THE NUTRITIONAL ECOLOGY OF BLACK-TAILED DEER By AMY COLLEEN UL

165 Pages·2015·3.45 MB·English

by  Amy Ulappa

USING FORAGING DYNAMICS TO ANSWER LANDSCAPE MANAGEMENT
QUESTIONS: THE NUTRITIONAL ECOLOGY OF BLACK-TAILED DEER

By
AMY COLLEEN ULAPPA

A dissertation submitted in partial fulfillment of
the requirements for the degree of

DOCTOR OF PHILOSOPHY

WASHINGTON STATE UNIVERSITY
School of the Environment

DECEMBER 2015

© Copyright by AMY COLLEEN ULAPPA, 2015
All Rights Reserved
© Copyright by AMY COLLEEN ULAPPA, 2015
All Rights Reserved
To the Faculty of Washington State University:

The members of the Committee appointed to examine the dissertation of AMY
COLLEEN ULAPPA find it satisfactory and recommend that it be accepted.

______________________________________________
Lisa A. Shipley, Ph.D., Chair

______________________________________________
Rachel C. Cook, Ph.D.

______________________________________________
Janet L. Rachlow, Ph.D.

______________________________________________
Mark E. Swanson, Ph.D.

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ACKNOWLEDGMENTS

It takes a village to complete a graduate project and that is especially true in my case.
First, I want to thank my major advisor, Lisa Shipley, for being a patient teacher, advocate, and
inspiring example, both professionally and personally. Lisa never seemed to doubt that I would
succeed and that was invaluable to me during every step of the project and meant more than I can
express. I would like to thank Rachel Cook for teaching me how to do all the field methods, to
knit, and for encouraging me to be a thorough scientist. I am grateful to Rachel for helping me
raise fawns, mentoring me in the field the first summer (and keeping me from going crazy),
teaching me to turn an 18 foot stock trailer around on a dirt road, and for being a committee
member as well. Thank you to Mark Swanson for all of his forestry expertise, statistics advice,
being a committee member, and for coming out to the field last-minute to chainsaw a perimeter
to build a deer pen. I would like to thank Janet Rachlow for the insight she has given me on
habitat ecology and for being a committee member as well. All the members of my committee
have been generous of their time, knowledge and their suggestions have shaped my dissertation;
I am grateful for their guidance.
This project was possible through collaboration with many people and organizations. I
would like to thank John Cook for his work getting the project off the ground and continued
input throughout the project. Thank you to Bruce Davitt and the Wildlife Nutrition Lab at WSU
for all the sample analysis and for welcoming me into the lab. The private land owners and
companies we worked with provided in-kind support in the form of time and logistical help.
Mike Rochelle, Julie Kehough, Karen Temen, Mark Sheldahl and Rod Meade from
Weyerhaueser Timberlands, Tim McBride from Hancock Timber Resource Group, Blake
Murden from Port Blakely Tree Farms, Randall Greggs from Green Diamond Resource

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Company, Candace Cayhill from Rayonier Forest Resources, Daniel Ravenal from the Quinault
Division of Natural Resources, and Dave Vales from the Muckleshoot Wildlife Department all
located sites, answered questions, and provided useful advice and logistical support. I especially
want to acknowledge Rod Meade for the weeks he spent driving me to all the sites, marking
maps so I wouldn’t get lost, and teaching me so much about the history of western Washington.
The City of Tacoma and Seattle Public Utilities also allowed us access to their land. All the
collaborators I worked with on this project were welcoming, helpful, and I have benefited greatly
from their expertise.
I am grateful for all the volunteers and students who have helped me care for animals,
and do field and lab work. Specifically, thank you to Emily Reinig, Zorah Oppenheimer, Jarrett
and Bobbilee Schuster, Marley Iredale, Jennifer Smith, Cheryl Payne, Alysa Adams for their
help raising fawns for the project. I want thank several fellow graduate students, Sara Wagoner,
Sarah McCusker, Jeremy Brown, Laura Felicetti, Miranda Crowell, and Meghan Camp for
answering my questions, assisting with deer emergencies, and helping to keep me sane. The field
technicians who worked with me deserve special thanks. They worked (and slept) in the rain,
heat, and dirt for 4 months, and survived veg ring, the black mesh, Stinky Pete, and so many
more trials. Thank you to Matt Thomas, Nick Baumgarten, Jameson Demeglio, Rachel
Granberg, Dave Keiter, Mason Cole, Amanda Rudie, Sam Stripes and Stephanie Berry (I am so
glad you were crazy enough to come back for a second year!) for your hard work, making me
laugh, and for teaching me so much about teamwork. All the field work resulted in thousands of
samples to be weighed and analyzed and piles of data to be entered. I could not have gotten it
done by myself so thank you to Amanda Sundahl, Curtis Flolid, Nick Steveson, Dee Orr, Stacy
Hanson, Krista Rensi, Erin Ranney, Katie Meline, Dan Nyquist, Alex Biswas, Sandee Ditt,

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Miranda Crowell, Marley Iredale, and Bobbilee Schuester for all your work in the lab and
entering data.
Finally, thank you to my family. I appreciate the support of my parents, Thomas Ulappa
and Janis and Gary Kerby, in my decision to be a student for a very long time and for always
showing me love and encouragement. Thank you to my husband, Shannon, for more than I can
list, including, encouraging me to pursue my Ph.D. and for all the sacrifices that went along with
that. He has been pooped on by every animal species I have studied, fixed field gear and trucks,
driven thousands of miles to see me on the weekends, and talked me down from many rants. I
want to thank Shannon and our young son, Silas, for bringing me so much joy to my life. I also
want to thank Olivia and Jake Simpson for watching Silas so I could work and for showing me
how great a big family is. Lastly, and sort of like family, I want to thank my deer friends (Dora,
Gail, Georgia, Freeda, Sarah 2, Mabel, Wednesday and Xena) for all their work in the field and
all the deer at the Wild Ungulate Facility. It was a privilege to work with such magnificent
animals.
This dissertation was made possible by many funding sources and I would like to
acknowledge Washington Department of Fish and Wildlife, the National Council for Air and
Stream Improvement, and the Muckleshoot tribe for their substantial contributions. I also
appreciate funding I received from the Washington chapter of the Wildlife Society, the Safari
Club, and Washington State University School of the Environment. Additionally, this work was
supported by the USDA National Institute of Food and Agriculture, Hatch Project 1005876.

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USING FORAGING DYNAMICS TO ANSWER LANDSCAPE MANAGEMENT
QUESTIONS: THE NUTRITIONAL ECOLOGY OF BLACK-TAILED DEER
Abstract

by Amy Colleen Ulappa, Ph.D.
Washington State University
December 2015

Chair: Lisa A. Shipley

Black-tailed deer (Odocoileus hemionus columbianus) and other forest herbivores depend
on abundant and nutritious understory vegetation found in open-canopy and young, early-seral
forests. Forage management activities such as logging, thinning and using silvicultural
herbicides have the potential to alter the quality and quantity of forage available to herbivores.
Therefore, I compared nutrient intake and estimated nutritional carrying capacity of black-tailed
deer between stands that received herbicide treatments paired with those that did not in Douglas-
fir/western hemlock forests of western Washington, and investigated how these responses change
as forests age from early seral stages to canopy closure (2 – 20 years post-harvest). I measured
understory vegetation biomass, plant species nutritional quality, and overstory characteristics of
each forest stand. Additionally, I measured diet composition, diet quality and nutrient intake of
tractable black-tailed deer within each paired stand. I found that herbicide use decreased the
amount of understory biomass useable for foraging deer and decreased their daily digestible
energy intake, especially in the first 3 years of stand establishment. Even so, early seral habitat,
regardless of herbicide use, provided more forage and higher daily energy intake for deer than
closed canopy stands (14 yrs +) or mid-seral stands. Using data from these experiments, I also
investigated how the resolution of data collection (e.g., plants vs. plant parts) influences

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estimates of plant selection and nutritional carrying capacity, and how well measures of forage
abundance and quality predict nutrient intake of black-tailed deer. I found that treating plant
parts (i.e., stems and leaves) as separate food items did not influence how plant species were
categorized as selected, neutral or avoided, but increased estimates of nutritional carrying
capacity. In addition, measures of forage resources that included both biomass and quality or
selection status by deer better predicted daily digestible energy intake of tractable black-tailed
deer.

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TABLE OF CONTENTS

Page
ACKNOLEDGEMENTS...............................................................................................................iii
ABSTRACT...................................................................................................................................vi
LIST OF TABLES..........................................................................................................................xi
LIST OF FIGURES......................................................................................................................xiv
BACKGROUND AND ATTRIBUTION........................................................................................1
LITERATURE CITED......................................................................................................11
CHAPTER ONE
1. INTRODUCTION.........................................................................................................17
Study area...............................................................................................................22
2. METHODS....................................................................................................................23
Experimental design and treatments......................................................................23

Overstory characteristics........................................................................................25
Biomass, species composition and available forage..............................................26

Estimated useable biomass and nutritional carrying capacity...............................27

Diet composition, selection and foraging behavior of deer...................................28

Nutritional quality of forages and deer diets..........................................................31

Deer nutrient intake................................................................................................32
Statistical analysis..................................................................................................33
3. RESULTS......................................................................................................................34

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Characteristics of forest overstory.........................................................................34
Biomass and composition of forest understory.....................................................34

Estimated nutritional carrying capacity.................................................................37
Composition and selection of deer diets................................................................37
Nutritional quality of deer diets.............................................................................38

Deer nutrient intake................................................................................................38
Unmanaged mid-seral stands compared to stands <20 yrs old..............................39
4. DISCUSSION................................................................................................................41

5. LITERATURE CITED..................................................................................................54

6. TABLES........................................................................................................................65

7. FIGURES......................................................................................................................72

CHAPTER TWO

1. INTRODUCTION.........................................................................................................80

Study area...............................................................................................................83

2. METHODS....................................................................................................................84

Habitat characteristic measurements......................................................................84
Diet composition, foraging behavior and daily nutrient intake of deer.................85
Nutritional quality of forages and deer diets.........................................................87
Deer nutrient intake...............................................................................................88
Plant selection by deer..........................................................................................89
Estimated Nutritional Carrying capacity..............................................................89
Statistical analysis.................................................................................................90
3. RESULTS.....................................................................................................................91

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