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ZOOGEOMORPHOLOGY

Animals as geomorphic agents have primarily been considered "curiosities" in

the literature of geomorphology, whose spatial and quantitative influences have
been seen as both limited and minor. Zoogeomorphology: Animals as Geo-
morphic Agents examines the distinct geomorphic influences of invertebrates,
ectothermic vertebrates, birds, and mammals, and demonstrates the importance
of animals as landscape sculptors. Specific processes associated with the diversi-
ty of animal influences in geomorphology are examined, including burrowing
and denning, nesting, lithophagy and geophagy, wallowing and trampling, food
caching, excavating for food, and dam building by beavers. Particular empha-
sis is placed on terrestrial animals, although aquatic animals are also discussed
where appropriate.
This book, the only one available wholly devoted to this topic, will interest
graduate students and professional research workers in geomorphology, ecolo-
gy, environmental science, physical geography, and geology.
ZOOGEOMORPHOLOGY
ANIMALS AS GEOMORPHIC AGENTS

DAVID R. BUTLER
University of North Carolina, Chapel Hill

CAMBRIDGE
UNIVERSITY PRESS
To Janet and William Butler

CAMBRIDGE UNIVERSITY PRESS

Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, Sao Paulo

Cambridge University Press

The Edinburgh Building, Cambridge CB2 8RU, UK

Published in the United States of America by Cambridge University Press, New York

www.cambridge.org
Information on this title: www.cambridge.org/9780521433433
© Cambridge University Press 1995

This publication is in copyright. Subject to statutory exception

and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without the written
permission of Cambridge University Press.

First published 1995

This digitally printed version 2007

A catalogue record for this publication is available from the British Library

Library of Congress Cataloguing in Publication data

Butler, David R.
Zoogeomorphology : animals as geomorphic agents / David R. Butler
p. cm.
Includes bibliographical references and index.
ISBN 0-521-43343-6 (he)
1. Biogeomorphology. I. Title.
QH542.5.888 1995
591.52'2-dc20 94-44726
CIP

ISBN 978-0-521-43343-3 hardback

ISBN 978-0-521-03932-1 paperback
Contents

Acknowledgments P&ge y ii

1 Introduction 1
Definitions, scope, and limitations 3

2 The geomorphic influences of invertebrates 11

Geomorphic effects of terrestrial invertebrates 11
Aquatic invertebrates 27
Conclusions 28

3 The geomorphic accomplishments of ectothermic

vertebrates 29
The geomorphic role of fish 29
The geomorphic role of amphibians 34
The geomorphic role of reptiles 35
Conclusions 40

4 Birds as agents of erosion, transportation,

and deposition 42
Introduction 42
Internal clast transport as gastroliths 42
Geophagy 43
Clast transport for use as tools 43
Transport of materials for use in nest building 44
Mound building 46
Vegetation removal and its geomorphic effects 47
Burrowing and nest-cavity excavations 54
Conclusions 58
vi Contents

5 The geomorphic effects of digging for

and caching food 60
Digging for food 60
Caching of food 80

6 Trampling, wallowing, and geophagy by mammals 82

Trampling 82
Wallowing 92
Geophagy 95
Trampling erosion associated with a natural salt lick 97
Conclusions 107

7 The geomorphic effects of mammalian burrowing 108

Introduction 108
Burrowing and denning 109
The question of Mima mounds 142
Conclusions 146

8 The geomorphic influence of beavers 148

Beaver species and morphology 148
Historical and modern distribution of the beaver 149
Dam building and its geomorphic effects 151
The beaver pond environment and associated geomorphic
influences 159
Sedimentation and sedimentation rates in beaver ponds 170
Catastrophic dam failure and its effects 177
Conclusions 183

9 Concluding remarks 184

References 187
Index 221
Acknowledgments

The creation of a book such as this is a long process that has its roots in
many years of observation, classwork training, research, and travel. Many
people and organizations have directly or indirectly aided in the develop-
ment of my interests in animals as geomorphic agents, and deserve recogni-
tion. I would first like to express my gratitude to several individuals who
served as mentors during my training in geomorphology: Dr. Nicholas
Bariss and Dr. Jack Shroder, University of Nebraska-Omaha; and Dr. Curt
Sorenson, Dr. Bill Johnson, and Dr. Wake Dort, University of Kansas. I al-
so thank Dr. Jack Vitek for his friendship and guidance during my early
postgraduate years.
Field assistants and graduate students are some of the unsung heroes of
academia. Often for no salary, a limited food budget, and marginal housing,
they serve just to be there and to learn. I am very grateful to Susan Panciera,
Jack and Lori Oelfke, Dan Brown, Katherine Schipke, Dave Cairns, and
Bill Welsh for their help, comraderie, comments, and insight, and critical
mass when hiking in grizzly terrain. I also thank Marilyn Wyrick for her ex-
pert library ferreting.
Financial assistance for my work on zoogeomorphic topics has come
from a variety of sources. I thank Dr. Jim Fisher and the Department of
Geography, University of Georgia, for their support over the six-year period
of my residence there. The Department of Geography at the University of
North Carolina-Chapel Hill, and Dr. John Florin, Chair, have also provided
solid support for this project, not the least of which was in the form of a
Semester of Research and Study during the fall of 1993. Several grants and
agencies funded fieldwork in Glacier National Park, Montana, which ex-
posed me to a diversity of geomorphic effects produced by animals. I thank
the University of Georgia Research Foundation, Inc. (summer 1987), the
National Geographic Society (Grant 3831-88; summer 1988), Dr. Steve

vn
viii Acknowledgments

Walsh and his NASA grant (summer 1990), the Geography and Regional
Science Division of the National Science Foundation (Grant No. SES-
9109837) (summers 1991-2), the J. D. Eyre Fund of the Department of
Geography, UNC-Chapel Hill (January and October 1993), and the UNC
University Research Council (October 1993).
Officials in the Science Office of Glacier National Park provided research
collecting permits, logistical support, and housing. I particularly thank Clif-
ford Martinka, former Chief Scientist and current Senior Scientist, for his
efforts during my early research forays into the park; Carl Key, Park Geog-
rapher; and Dr. Dan Fagri, director of the park's climate change program.
Ms. Beth Ludeau-Denevan of the Park Library has also been most helpful.
Local individuals in the Glacier Park area have provided valuable infor-
mation and advice about field sites and conditions, accommodations, and
access through the years. Of particular note were Suzie, Sandy, and Megan
during the summer of 1988, who kept the cold ones coming and provided
occasional additional ones at no cost!
Dr. Robin Smith, Life Sciences Editor, and Michael Gnat, Production
Editor, have been extremely helpful and supportive of my efforts. I sincere-
ly thank them both.
Dr. Steve Walsh and Dr. George Malanson have, over the past twelve
years, been constant sources of camaraderie, friendship, and inspiration.
Each has been "infected" by the Glacier Park virus, for which I claim full
responsibility. They have collaborated together and separately with me on
many of the field efforts that resulted in my observations of animals as geo-
morphic agents, and I thank them sincerely. They each deserve a jug of
wine, a loaf of bread, and a huge hunk of Pecorino!
Finally, my family has, through the years, been my most consistent
source of support, love, and inspiration. My parents, Ray and Marian But-
ler, instilled in me a love for the natural world, and have been supportive of
my research efforts even when they had to defend to their friends why their
son was studying beavers! My brother, Mike Butler, has also been there as a
supporter and frequent field companion. Most important, my wife and son,
Janet and William Butler, have provided love and unwavering support. They
have understood why "Daddy has to go to Montana again." No one could
be luckier than I have been.
Introduction

Geomorphology is the study of surface processes and landforms (Easter -

brook 1993). Most geomorphologists, including Easterbrook, consider that
geomorphology also encompasses the evolution of landforms and interpre-
tations as to their origin. Geomorphology therefore examines the processes
currently or recently operative on the earth's surface that erode, transport,
and deposit sediment and that create landforms. Rhoads and Thorn (1993,
p. 288) succinctly summarized the discipline of geomorphology while ex-
tending its reach both temporally and spatially, by stating that the discipline
is "the study of past, present, and future landforms, landform assemblages
(physical landscapes), and surflcial processes on the earth and other plan-
ets."
In typical introductory geomorphology textbooks, a variety of surflcial
and internal processes are described. A common list of topics covered in
such books would include diastrophic forces of folding and faulting, inter-
nal and surface volcanism, weathering and soil development, gravity and
mass movement, the work of running water on and under the surface, the
work of glacial ice and ground ice, wind, and wave and current action. (See
Gregory [1988] for a discussion of recent curriculum trends in geomorphol-
ogy.) Unfortunately, these processes are frequently presented as if they
were operative in a sterile, nonliving void. When the biosphere is even ac-
knowledged or recognized in such works, it is usually restricted to large-
scale overviews of geomorphic processes operative in different climatic re-
gions or biomes (i.e., climatic geomorphology).
Animals are, however, a conspicuous element of the earth's physical
landscape and its environmental systems, but one that is typically glossed
over or completely ignored in earth-science texts and classes at the primary -
and secondary-school levels. Even in college and university classes in phys-
ical geography, animals are at best mentioned parenthetically. Only in ad-

1
2 Introduction

vanced university classes in biogeography are animals closely examined as

important elements of the physical landscape, and their role as geomorphic
elements is not typically emphasized in such classes.
Why has the role of animals been ignored in most twentieth-century geo-
morphology? Viles (1988a) identified the significance of Davisian geomor-
phology as the primary reason. Davisian models of landscape development
focused in large part on landscape history at the regional macro scale of in-
vestigation, rather than landscape processes operative at the meso or micro
scale. With the decline of Davisian geomorphology after World War II, the
study of processes received more attention at the meso scale. Processes op-
erative at the micro scale, such as biological organisms, have only recently
begun to be examined closely (Viles 1988a).
Even with the increased emphasis on micro-scale processes, however,
geomorphologists have not suddenly "jumped on the bandwagon" of study-
ing the effects of animals on the landscape. This is probably a result of the
perspective of many geomorphologists trained in the earth and physical sci-
ences rather than the biological sciences. The background and training of
geomorphologists, then, translates into an approach to the living landscape
in which they typically examine how surface processes affect environmen-
tal and ecological systems, rather than how elements of the biological land-
scape affect and may act as geomorphic processes (e.g., see Swanson et al.
1988). A certain parochial attitude among some geologically trained geo-
morphologists may also be a factor, reflected by an unfamiliarity with, or
unwillingness to utilize, literature sources from outside the earth sciences.
The cross-disciplinary nature of the discipline of geography makes geomor-
phologists trained in that tradition perhaps more aware of, and attentive to,
scientific literature in the biological sciences.
As the twentieth century draws to a close, commonly used geomorpholo-
gy texts (Thornbury 1969; Twidale 1975; Rice 1977; Chorley, Schumm, and
Sugden 1985; Selby 1985; Ritter 1986; Rohdenburg 1989; Bloom 1991;
Summerfield 1991; Easterbrook 1993), as well as summaries of different na-
tional research efforts in geomorphology (Ahnert 1989; Le Groupe Frangais
de Geomorphologie 1989; Suzuki 1989), still completely ignore the funda-
mental role of animals as geomorphic agents of erosion, transportation, and
deposition (but see Lobeck [1939] for an example of an early geomorpholo-
gy text that did include coverage of animals). The only exception to this rule
is in the study of coral reefs as biogenic landforms, which occasionally
gains mention in geomorphology textbooks and has a long tradition of study
in geomorphology going back (again) to the later works of William Morris
Definitions, scope, and limitations 3

Davis (Guilcher 1988). Indeed, the scholarly journal Coral Reefs is devoted
entirely to various aspects of the study of these unusual landforms.
This is not to say that geomorphologists have been completely derelict in
the study of animals as agents of erosion. Several papers have appeared
since the mid-1970s in major journals of geomorphology and physical ge-
ography dealing with topics such as the geomorphic role of beavers, ground
squirrels, rabbits, badgers, porcupines, grizzly bears (Figs. 1.1, 1.2), and
some invertebrates. Several chapters in Viles's (1988d) compilation on bio-
geomorphology examined aspects of animals as erosional agents, but the
emphasis there was primarily on geomorphological interactions with vege-
tation.(Also see the book edited by Thornes [1990], including Viles's [1990]
paper therein.)
Interestingly, however, animal ecologists and wildlife managers have al-
so recently made strides in the study of animals as geomorphic agents.
These studies typically examine animals from a paradigmatic framework of
landscape ecology or wildlife behavioral ecology, however, rather than from
a truly geomorphological perspective.
My goals in writing this book are to bring together this diverse research
literature covering the study of animals as geomorphic agents, whether from
a process geomorphology or a landscape/wildlife ecology paradigmatic
background, with my own experiences in the study of animals as geomor-
phic agents; to illustrate the levels of geomorphic work carried on in the
animal kingdom; and to compare those levels with the more traditionally
studied geomorphic processes. It has been suggested that "while much of
biogeomorphology is obviously fun, it is not so obviously fundamental"
(Cox 1989, p. 623). On the other hand, Johnson (1993, p. 69) stated that
"biota play fundamental roles in Earth's various systems, including soil-
slope systems." I hope to show that biogeomorphology, and more specifi-
cally zoogeomorphology, is both "fun" and indeed fundamental to a thor-
ough understanding of earth-surface processes and landforms. My approach
is that of a geomorphologist trained as a physical geographer.

Definitions, scope, and limitations

Several of Thornbury's (1969) concepts of geomorphology illustrate the im-
portance of knowledge of the past; yet at the same time, from a geomorphic
perspective, most landscapes postdate the bulk of the earth's history. In this
book, I examine only the late Holocene and its geomorphic processes asso-
ciated with animals. It would be fascinating to speculate on the geomorphic
 Introduction

Figure 1.1. Mountain slope excavated by grizzly bear (Ursus arctos horribilis) in search of
ground squirrel. Note 49-mm lens cap and pen for scale.
Figure 1.2. Trampled soil and grizzly bear pawprint, Otatso Creek drainage, Glacier National Park, Montana. Lens-cap diameter
is 49 mm.
6 Introduction

influence of Pleistocene megafauna, such as woolly mammoth wallowing

(and, indeed, Viles [1988c] briefly spoke to this topic); or on the trampling
effects of a herd of megadinosaurs such as Brachiosaurus. Without modern-
day observations or a sufficient historical data base, however, such specula-
tions must necessarily remain outside the confines of this book.
Viles (1988a, p. 1) defined biogeomorphology as "an approach to geo-
morphology which explicitly considers the role of organisms." Subsequent-
ly, I specifically defined zoogeomorphology as the study of the geomorphic
effects of animals (Butler 1992). By animals, I mean both endothermic
(warm-blooded) and ectothermic (cold-blooded) vertebrates as well as inver-
tebrates. I exclude from this definition the anthropogenic changes brought
about by human use of, and disruption of natural geomorphic processes on,
the landscape. Excellent coverage and reviews of this topic already exist
(Nir 1983; Trimble 1990; Phillips 1991; Sandgren and Fredskild 1991; Gou-
die 1993). Along similar lines, this book does not examine in any detail the
role of animals kept for agricultural purposes, but readers interested in that
topic are referred to Trimble (1988). The emphasis here is on free-ranging,
natural populations of animals (at least, as natural as possible in such a
human-impacted world). Animals that have been introduced and become
established in previously unoccupied ranges are, where appropriate, also dis-
cussed.
Because they have been extensively studied in their own right from both
geomorphic and biologic perspectives, I exclude coral reefs from inclusion
in the topics under discussion in this book. Readers interested in these bio-
genic landforms (in which only deposition occurs zoogeomorphically; ero-
sion and transportation of coral sediments are carried out by other geomor-
phic agents) are referred to Guilcher (1988), Spencer (1988), Viles's paper
(1988b) in her edited volume (Viles 1988d, and references therein), and the
aforementioned journal, Coral Reefs.
Obvious difficulties exist in identifying and documenting ocean- or lake-
floor geomorphic impacts by aquatic animals. Where such impacts have
been identified and have lasting geomorphic influence, they are described in
this book. The emphasis here, however, is on terrestrial animals. Vertebrates
are examined more closely than invertebrates, not because of any real or
imagined hierarchy of importance to geomorphic processes, but rather be-
cause of personal experiences and interests. In addition, the role of inverte-
brates has received a larger degree of synthesizing attention in recent liter-
ature, and readers interested in greater detail are referred to those works
where appropriate.
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