, as well as their locations, were monitored and recorded over 14 years. Collision locations were highly site- specific and were positively correlated to the proximity of bends in the railway. By relating the number of annual elk-train collisions to various climate factors, I found that collision rates were positively related to snow depth. By analyzing field camera data, I found that elk used the railway mostly during the fall and spring, when elk commonly travel to and from wintering grounds. However, by examining VHF telemetry locations, I determined that elk were closer to the railway in winter than in any other season. Railways likely are perceived by elk as easy travel corridors, especially in the winter, and deep snow might prevent escape from oncoming trains. Black bear (Ursus americanus) and wolves (Canis lupus) were the major predators of elk in the Burwash population. White-tailed deer (Odocoileus virginianus), elk (Cervus elaphus), and moose (Alces alces), were the ungulate prey species available to both predators. To determine if predators prefer one ungulate species over another, and to identify which predator species is likely to have a greater impact on elk survival, I investigated predator diets. To compare rates of iv ungulate use by predators in relation to prey availability, I calculated the relative abundance of each ungulate species. I found that wolves used juvenile and adult elk as their primary ungulate prey in greater proportions in comparison to their availability. Bears on the other hand, tended to use all ungulate species in proportion to their availability. Climate is well known to affect ungulate population dynamics; however, several factors (e.g.: density, predator presence), can govern the response. Relating the annual growth rates of 7 elk populations to various climate factors I found that responses were population specific. Increased annual snow fall was associated with declines in population growth rates for 2 of the 7 populations assessed and only 1 population responded negatively to increased summer temperatures. Climate likely interacts with other environmental variables to influence fluctuations in annual population growth rates which warrants further investigation. The results of this research will contribute to informed planning of future elk reintroductions and should support development through improved management. In addition, this research highlights the importance of using within- and among- populations approaches to investigating factors that influence elk reintroduction success. v Dedication Nothing has inspired me to pursue my dreams more than my children, Shyla and Hunter Popp. Their very existence is a constant reminder to me that anything is possible. My children, and my husband, Michael Popp, have made me smile every single day, supported me relentlessly, and fueled my desire to make my dreams come true. I dedicate this dissertation to the most amazing people in my world, Shyla, Hunter, and Mike. vi Acknowledgements Numerous people contributed to the development of this dissertation; however, my supervisors, Dr. Frank F. Mallory and Dr. Joseph Hamr were critical to its successful completion. Their guidance and unwavering support have been appreciated deeply throughout the years. Frank has graciously accepted my ongoing desire to work in his lab throughout several degrees and has unremittingly contributed to the development of my career. Joe has bestowed a wealth of elk knowledge upon me and in addition to contributing greatly to my new passion in life (elk!), has contributed significantly to the direction of my research. I will be forever grateful for their model supervision surrounding my project and everything they have done above and beyond what is expected. In addition, my committee members provided ongoing expertise through time spent in scheduled committee meetings, but also by making themselves available on a regular basis outside of meetings. Each of these members contributed significantly to the development of my thesis. I am very grateful for the time and expertise given to me by Dr. Mark S. Boyce, Dr. Jeff L. Larkin, and Dr. Michael Persinger. Over the years, two fellow grad students were of particular importance to the development of this dissertation. Tori Donovan and Dave McGeachy spent many days (and nights) listening to my statistics and study design rants, and were great friends
Load more

Recommended publications
  • INFLUENCE of a POPULATION IRRUPTION by ROOSEVELT ELK on a VEGETATION INDEX by Heath D. Starns, B.S. a Thesis Submitted To
    INFLUENCE OF A POPULATION IRRUPTION BY ROOSEVELT ELK ON A VEGETATION INDEX by Heath D. Starns, B.S. A thesis submitted to the Graduate Council of Texas State University in partial fulfillment of the requirements for the degree of Master of Science with a Major in Wildlife Ecology May 2014 Committee Members: Floyd W. Weckerly, Chair M. Clay Green Thomas R. Simpson COPYRIGHT by Heath D. Starns 2014 FAIR USE AND AUTHOR’S PERMISSION STATEMENT Fair Use This work is protected by the Copyright Laws of the United States (Public Law 94-553, section 107). Consistent with fair use as defined in the Copyright Laws, brief quotations from this material are allowed with proper acknowledgment. Use of this material for financial gain without the author’s express written permission is not allowed. Duplication Permission As the copyright holder of this work I, Heath D. Starns, authorize duplication of this work, in whole or in part, for educational or scholarly purposes only. ACKNOWLEDGEMENTS I am deeply grateful to the three members of my advisory committee for their support and guidance through this process. Butch Weckerly provided useful insight into the statistical procedures used and kept me focused while writing the manuscript. I am also indebted to Mark Ricca for his advice regarding the method used to obtain vegetation data, as well as to all the students involved with counting elk throughout the time of the study: R. Keleher, J. Hunt, D. Lancaster, M. Longoria, R. Luna, M. O’Dell, K. Richardson, S. Robinson, S. Shelton, G. Street, and D. Wolcott. Further assistance with counts, in addition to logistical support, was provided by staff biologists of Redwood National and State Parks.
    [Show full text]
  • UC Berkeley UC Berkeley Electronic Theses and Dissertations
    UC Berkeley UC Berkeley Electronic Theses and Dissertations Title Spatial Ecology and Population Dynamics of Tule Elk (Cervus elaphus nannodes) at Point Reyes National Seashore, California Permalink https://escholarship.org/uc/item/2wt3h3rc Author Cobb, McCrea Andrew Publication Date 2010 Peer reviewed|Thesis/dissertation eScholarship.org Powered by the California Digital Library University of California Spatial Ecology and Population Dynamics of Tule Elk ( Cervus elaphus nannodes ) at Point Reyes National Seashore, California by McCrea Andrew Cobb A dissertation submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy in Environmental Science, Policy and Management in the GRADUATE DIVISION of the UNIVERSITY OF CALIFORNIA, BERKELEY Committee in charge: Professor Reginald H. Barrett, Chair Professor Justin S. Brashares Professor Maggi Kelly Professor Eileen A. Lacey Spring 2010 1 Spatial Ecology and Population Dynamics of Tule Elk ( Cervus elaphus nannodes) at Point Reyes National Seashore, California © 2010 by McCrea Andrew Cobb 2 Abstract Spatial Ecology and Population Dynamics of tule elk ( Cervus elaphus nannodes ) at Point Reyes National Seashore, California by McCrea Andrew Cobb Doctor of Philosophy in Environmental Science, Policy and Management University of California, Berkeley Professor Reginald H. Barrett, Chair The tule elk ( Cervus elaphus nannodes ) was reintroduced to Point Reyes National Seashore, California in 1978. After exhibiting irruptive growth for 20 years, elk abundances have fluctuated. Three subpopulations of elk currently inhabit Pt. Reyes: a fenced subpopulation at high density; a free-ranging subpopulation in designated wilderness at low density; and a free- ranging subpopulation on active ranchland at low density. Little is known about the spatial ecology and population dynamics of tule elk.
    [Show full text]
  • Background of the Field
    © Jones & Bartlett Learning, LLC. NOT FOR SALE OR DISTRIBUTION PART I Background of the Field 78903_ch01_5807.indd 1 11/10/10 5:06:36 PM © Jones & Bartlett Learning, LLC. NOT FOR SALE OR DISTRIBUTION 78903_ch01_5807.indd 2 11/10/10 5:06:36 PM © Jones & Bartlett Learning, LLC. NOT FOR SALE OR DISTRIBUTION CHAPTER 1 Introduction: The Environment at Risk content areas of environmental health such as air quality, LEARN I NG OBJECT I VES water quality, food safety, and waste disposal. By the end of this chapter the reader will be able to: Maintaining environmental quality is a pressing task for • Describe how environmental health problems influence our the 21st century. Improvement in environmental quality is an lives official goal of the US government, as articulated in Healthy • Describe the potential impacts of population growth upon the environment People 2010. This goal (number 8, Environmental Health) is • State a definition of the term environmental health formatted as follows: “Promote health for all through a healthy • List at least five major events in the history of environmental environment.”1(p55) A list of environmental objectives is shown health in Table 1-1. Note that Healthy People 2020, which will provide • Identify current issues in the environmental health field updated objectives, was not available as of this writing. • Describe employment opportunities in the environmental health field According to Healthy People 2010: Physical and social environments play major roles in the health of individuals and commu- INTRODUCTION nities. The physical environment includes the air, water, and soil through which exposure to This chapter will illustrate how environmental health relates chemical, biological, and physical agents may to our lives and delimit the scope of the environmental health occur.
    [Show full text]
  • IM 11 Environmental Science
    MATSEC Examinations Board Marking Scheme IM 11 Environmental Science Main Session 2019 Marking Scheme (Main Session 2019): IM Environmental Science Marking schemes published by the MATSEC Examination Board are not intended to be standalone documents. They are an essential resource for markers who are subsequently monitored through a verification process to ensure consistent and accurate application of the marking scheme. In the case of marking schemes that include model solutions or answers, it should be noted that these are not intended to be exhaustive. Variations and alternatives may also be acceptable. Examiners must consider all answers on their merits, and will have consulted with the MATSEC Examinations Board when in doubt. Page 1 of 10 Marking Scheme (Main Session 2019): IM Environmental Science Question Marking scheme and suggested answers Marks Section A 1. a Any THREE from the following or suitable alternative: 3 Lowering of water table / Reduced surface water supplies / Land subsidence / Water quality concerns / Salt water contamination b Reverse osmosis; is the process in which seawater is pushed through an ultra-fine semi- 1, 2 permeable membrane. Only the water molecules can pass through the membrane, while salt and other chemicals cannot. c Any THREE from the following or suitable alternative: 3 Installing low flow shower heads / Turning off the tap while brushing your teeth or shaving / Placing water bottles in flushing tank / Buy washing appliances which are more water and energy efficient / Washing appliances are run only when they are full Total: 9 marks 2. a Insolation is the amount of incoming solar radiation that reaches the Earth’s surface.
    [Show full text]
  • Global Re-Introduction Perspectives: 2013 Further Case-Studies from Around the Globe Edited by Pritpal S
    Global Re-introduction Perspectives: 2013 Further case-studies from around the globe Edited by Pritpal S. Soorae Global Re-introduction Perspectives: 2013 Further case-studies from around the globe Edited by Pritpal S. Soorae IUCN/SSC Re-introduction Specialist Group (RSG) i ii The designation of geographical entities in this book, and the presentation of the material, do not imply the expression of any opinion whatsoever on the part of IUCN or any of the funding organizations concerning the legal status of any country, territory, or area, or of its authorities, or concerning the delimitation of its frontiers or boundaries. The views expressed in this publication do not necessarily reflect those of IUCN. Published by: IUCN/SSC Re-introduction Specialist Group & Environment Agency-ABU DHABI Copyright: © 2013 International Union for Conservation of Nature and Natural Resources Citation: Soorae, P. S. (ed.) (2013). Global Re-introduction Perspectives: 2013. Further case studies from around the globe. Gland, Switzerland: IUCN/ SSC Re-introduction Specialist Group and Abu Dhabi, UAE: Environment Agency-Abu Dhabi. xiv + 282 pp. ISBN: 978-2-8317-1633-6 Cover photo: Clockwise starting from top-left: i. Fen Raft Spider, UK © Helen Smith ii. Manglietia longipedunculata © Late Prof. Qingwen Zeng iii. European Tree Frog, Latvia © Andris Eglitis iv. Red Wolf © USA John Froschauer/PDZA v. Hungarian Meadow Viper © Tamás Péchy vi. Westslope Cutthroat Trout, USA © Carter Kruse, Turner Enterprises, Inc./Turner Endangered Species Fund vii. Oriental White Stork, Japan © Yoko Mitsuhashi Cover design & layout by: Pritpal S. Soorae, IUCN/SSC Re-introduction Specialist Group Produced by: IUCN/SSC Re-introduction Specialist Group & Environment Agency-ABU DHABI Download at: www.iucnsscrsg.org / www.iucn.org iii iv IUCN IUCN, International Union for Conservation of Nature, helps the world find pragmatic solutions to our most pressing environment and development challenges.
    [Show full text]
  • At Point Reyes National Seashore, California By
    Spatial Ecology and Population Dynamics of Tule Elk ( Cervus elaphus nannodes ) at Point Reyes National Seashore, California by McCrea Andrew Cobb A dissertation submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy in Environmental Science, Policy and Management in the GRADUATE DIVISION of the UNIVERSITY OF CALIFORNIA, BERKELEY Committee in charge: Professor Reginald H. Barrett, Chair Professor Justin S. Brashares Professor Maggi Kelly Professor Eileen A. Lacey Spring 2010 1 Spatial Ecology and Population Dynamics of Tule Elk ( Cervus elaphus nannodes) at Point Reyes National Seashore, California © 2010 by McCrea Andrew Cobb 2 Abstract Spatial Ecology and Population Dynamics of tule elk ( Cervus elaphus nannodes ) at Point Reyes National Seashore, California by McCrea Andrew Cobb Doctor of Philosophy in Environmental Science, Policy and Management University of California, Berkeley Professor Reginald H. Barrett, Chair The tule elk ( Cervus elaphus nannodes ) was reintroduced to Point Reyes National Seashore, California in 1978. After exhibiting irruptive growth for 20 years, elk abundances have fluctuated. Three subpopulations of elk currently inhabit Pt. Reyes: a fenced subpopulation at high density; a free-ranging subpopulation in designated wilderness at low density; and a free- ranging subpopulation on active ranchland at low density. Little is known about the spatial ecology and population dynamics of tule elk. This unusual situation provided an opportunity to quantify and compare factors affecting the home ranges, group sizes, resource selection, small- scale movements and population dynamics of tule elk. I collected biweekly fixes on radio- collared elk cows between 2005 and 2008 to examine herd structures, space use patterns, grouping behavior and survival rates of tule elk.
    [Show full text]
  • Proceedings of the 18Th Symposium
    SUMMER DIET AND FEEDING LOCATION SELECTION PATTERNS OF AN IRRUPTING MOUNTAIN GOAT POPULATION ON KODIAK ISLAND, ALASKA MCCREA A. COBB1, U.S. Fish and Wildlife Service, Kodiak National Wildlife Refuge, 1390 Buskin River Road, Kodiak, AK 99615, USA HEIDI HELLING, U.S. Fish and Wildlife Service, Kodiak National Wildlife Refuge, 1390 Buskin River Road, Kodiak, AK 99615, USA BILL PYLE, U.S. Fish and Wildlife Service, Kodiak National Wildlife Refuge, 1390 Buskin River Road, Kodiak, AK 99615, USA Abstract: The introduced mountain goat (Oreamnos americanus) population on Kodiak Island, Alaska, exhibited 60 years of rapid growth, resulting in an irruptive wave that expanded across the island from the initial introduction site. This unusual situation provided a rare opportunity to quantify and compare mountain goat summer diet and feeding location selection patterns at different stages along an irruptive growth cycle for an ungulate population. Diet composition analyses (via microhistological analyses of fecal pellets) indicated a temporal shift in summer mountain goat diets, which was likely driven by the onset of alpine plant growth following snowmelt. Sedges and forbs were important forage items and were increasingly consumed throughout the summer (June–August). Fern rhizomes were important in June (>16% of pellets), but less so (<1% of pellets) in July and August. Shrubs, mosses, and lichens were consistently consumed in small quantities (<5% of pellets), and therefore likely do not represent favored mountain goat summer forage on Kodiak Island. Consistent with our predictions, areas on Kodiak where mountain goats had completed an irruptive cycle (initial rapid population growth, decline to a lower abundance, and stabilization at a stochastic carrying capacity) had less forage cover at a lower diversity.
    [Show full text]
  • 1 AP Environmental Science Unit: Population 1. in the Real World
    AP Environmental Science Unit: Population Exam III: Chapter 6 (Population Biology) & Chapter 7(Human Population) 1. In the real world, many factors determine the numbers of organisms in any one population. Yet, a "superfly" with unlimited food and no mortality would show what type of growth? A. Carrying capacity geometric increase B. Irruptive growth C. Exponential growth D. Logistic growth E. Malthusian growth 2. A population growing at 1 percent per year should double in about ___ years. A. 30 B. 70 C. 10 D. 2 E. 0.7 3. Carrying capacity is the population A. That an environment can support in an optimal year B. Of a species that an environment can support on a long-term basis C. That remains after a catastrophic dieback has occurred D. Of a species without limiting factors E. Of a species without predators 4. In the real world, population explosion is usually followed by A. Continuous high population levels B. A gradual decrease in population as food supplies dwindle C. A tremendous increase in genetic diversity D. A gradual increase in population due to the availability of mates E. A population crash 5. A dieback or population crash, often occurs after a species ________ its environmental carrying capacity. A. Meets B. Undershoots C. Overshoots D. Oscillates around E. Decreases 1 6. An island has formed off the coast of the Pacific Northwest. The Department of Natural Resources (DNR) has determined that there is enough land area for a herd of 100 elk, yet they say the carrying capacity is only 60 elk.
    [Show full text]
  • Density Dependence in Ungulates: a Review of Causes, and Concepts with Some Clarifications
    550 CALIFORNIA FISH AND GAME Vol. 100, No. 3 California Fish and Game 100(3):550-572; 2014 Density dependence in ungulates: a review of causes, and concepts with some clarifications R. TERRY BOWYER*, VERNON C. BLEICH, KELLEY M. STEWART, JERICHO C. WHITING, AND KEVIN L. MONTEITH R. Terry Bowyer*, Department of Biological Sciences, Idaho State University, Pocatello, ID 83209, USA Vernon C. Bleich, Sierra Nevada Bighorn Sheep Recovery Program, California Department of Fish and Wildlife, 407 W. Line St., Bishop, CA 93514, USA Kelley M. Stewart, Department of Natural Resources and Environmental Science, 1664 N. Virginia St, Mail Stop 186, University of Nevada, Reno, NV 89557, USA Jericho C. Whiting, Gonzales-Stoller Surveillance, 120 Technology Drive, Idaho Falls, ID 83401, USA Kevin L. Monteith, Wyoming Cooperative Fish and Wildlife Research Unit, Department of Zoology and Physiology, Dept. 3166, 1000 E. University Ave., University of Wyoming, Laramie, WY 82071, USA Current address: Department of Biological Sciences, Idaho State University, Pocatello, ID 83209, USA (VCB) *Correspondent: [email protected] Understanding population dynamics of ungulates is not conceptually simple, in part, because there are many counterintuitive processes. We attempt to disentangle important concepts, including density dependence, density independence, limitation, regulation, compensatory and additive mortality, and top-down versus bottom-up forcing by examining how those ideas are related to the carrying capacity (K) of the environment. We contend that the K-selected, life-history characteristics of ungulates account for major components of their population dynamics. Those density-dependent attributes of ungulates require different management strategies than for species with attributes that are influenced primarily by density-independent processes.
    [Show full text]
  • Predicting Population Trends Under Environmental Change: Comparing
    PREDICTING POPULATION TRENDS UNDER ENVIRONMENTAL CHANGE: COMPARING METHODS AGAINST OBSERVED DATA Fiona Elizabeth Bridget Spooner A thesis submitted in partial fulfilment of the requirements for the degree of: Doctor of Philosophy of University College London 2019 Primary supervisor: Prof. Richard G. Pearson Secondary supervisor: Dr Robin Freeman I, Fiona Elizabeth Bridget Spooner, confirm that the work presented in this thesis is my own. Where information has been derived from other sources, I confirm that this has been indicated in the thesis ABSTRACT In this thesis I examine a range of approaches for predicting the impact of recent climate change and land use change on observed population trends. The thesis is split into three main parts. Firstly, I used linear mixed effects models to provide the first global assessment of the effects of environmental change on bird and mammal population trends. I find that populations have declined more rapidly in areas which have experienced rapid warming, this effect was more pronounced in bird populations. Secondly, I built habitat suitability models for 16 mammal species to explore the relationship between predicted habitat suitability and population abundance. I explored the correlations between time-series of rates of change in habitat suitability and corresponding time-series of observed population growth rates. There was little evidence to support the idea that population growth rates are directly linked to habitat suitability. However, when lagged responses are considered there is a stronger positive relationship between changes in habitat suitability and population growth rates, highlighting the importance of biodiversity time-series. Lastly, I built coupled niche-demographic (CND) models for three mammal species: Alpine ibex, brown bear and red deer.
    [Show full text]