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Carnivoran Ecology
Carnivoran Ecology
The Evolution and Function of
Communities
Steven W. Buskirk
Professor Emeritus, University of Wyoming
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
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Published in the United States of America by Oxford University Press
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Preface
This book arose from my several decades of interest a small part of our growing understanding of
in and research on carnivorans. Like many young what limits the distributions and abundances of
people with naturalist tendencies, I was particu- carnivorans.
larly intrigued by my early encounters with wild During those early years, I depended on the stan-
carnivorans—their rarity, elusiveness, and implicit dard academic references of the day: journal arti-
threat to my well-being. I vividly recall seeing my cles, monographs, book chapters, theses, disserta-
first bear track on a solo backpacking trip in the tions, and R. F. Ewer’s (1973) The carnivores. Ewer’s
Sierra Nevada of California, followed by a sleep- monograph was the definitive source for carnivo-
less night spent contemplating my fate. I recall the ran biology—especially paleontology and behavior.
horror of my older sister when I asked her to stop Writing the current account nearly fifty years on,
our parents’ car so that I could retrieve a road-killed I benefitted from a wealth of high-quality material
domestic cat and add its skull to my collection. In facilitated by the revolution in publishing—a prolif-
the summers of my college years, working as a tour eration of journal titles, many of them open access,
guide in Mount McKinley (now Denali) National expanded opportunities for scientists in develop-
Park, Alaska, I observed regular interactions involv- ing countries to publish their work, and many more
ing wolves, caribou, brown bears, moose, red foxes, women working and publishing as scientists. The
and other species. Some of these observations were period since 2000 has offered unprecedented oppor-
dramatic and photogenic, but curious as well. Why tunities to write a book such as this, and the pan-
did 150-kg brown bears invest so much effort to demic of 2020–22 gave my isolation and focus new
capture 200-g ground squirrels? Why did wolves purpose.
give birth in the same dens for decades on end, What qualifies me to write a book about all car-
even though they were well known and prone nivorans? I am not the most prolific author of car-
to human disturbance? Why were coyotes deathly nivoran papers, nor the one with the broadest geo-
afraid of being anywhere near wolves, while red graphic experience. Others have spent more time
foxes merely stayed out of their grasp? in the field, or are more quantitative than I. How-
My PhD research and subsequent faculty ever, my interests extend in multiple directions,
appointment in Zoology and Physiology at the all of which relate to how carnivorans succeed or
University of Wyoming gave me opportunities fail in the wild. I am as intrigued by answers to
to pursue these kinds of questions. While most big ecological questions as by solutions to specific
carnivoran research of that time gravitated to conservation problems. I am as satisfied by under-
intraspecific and predator–prey interactions, standing of some ecological puzzle as by watching
my interests ranged more widely. My research a large carnivore stalk an ungulate. I appreciate new
addressed various mechanisms by which car- discoveries in natural history as much as I do stud-
nivorans might be limited: thermal energetics, ies of functional genomics. I also value those who
allometry, tooth morphology, fasting endurance, study carnivorans and share their findings with sci-
genetic variability, and interspecific competition. entists and the public. This group overlaps strongly
I learned that predator–prey interactions were with those committed to assuring the presence and
vi P R E FA C E
Many people supported and aided me in writing Van Valkenburgh, Lars Werdelin, and Andrzej
this book. My wife Beth encouraged me at every Zalewski. Collectively, the reviewers corrected
stage and tolerated many interruptions to our rou- errors of fact, identified important omissions, pro-
tine so that I could spend time writing. Close col- vided more pertinent or more recent references,
leagues Dennis Knight and Carlos Martinez del Rio and improved the organization and presentation.
were reliable discussants and sources of encour- Without their help the project would not have been
agement early in the conception and early writing possible.
phases. Hannah Sease produced all graphic arts The illustrations strengthen the narrative
work, keeping pace with my requests throughout throughout, and some of the finest artwork and
her graduate studies in graphic arts. photography were contributed gratis or licensed
The William Robertson Coe Library at the Univer- at reduced rates. I particularly thank Justin Binfet,
sity of Wyoming provided outstanding support, fill- Darin Croft, Walton Ford, Stan Gehrt, Don Gutoski,
ing scores of requests for materials, and the J. Cloyd Esperanza Iranzo, Jeffrey Kerby, Janet Kessler,
Miller Library at Western New Mexico University Débora Kloster, Susan McConnell, Chris Mills,
provided additional assistance. My academic home, Larissa Nituch, Velizar Simeonovski, Alejandro
the Department of Zoology and Physiology at the Travaini, Juan Zanón, the Philmont Scout Ranch,
University of Wyoming, provided office space and and Kasmin Gallery. Although I have tried to
other support for most of duration of the project. select photos taken under natural conditions, I
I benefitted from reviews by over thirty scien- cannot assure that none was staged or in some way
tists, eight of them commissioned by Oxford Uni- contrived.
versity Press, and others solicited by me, who The staff of Oxford Press were most support-
generously reviewed chapters or shorter sections: ive and encouraging. Ian Sterling was immediately
Benjamin Allen, Rudy Boonstra, Jeff Bowman, receptive when I approached him with my pro-
Joseph Bump, Emiliano Donadio, Jacob Goheen, posal, and consistently improved my presentation
Henry Harlow, Dennis Knight, Serge Lariviere, as well as my understandings of how the book could
Paul Leberg, Jason Lillegraven, Carlos Martinez be made most useful. Charlie Bath provided excel-
del Rio, Sterling Miller, Robert Naiman, Richard lent suggestions on draft chapters, and Katie Lakina
Ostfeld, Jonathan Pauli, James D. Rose, Oswald shepherded the project through the editorial and
Schmitz, John Schoen, Qian-Quan Sun, Blaire production processes.
Contents
2 Functional morphology 11
2.1 The skull 11
2.1.1 Dentition 12
2.2 Post-cranial skeleton 15
2.2.1 Fossorial movement 16
2.2.2 Running and walking 16
2.2.3 Climbing 17
2.2.4 Swimming and deep diving 18
2.3 Other adaptations to aquatic living 19
2.4 Gut morphology 20
2.5 The integument 20
2.6 The major ecomorphotypes 22
2.6.1 Scansorial ecomorph 23
2.6.2 Dog-like ecomorph 23
2.6.3 Cat-like ecomorph 23
2.6.4 Scavenger ecomorph 24
2.6.5 Semi-fossorial ecomorph 24
2.6.6 Semi-aquatic ecomorph 24
2.6.7 Marine ecomorph 25
2.6.8 Intermediate and unique ecomorphs 25
Key points 25
References 26
4 Physiological ecology 49
4.1 Digestion 49
4.1.1 Soluble carbohydrates 50
4.1.2 Gut passage 51
4.2 Dietary requirements 52
4.2.1 Amino acids and fatty acids 52
4.2.2 Macronutrients 53
4.3 Metabolism and growth 53
4.3.1 Metabolism 53
4.3.2 Growth 55
4.4 Body temperature and torpor 55
4.4.1 Body temperature 55
4.4.2 Torpor 56
4.5 Energy storage and fasting 56
4.6 Osmoregulation and kidney function 58
4.7 Detoxification and self-medication 60
4.8 Reproduction 61
4.8.1 General patterns 61
4.8.2 Embryonic diapause 61
4.8.3 Induced ovulation 63
4.9 Scaling physiology to populations 64
Key points 65
References 65
6 Community ecology 87
6.1 Nutrient cycling and transport 87
6.2 Direct effects on soil 90
CONTENTS xi
9 Cascades 153
9.1 Ecological cascades 153
xii CONTENTS
Glossary 259
Index 263
CHAPTER 1
Order Carnivora represents one of the most species- emotions are stronger when we imagine ourselves
rich, phenotypically diverse, widely distributed, or animals we own as prey. As a result, we have long
and ecologically influential mammalian lineages. imbued carnivorans with spiritual powers to match
Its extant members live on all continents and in their impressive physical abilities, and our ances-
all oceans and range from vole-sized (c. 30 g) tors represented carnivorans in some of the earliest
to larger than a rhinoceros (c. 4,000 kg). They figurative art (Figure 1.3) (Hart and Sussman, 2008;
eat diverse foods including leaves, fruits, insects, Azéma, 2015). Today, we continue to use carnivo-
honey, marine invertebrates, and mammals larger rans to symbolize wildness, ferocity, and indepen-
than themselves. Some species live below ground dence in visual and literary arts. Every pocket and
for weeks at a time, a few live mostly in the fold of most human cultures—languages, parables,
forest canopy, and others live at sea for months spiritual beliefs, and symbols—is rich with carnivo-
on end. Many are wilderness dwellers, wary of ran references.
humans and their activities, while some non-
domestic species thrive in major cities, largely
1.1 “Carnivoran” vs. “carnivorous”
dependent on humans for food, shelter, or protec-
tion from larger, wilder carnivorans. No other mam- The terms “mammalian predator,” “mammalian
malian order approaches Carnivora in the breadth carnivore,” and “carnivoran” are not precisely syn-
of adaptive suites shown and ecological niches onymous. Predators are animals that kill and con-
occupied. sume multicellular animals (Taylor, 1984), whether
Humans have always been keenly interested in one at a time, or filtered from water by the thou-
carnivorans. Our hominin ancestors were preyed sands. Eagles, dragonflies, and blue whales are
on by carnivorans and competed with them for predators. By contrast, a carnivore consumes the
food. Both species hunted the same prey and drove flesh of animals, whether it kills or scavenges it; vul-
each other away from prey carcasses (Figure 1.1) tures, snakes, and Venus fly traps are carnivores and
(Espigares et al., 2013). Paleolithic humans con- carnivorous. Carnivorans, the subject of this book,
verted a potential competitor, the wolf, to a partner. are exclusively mammals in Order Carnivora—a
The resultant dog was the first domesticated animal, branch of the tree of life. Species in the order
and became essential to human lives, providing a may pursue and kill vertebrates, eat termites exca-
food source, vigilance against intruders, transport vated from soil, or subsist entirely on plant parts
of possessions, and assistance in hunting and herd- (Figure 1.4). “Hypercarnivore” sometimes indicates
ing (Figure 1.2). Dogs became so important to early a species with a diet exceeding some threshold level
humans that they are credited with shaping human of vertebrate prey, typically killed rather than scav-
evolution as much as humans shaped theirs (Pierotti enged. Most members of the cat family, the Felidae,
and Fogg, 2017). are considered hypercarnivores. Other terms, such
The life-or-death nature of predation elicits strong as “mesopredator,” “apex predator,” and “keystone
human emotions—either the predator eats and predator,” have imprecise meanings that I parse as
lives, or the prey escapes and survives. These they arise in the book.
Carnivoran Ecology. Steven W. Buskirk, Oxford University Press. © Steven W. Buskirk (2023). DOI: 10.1093/oso/9780192863249.003.0001
2 CARNIVORAN ECOLOGY
Figure 1.1 The first contact between humans and the saber-toothed Smilodon in America, interpreted by Velizar Simeonovski. Carnivorans have
been competitors with and potential predators of hominins from before we became humans through to today.
Painting: © Velizar Simeonovski.
I N T R O D U C T I O N TO C A R N I V O R A N E C O L O G Y 3
Figure 1.2 Lions (right) depicted on the walls of Salle du Fond in Chauvet-Pont d’Arc Cave, France, dating to 32,000–30,000 years ago. The lions
appear to be watching a rhinoceros on a distant panel. Carnivorans were subjects of some of the earliest figurative art. Vertical scratches in the
lower right and on the rhinoceros painting were made by cave bears trapped in deep chambers.
Photo: J. Clottes in Azéma (2015), CC 4.0.
As is often the case in biology, the defining traits defining the group, because all modern species have
of the Carnivora—the traits held by all members that trait, but some early carnivorans did not.
of the lineage, both living and extinct—are difficult Predatory placental mammals—those that pri-
to state without qualification. Traditionally, biolo- marily kill other animals for food—occur in several
gists have cited the presence of cheek teeth that have orders, including shrews and moles (Order Sorico-
shearing functions, comprising the upper fourth morpha), bats (Chiroptera), whales (Cetacea), pan-
premolar (P4 ) and lower first molar (M1 ). All living golins (Pholidota), and hairy anteaters (Pilosa). If
carnivorans have ancestors with this trait. However, we broaden our frame of reference to include mar-
living genets, bears, and seals have secondarily lost supial predators, several additional orders must
the shearing function of those teeth, and at least be included as mammalian carnivores. Even some
one species has lost those teeth completely. Some obligate herbivores, among them ungulates and
prehistoric linages of other carnivorous mammals rodents, prey on or scavenge vertebrates oppor-
had shearing cheek teeth, but they occupied other tunistically (Boonstra et al., 1990; Dudley et al., 2016).
positions in the tooth row, for example M1 and M2 . Finally, many extinct non-carnivoran lineages were
They were not homologous to modern carnassials, at least partially carnivorous. Clearly, Order Car-
and the lineages that exhibited them have no living nivora is not unique among mammals in killing and
descendants. The fused scaphoid and lunate carpal eating vertebrates. This book is about a mammalian
bones (the scapholunate) are sometimes regarded as lineage, not a foraging style or trophic niche.
4 CARNIVORAN ECOLOGY
Figure 1.3 Rock art in northwestern Saudi Arabia showing dogs resembling modern Canaan dogs assisting with lion hunting. Various glyphs
from this site include the earliest depictions of dogs on leashes, 12,000–10,000 years old. The upper image is shaded to show relief that is less
visible in the lower, unaltered image.
Photo: Guagnin et al. (2018, Figure 10) by permission.
I N T R O D U C T I O N TO C A R N I V O R A N E C O L O G Y 5
Herbivorous Carnivorous
mammals Carnivora mammals
1.2 The carnivorans—who and where? nearly cosmopolitan, found on all continents except
Antarctica and Australia before humans trans-
The approximately 287 extant species of carnivo-
ported them. On the other hand, the eight species
rans are organized into fifteen currently recognized
of Eupleridae occur only on Madagascar Island,
families (Appendix I) and occur on all continents, if
their common ancestor having rafted there from
we include seals that haul out on Antarctic beaches.
the African mainland around 20 million years ago
These numbers change as we learn more about
(Ma). The thirty-three extant species of seals, sea
how mammal lineages are related. For example, the
lions, and walrus make up the pinniped (ear foot)
neotropical olinguito recently has been identified as
group—a lineage comprising two or three families
distinct from other olingos, and the African golden
that arose from a single aquatic ancestor. “Fissiped”
wolf was judged a separate species from the golden
(split foot), on the other hand, denotes the remain-
jackal, (Gaubert et al., 2012; Helgen et al., 2013). On
ing, mostly terrestrial, non-pinniped carnivorans.
the other hand, the long-recognized red wolf of east-
ern North America is now regarded as an ancient
hybrid of the wolf and the coyote, with uncertain 1.3 The growth of knowledge
endangered species status (vonHoldt et al., 2016, but
Before 1900, understanding of carnivoran ecology
see Hohenlohe et al., 2017). At the level of taxo-
(as opposed to natural history) was limited, often
nomic families, the skunks and stink badgers were
based on lore and conjecture. Much of our knowl-
grouped with weasels and otters in the Mustelidae
edge of their genetics, behavior, and reproduction
before Family Mephitidae was recognized as war-
at that time resulted from observing domestic cats,
ranting recognition (Dragoo and Honeycutt, 1999).
ferrets, and dogs, and farmed minks and foxes
With each such discovery, our knowledge of car-
(Figure 1.5). Very little scientific knowledge about
nivoran classification becomes more reflective of
wild carnivorans existed, and most interest cen-
evolutionary history, as revealed through genetic
tered on the value of their furs or other body
and morphological studies.
parts and the threat they posed to agriculture. They
Carnivoran species are distributed unevenly
were difficult to study directly because of their low
across lineages and continents. Family Mustel-
densities, elusive behaviors, and constant persecu-
idae holds sixty species, whereas the Nandiniidae,
tion near humans. Well into the twentieth century,
Ailuridae, and Odobenidae hold a single species
the leading ecological questions about carnivorans
each. The Felidae, Canidae, and Mustelidae are
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