Introduction
Vincenzo Penteriani and Mario Melletti
I consider the mountainside a special place, a place with power,
as I do certain other valleys and basins . . . where grizzlies still
roam. I return to these places year after year, to keep track of
the bears and to log my life. The bears provided a calendar for
me when I got back from Vietnam, . . . I had trouble with a
world whose idea of vitality was anything other than the naked
authenticity of living or dying. The world paled, as did all that
my life had been before, and I found myself estranged from my
own time. Wild places and grizzly bears solved this problem.
Doug Peacock, 1990, Grizzly Years
Bears have fascinated people since ancient times. The relation-
ship between bears and humans dates back tens of thousands
of years, during which time we have also competed with bears
for shelter and food. Our strong link with bears is also attested
to by the Neanderthal burial of “Le Regourdou,” in France,
where the skeleton of a Neanderthal in a fetal position was
found under a funeral slab surrounded by the bones of a
brown bear, probably sacrificed for the burial. Bears were also
represented in rock paintings in caves inhabited by our ances-
tors in Europe. The bears depicted by our ancestors were cave
bears, which roamed Eurasia until about 24,000 years ago
when they became extinct during the Last Glacial Maximum.
Recently, gene flow between extinct cave bears and brown
bears has been discovered, providing direct evidence for ances-
tral hybridization between the two species which resulted in
the modern Ursus arctos that we all know (Chapter 1).
In human culture, bears also represent an important figure
in Native American mythologies. For example, the bear is a
symbol of power and strength. In fact, warriors of some tribes
wore necklaces of bear claws. Bears also play a major role in
several religious ceremonies in many North American tribes,
which used to have a bear dance as part of their tribal trad-
itions, and they represent an important clan animal in some
native cultures, for example in tribes such as the Cherokees,
Creeks, Hurons, and Navajos. Furthermore, bears are often
found carved on totem poles of several tribes of north-western
North America.
Bears have also influenced the culture of many tribes in
Asia. In fact, they are important animals for some tribes in
Siberia, and the people of the Hokkaido and Ryukyu islands
in Japan. For example, the Ainu people in Japan consider the
bear as the “Spirit of the Mountains.” In Russia, at a Fat’ya-
novo cultural site dated to around 1500 BC, necklaces made
with bear teeth were found, and other Neolithic findings have
been discovered as far north as Lake Ilmen, in the Russian
oblast of Nóvgorod. Several bear claws with bronze mounting,
dated between the ninth and eleventh centuries, were also
discovered among a Finno-Ugrian group located along the
River Tsna (a river in the Tambov and Ryazan oblasts of
Russia), whereas another Finno-Ugrian group in the Urals
venerated the bear as a symbol of heroism.
The images of bears in popular culture have helped them to
become an icon that most people know and love. The most
famous example is the teddy bear, which has been one of the
most popular stuffed animals since the early 1900s and con-
tinues to be a favorite of children. Developed almost simultan-
eously by toymakers Morris Michtom in the US and
Margarethe Steiff in Germany, and then named after US Presi-
dent Theodore “Teddy” Roosevelt, the teddy bear is an iconic
children’s toy celebrated in stories, songs, and films. More
recently, Baloo from The Jungle Book, Winnie the Pooh, Yogi
and Bubu, and Masha and the Bear tell us that the strong link
between people and bears, which started more than 80,000
years ago, continues today.
At the present time eight bear species are recognized, from
the very popular polar bear, giant panda, brown bear, and
American black bear to the lesser-known Andean bear, sun
bear, sloth bear, and Asiatic black bear. In Chapter 1, the
authors follow a different taxonomy for sun bear (Helarctos
malayanus) and Asiatic black bear (Ursus thibetanus),
ascribing them to the genus Melursus. However, we recognize
that further genetic and morphometric studies are still
required in order to fully understand the taxonomy of these
two bear species. Therefore, and pending further investigation
on Melursus taxonomy, we will still refer in this book to the
genus Helarctos for sun bear and Ursus for Asiatic black bear,
acknowledging that they might deserve a different genus in
future. The conservation issue facing some of these species is a
big conservation challenge today. Human activities, population
encroachment, and poaching in bear landscapes continue to
1
 Introduction
represent serious threats for some bear species or populations.
As an example, in more recent times, human–bear conflicts
have been exacerbated by the increasing number of people
sharing the same landscapes with bears. Such coexistence has
engendered an increase in conflicts, such as damage to live-
stock, crops, and apiaries, as well as the fear of bear attacks
(Chapters 15–17, 20, 28). Moreover, in some regions people
continue to keep bears in terrible captive conditions to extract
bile and other body parts, mainly in Asia where the trade in
these kinds of products still flourishes (Chapter 26). For
instance, in China and Vietnam, thousands of bears are kept
in tiny cages and bred for their bile. Furthermore, an emerging
threat to bear conservation is related to the effect of global
warning. This is the case, for example, for polar bears, which
are now more endangered than even a few decades ago due to
the dramatic reduction of sea ice. In fact, polar bears depend
on the ice shelf for feeding, breeding, and movement, and they
can only persist where the temporal and spatial availability of
sea ice provides adequate access to their marine mammal prey
(Chapter 21). The panda, another iconic bear species, is also
threatened by climate change. The panda is an extremely
specialized species that relies on bamboo for 99% of its diet,
occurs in a very restricted range, and has one of the lowest
reproduction rates among bears (Chapters 6 and 21). The
challenge that bear conservation and management represents
around the world makes this book extremely important and
timely because it provides informative and complete accounts
of everything you want to know on bears to a broad audience.
However, even if our general knowledge of bear ecology and
behavior has significantly increased in the past decades, we still
have a lot to learn about this group of species, and in particular
tropical bears (Chapters 4, 7–10, 22, 24–28).
The idea to edit a major volume on the ecology and conser-
vation of bears started in 2017 from the need to provide a
comprehensive book on all the bear species that inhabit our
planet, as well as a useful tool for both the general public and
people more directly involved in the fields of animal ecology,
behavior, and conservation such as researchers, wildlife man-
agers, conservationists, stakeholders (such as farmers and
hunters), and students. In our minds, this book would also
been able to demonstrate why the study of human–bear inter-
actions, and stakeholder perception and involvement, are cru-
cial for bear conservation and management, with detailed
examples and case studies from all the continents inhabited
by bears. Such a book would also highlight the urgency of
conservation actions that need to be put into practice in the
different regions of the world, mainly for lesser-known bear
species in developing countries. Additionally, and despite the
long-term interest in bears, as well as the many groups
2
working on bears for decades and the large literature on these
species around the world, a comprehensive and very detailed
book on all the bear species in the world had never been
published before. To make this idea a reality, we started by
contacting more than 250 people among the best bear biolo-
gists in the world. Many of them replied enthusiastically to our
invitation, saying that such a book was very much welcomed
and long overdue. Of course, this positive feedback gave us
further motivation to move forward with this project, in which
200 authors ultimately participated, many of them having
spent their entire lives studying bears. The authors come from
33 countries spread across five continents, and work at very
diverse institutions, such as research centers, universities,
IUCN SSC Bear Specialist Group and Polar Bear Specialist
Group and IBA, and non-governmental organizations
(NGOs). To give you an idea of the heterogeneity of the book’s
contributors, people involved in this project are from (in
alphabetical order): Bhutan, Bolivia, Canada, China, Croatia,
Denmark, Ecuador, Finland, France, Germany, Greece, Hun-
gary, India, Iran, Italy, Japan, Malaysia, Mexico, Nepal, the
Netherlands, Norway, Pakistan, Poland, Romania, Russia, Slo-
venia, Spain, Sri Lanka, Sweden, Turkey, UK, USA, and
Venezuela.
Together with the most important information on the
ecology and behavior of bear species, the volume also includes
specific chapters on taxonomy, phylogeny and genetics, popu-
lation status and trends, as well as conservation status, man-
agement, and climate changes. The book is composed of
28 chapters subdivided into four sections: Part I – Systematics,
Ecology, and Behavior (Chapters 1–5); Part II – Species
Accounts (Chapters 6–14); Part III – Human–Bear Coexist-
ence (Chapters 15–19); and Part IV – Conservation and Man-
agement (Chapters 20–28).
We hope that you will enjoy this book at least as much as
we have enjoyed its long preparation and our close collabor-
ation with chapter contributors, and that the huge effort made
by all the authors will be appreciated by the public and scien-
tific community. We will consider that our work has achieved
its goal if it is rewarded by an increase in the understanding of
bears and their effective conservation. However, the future of
bear species will depend on our capacity to find pragmatic
solutions that should represent a trade-off between human
growth and the needs of bears and their habitats. What bears,
among the most charismatic creatures on the planet, do for
people has an inestimable value for our most intimate life, and
their loss will create a void impossible to fill. Being in a bear
country captivates our minds and, at the same time, offers a
lesson in humility by giving us the feeling that something more
powerful than us is out there.
 Part I Systematics, Ecology, and Behavior
Chapter
Systematics, Evolution, and Genetics of Bears

1 Andrew C. Kitchener, Eva Bellemain, Xiang Ding, Alexander Kopatz,

Verena E. Kutschera, Valentina Salomashkina, Manuel Ruiz-Garcı́a,
Tabitha Graves, Yiling Hou, Lars Werdelin, and Axel Janke

Introduction morphological characters, including carnassial teeth, i.e. the

fourth upper premolar (P4) and first lower molar (m1), which
Bears, Family Ursidae, are among the largest of the Carnivora,
form two shearing blades for slicing through body tissues, and
but the Ursidae is among the least speciose carnivoran family
the presence of three elements in the auditory bulla: the caudal
with only eight extant species. Owing to the bears’ widespread
and rostral entotympanic and ectotympanic bones (Hunt 1998;
distribution in Eurasia, North America, and north-western
Wozencraft 1989).
South America, they occupy many habitats from the northern
Based on basicranial characteristics, the Carnivora com-
polar ice cap to tropical rainforests. Despite having few species,
prises two suborders: the Feliformia (felids, viverrids, hyaenas,
relationships between the Ursidae and other Carnivora and
and related families) and the Caniformia (canids, ursids, pinni-
between ursid species (and indeed what constitutes an ursid)
peds, and musteloids). The Caniformia comprises two Infra-
have been controversial. New molecular genetic techniques in
orders, the Cynoidea (or Canoidea), including the Canidae,
recent years have allowed relationships to be explored in new
and the Arctoidea, including the remaining caniforms. Rose
ways, leading to some clarification, but debates continue owing
(2006) characterized the Arctoidea as having a suprameatal
to incomplete lineage sorting and ancient hybridization.
fossa (a hollow in the dorso-lateral wall of the middle-ear
Bears are threatened by humans; conflict with human land
cavity) and loss of the third upper molar (M3), and that most
uses and exploitation for fur and other body parts, coupled
arctoids have a single-chambered auditory bulla comprising
with habitat loss and extirpation through direct hunting, have
mostly the ectotympanic bone. Some relationships within the
led to fragmentation of populations of all species. Molecular
Arctoidea remain controversial, although a consensus is grow-
genetics are key to understanding current and historical rela-
ing. The main contentious areas include the relationship
tionships between isolated populations, including species’ col-
between the giant panda, Ailuropoda melanoleuca, the red
onization during glacial–interglacial cycles, to determine
panda, Ailurus fulgens, and the Ursidae (see below), whether
viability of local populations, needs for habitat corridors, and
the superfamily Pinnipedia is monophyletic, and the relation-
other aspects of population management, especially where
ships between pinnipeds, the Ursidae, and Superfamily Mus-
bears are harvested for sport, etc. As natural habitats shrink,
teloidea. Traditionally, the systematics of the Carnivora were
some bear species will inevitably require high levels of man-
based on morphological studies of extant and fossil taxa (e.g.
agement, perhaps combining captive and wild populations
Wozencraft 1989), but new molecular techniques have clarified
following the IUCN’s One Plan Approach. In this chapter
many relationships, especially where homoplasy or conver-
we review the systematics of the Ursidae and its relationships
gence gave ambiguous or incorrect affinities between taxa.
with other Carnivora, the molecular phylogenetics of
However, some relationships remain uncertain, owing to
extant ursid species, the phylogeography of and morphological
incomplete lineage sorting and ancient hybridization. The
variation within each species, and the use of molecular genetics
problematic systematics of the Arctoidea are discussed below.
to monitor bear populations for management and
The Superfamily Pinnipedia includes seals (Family Phoci-
conservation.
dae), walrus (Family Odobenidae), and sea lions and fur seals
(Family Otariidae). Morphological studies since the nineteenth
Systematics of the Ursidae century suggested a sister relationship between the Ursidae and
Wozencraft (1989) summarized the systematic history of the Otariidae/Odobenidae based, for example, on the structure of
Carnivora, hence only a brief overview is given here. The the auditory bulla, and between the Mustelidae and Phocidae,
classification of the Order Carnivora based on the morphology based, for example, on the absence of an alisphenoid canal,
of the basicranium began with Turner (1848), was further thereby supporting a diphyletic origin of the Pinnipedia. How-
developed by Flower (1869), and has since been refined, e.g. ever, recent morphological studies demonstrated a monophy-
by Hunt (1974), who characterized the development and evo- letic Pinnipedia (e.g. Berta et al. 2015, but see Koretsky et al.
lution of auditory bullae. The Carnivora is characterized by 2016).

3
 Systematics, Ecology, and Behavior

Table 1.1 Classification of the extant Ursidae (Tedford 1976; Bryant 1996; Wagner 2010).

Class Mammalia Linnaeus, 1758; 1

Order Carnivora Bowdich, 1821; 33
Suborder Caniformia Kretzoi, 1943; 16, 194
Infraorder Arctoidea Flower, 1869; 15
Superfamily Ursoidea Batsch, 1788; 110
Family Ursidae Batsch, 1788; 110
Subfamily Ursinae Batsch, 1788; 110
Genus Ursus Linnaeus, 1758; 47
Ursus arctos Linnaeus, 1758; 47
Ursus maritimus Phipps, 1774; 185
Ursus americanus Pallas, 1780; 5
Genus Melursus Meyer, 1793; 155
Melursus ursinus (Shaw, 1790; pls. 58–59)
Melursus malayanus (Raffles, 1821; 254)
Melursus thibetanus (G. Cuvier, 1823; 325)
Subfamily Arctotheriinae F. Ameghino, 1902; 236
Genus Tremarctos Gervais, 1855; 20
Tremarctos ornatus (F. Cuvier, 1825)
Subfamily Ailuropodinae Grevé, 1894; 217
Genus Ailuropoda Milne-Edwards, 1870; 342
Ailuropoda melanoleuca (David, 1869; 12–13)

There is still no consensus on the relationship between the
Ursidae and other Arctoidea. Several studies showed a sister
relationship between the Ursidae and the Pinnipedia, but
excluded the Musteloidea (i.e. Mustelidae, Procyonidae, Ailur-
idae, and Mephitidae) (Luan et al. 2013), while in most others
the Ursidae was basal with the Pinnipedia and the Musteloidea
grouped together (Eizirik et al. 2010; Nyakatura & Bininda-
Emonds 2012; Doronina et al. 2015) (Figure 1.1), or grouped
the Ursidae with the Musteloidea (Luan et al. 2013).
The Ursidae comprises eight extant species, which many
authors consider to form two subfamilies (Wozencraft 1989;
Goswami 2010; Wagner 2010): the Ailuropodinae includes the
basal and highly derived giant panda, Ailuropoda melanoleuca,
which is adapted to feeding on bamboo; and the Ursinae,
which comprises two tribes, the Tremarctini, including the
Andean bear, Tremarctos ornatus, and the Ursini, comprising
the remaining six ursid species. Hunt (1998) divided the Ursi-
dae into four subfamilies: the Ursinae containing the extant
bears; the Ailuropodinae containing the giant panda; and two
extinct subfamilies, the Amphicynodontinae and the Hemi-
cyoninae. He further divided the Ursinae into three tribes –
Ursini, Tremarctini, and the ancestral Ursavini. Most recent
authors place the Andean bear in the subfamily Tremarctinae
(e.g. Soibelzon et al. 2005; Mitchell et al. 2016), but Wagner
(2010) suggested that the Tremarctinae Merriam and Stock,
1925 is pre-dated by the Arctotheriinae Ameghino, 1903, but it
actually dates to Ameghino (1902) (Table 1.1).
Currently, the extant Ursinae includes three genera:
Helarctos, for the sun bear, H. malayanus; Melursus, for the
sloth bear, M. ursinus; and Ursus, comprising the brown bear,
U. arctos, polar bear, U. maritimus, Asian black bear, U.
thibetanus, and American black bear, U. americanus. Previ-
ously, three genera were recognized for Asian bears, including
Melursus (sloth bear), Selenarctos (Asian black bear), and
Helarctos (sun bear). A recent molecular phylogenetic study
by Kumar et al. (2017) showed that the sloth and sun bears are
sister species in a clade with the Asian black bear, which is
basal. This suggests that either all Ursinae should be in the
genus Ursus as proposed previously (e.g. Wozencraft 1989;
Hunt 1998) or that the three Asian bears may be in a separate
genus, Melursus. Recently, Kitchener et al. (2017) employed
Hennig’s Rule, which discriminates genera if divergence times
are c.5 million years or more. Using Hennig’s Rule divides the
Ursinae into Melursus for Asian bears and Ursus for the rest
(Table 1.2).
The Ursidae has a primitive dental formula (I3/3 C1/1
PM4/4 M2/3) and a Type A auditory bulla (Hunt 1974),
comprising mostly the ectotympanic but also rostral and two

4
 Systematics, Evolution, and Genetics of Bears

Table 1.2 Classification of the Ursidae, including extinct taxa.

Family Ursidae Batsch, 1788

Subfamily Hemicyoninae Frick, 1926
Tribe Cephalogalini Bonis, 2013
Adelpharctos Bonis, 1971
Cyonarctos Bonis, 2013
Phoberogale Ginsburg & Morales, 1995
Filholictis Bonis, 2013
Cephalogale Jourdan, 1862
Tribe Phoberocyonini Ginsburg & Morales, 1995
Plithocyon Ginsburg, 1955
Phoberocyon Ginsburg, 1955
Tribe Hemicyonini Frick, 1926
Zaragocyon Ginsburg & Morales, 1995
Dinocyon Jourdan, 1861
Hemicyon Lartet, 1851
Subfamily Ursavinae Hendey, 1980
Ballusia Ginsburg & Morales, 1998
Ursavus Schlosser, 1899
Subfamily Agriotheriinae Kretzoi, 1929
Agriotherium Wagner, 1837
Subfamily Ailuropodinae Grevé, 1894 Figure 1.1 Phylogenetic tree showing a version of the relationships among
the Arctoidea (Koepfli et al. 2017).
Tribe Indarctini Abella et al., 2012
Miomaci Bonis et al., 2017
Indarctos Pilgrim, 1913
Tribe Ailuropodini Grevé, 1894
Kretzoiarctos Abella et al., 2012
Agriarctos Kretzoi, 1942
Ailurarctos Qi et al., 1989
Ailuropoda Milne-Edwards, 1870
Subfamily Arctotheriinae Ameghino, 1903
Plionarctos Frick, 1926
Arctodus Leidy, 1854 Figure 1.2 Dissected auditory region of a bear, showing the internal carotid
artery looped within the inferior petrosal venous sinus on the lateral edge of the
Arctotherium Burmeister, 1879 basioccipital bone. AL, alisphenoid; BO, basioccipital; BS, basisphenoid; PE,
petrosal (Hunt 1977).
Tremarctos Gervais, 1855
Subfamily Ursinae Batsch, 1788
Ursus Linnaeus, 1758 The Ursinae and Arctotheriinae share several morpho-
logical characters; a reduced lacrimal forming a vestigial bony
Melursus Meyer, 1793
rim around the naso-lacrimal foramen; a flat auditory bulla,
which is much smaller than the mastoid, squamosal, and
caudal entotympanic bones (Hunt 1998). The internal carotid basioccipital processes; reduced premolars, whereby P4 has
artery loops inside the petrosal venous sinus, covered by the lost most of its shearing function; the upper and lower molars
basioccipital, and may act as a countercurrent heat exchanger are quadrate (with four main cusps: paracone, metacone,
to cool the brain (Hunt 1998) (Figure 1.2). protocone, and a large metaconule, which migrated to the

5
 Systematics, Ecology, and Behavior
postero-internal border to form the molars’ oblong shape), and
the rear border of M2 is greatly enlarged and elongated,
forming a distinctive heel and making it the longest tooth in
the maxilla (Wozencraft 1989; Hunt 1998).
The enamel on the surface of the talonid and trigonid
basins of m2 of arctotheriines is tuberculated and forms a “z”
pattern on the talonid (Hunt 1998). The skull is shorter than
those of the Ursinae and there is a pre-masseteric fossa in the
mandible in Tremarctos and Arctodus, but not Plionarctos. The
diploid (2n) chromosome number is 52 biarmed autosomes in
Tremarctos compared with 2n = 74 mostly acrocentric
chromosomes of the Ursinae, and 2n = 42 mostly biarmed
autosomes in the giant panda (Chorn & Hoffmann 1978).
The giant panda’s classification was controversial since its
description by David (1869), who regarded it as a bear, but
Milne-Edwards (1870) classified it as a procyonid. Later authors
proposed placing it in a family (Ailuridae) with the red panda,
or in a monotypic family (Ailuropodidae). The controversy
continued as different morphological features were presented
for and against the two main hypotheses until molecular studies
demonstrated unequivocally that the giant panda is basal to the
Ursinae and Arctotheriinae. It is usually classified in the Ailur-
opodinae, although some still place it in its own family
(although not recently, e.g. Thenius 1989), owing to its highly
derived morphology and deep divergence time of c.12.5 million
years ago (Ma) (Kutschera et al. 2014).
A key morphological character of the Ailuropodinae is a
greatly enlarged radial sesamoid in the wrist, the so-called
panda’s thumb, which grasps stems, such as those of bamboo.
In most other extant ursids the radial sesamoid is not enlarged,
but recently the Andean bear’s moderately enlarged radial
sesamoid was described (Salesa et al. 2006). Therefore,
enlarged radial sesamoids are either plesiomorphic (i.e. an
ancestral character shared by two or more taxa), or they
evolved independently in primarily herbivorous ursids. Other
morphological characteristics of ailuropodines include the
highly domed skull for attachment of massive jaw muscles,
and enlarged molars and premolars for crushing tough
bamboo stems and shoots.
Fossil History of the Ursidae
There is no agreement regarding quite what a bear is, because
opinion differs as to which basal taxa should be included in the
Ursidae. For any mammal group the deeper one descends into
the fossil record, the more similar taxa become, until the origin
of the group is reached. Today’s bears differ greatly from those
of the late Eocene and Oligocene (c.35–23 million years ago).
Consequently, opinion differs as to which basal taxa should be
included in the Ursidae (Table 1.2). A group of genera, includ-
ing Amphicynodon, Amphicticeps, Parictis, Kolponomos, Allo-
cyon, and Pachycynodon, distributed in Europe, Asia, and
North America, constitute the amphicynodonts. This is either
a paraphyletic stem lineage within the Ursidae or a monophy-
letic family Amphicynodontidae that may or may not be sister
6
taxon to the Ursidae. The latter view point has gained credence
in recent years, but without consensus. Here, we consider the
amphicynodonts to constitute a distinct family and exclude
them from further discussion.
The earliest Ursidae are from the earliest Oligocene. The
family is subdivided into a number of subfamilies: Hemicyoni-
nae, Ursavinae, Agriotheriinae, Ailuropodinae, Arctotheriinae,
and Ursinae. The Hemicyoninae is the basal subfamily and
encompasses about 10 genera in three tribes. Among these is
the genus Cephalogale, often called “the earliest bear.” This
genus has recently been revised (de Bonis 2013), which showed
that traditional “Cephalogale” is an assemblage of variably
related, primitive Ursidae, within the tribe Cephalogalini,
which now comprises five genera – Cephalogale, Filholictis,
Cyonarctos, Adelpharctos, and Phoberogale – of jackal- to
wolf-sized, rather canid-like animals. Cephalogalini is known
from the earliest Oligocene (c.34 Ma) to the early Miocene (c.19
Ma). Their geographical range extended from Western Europe
to the Midwestern United States, with Cephalogale and Phober-
ogale occurring on both continents. De Bonis (2013, p. 810)
notes that no antecedents of Cephalogalini are known from the
Eocene and surmises that they (and the Ursidae) may have had
an Asian origin, where Cephalogalini are poorly known.
A second tribe of Hemicyoninae is the Phoberocyonini,
revised by Ginsburg and Morales (1998). This tribe comprises
Phoberocyon (not to be confused with Phoberogale) and Plitho-
cyon, two genera with wide geographical ranges, both known
from France to Florida. Phoberocyon is the older genus, with a
stratigraphic range encompassing the late Oligocene to late
middle Miocene (c.29–14 Ma), whereas Plithocyon is restricted
to the middle Miocene (c.16–11.5 Ma). These two genera
include larger, more derived species than the Cephalogalini,
with the largest, e.g. Phoberocyon aurelianensis, within the size
range of ursids today. Although phylogenetically close, Pho-
berocyon evolved toward hypercarnivory, with taller cusps on
the lower carnassial and a narrow, trenchant talonid, whereas
Plithocyon became more hypocarnivorous, with a low, wide
carnassial. The reasons for this ecological divergence are debat-
able and complicated by the presence of both genera on mul-
tiple continents, each with different competitors. Nevertheless,
the Ursidae would never be as hypercarnivorous again.
The Hemicyonini comprises two genera, Hemicyon and
Dinocyon. Hemicyon includes generalized, medium- to large-
sized hemicyonines of early and middle Miocene age with a
broad geographical range from western Europe (where it is
relatively common) to the midwestern United States, although
Hunt (1998) suggests that the latter (Hemicyon barbouri) rep-
resents a distinct genus. Hemicyon is also recorded from Kenya
by a single upper carnassial from the early Miocene (Schmidt-
Kittler 1987). This tooth is the only record of a bear from
Africa prior to Agriotherium in the latest Miocene (see below),
despite decades of searching at productive localities, raising
many presently unanswerable questions.
Dinocyon is much less common than Hemicyon. It is exclu-
sively known from western Europe, in sediments dating from

the middle Miocene to the earliest late Miocene (c.16–11 Ma).
Dinocyon includes some of the largest bears of all time and, at
least in tooth dimensions, rivaled giants such as Agriotherium
africanum and Arctotherium angustidens (see below).
Closer to crown-group Ursidae, relationships between sub-
families are a little better understood (Abella et al. 2012),
although the first, Ursavinae, is clearly a paraphyletic stem
lineage. It comprises the genera Ballusia and Ursavus. The
former is exclusively European from the early Miocene,
whereas the latter comprises several species from the early to
late Miocene of Eurasia and the middle Miocene of North
America. Ursavus is traditionally considered ancestral to more
derived bears. They are all small to medium-sized animals with
generalized omnivore characteristics.
Following the Ursavinae is the subfamily Agriotheriinae,
which includes only the genus Agriotherium. A. africanum is the
first unequivocal bear to be described from sub-Saharan Africa,
by Hendey (1980). Until that time, bears were only thought to
have reached North Africa in the late Pleistocene, but never to
have crossed the Sahara. Agriotherium africanum was consider-
ably larger and more carnivorous than modern Ursus.
Crown-group Ursidae (the common ancestor of all living
bears and all of its descendants) includes three subfamilies. The
Arctotheriinae encompasses the South American ursid radi-
ation, with Tremarctos ornatus as its surviving representative.
The arctotheriines have antecedents in North America, where
the ancestral taxon is Plionarctos of the late Miocene to Pliocene.
Tremarctos also originated in North America, where T. florida-
nus overlapped in time with Plionarctos. However, most char-
acteristic of the arctotheriine radiation are the short-faced bears,
Arctodus (Pliocene of North America) and Arctotherium (Pleis-
tocene of South America). Among the latter, Arctotherium
angustidens may have been the largest bear (and carnivoran)
of all time, with an estimated body mass of up to 1500 kg
(Soibelzon & Schubert 2011). Curiously, the South American
short-faced bear was more closely related to the living Andean
bear (which is not short-faced) than it was to the North Ameri-
can short-faced bear. This is a striking example of convergence
in size and morphology to ecological circumstance.
The Ursinae includes the extant genera Melursus and
Ursus. The antecedents of Melursus are poorly known except
for the Asian black bear. However, the genus Ursus is
extremely well known in the fossil record, particularly the cave
bear, U. spelaeus, which is studied from many thousands of
specimens across large parts of western Eurasia, where it
hibernated (and died) in caves. Numerous other species of
Ursus have been described, including fossil relatives of the
American black bear.
The final subfamily, the Ailuropodinae, includes the giant
panda and relatives. The palaeontology of this subfamily has
progressed greatly recently (Abella et al. 2012; de Bonis et al.
2017). The giant panda, once thought to be very isolated
phylogenetically among the Ursidae, is now known to be
among a large diversity of genera and species in the Ailuropo-
dinae. Two tribes are recognized, the Indarctini and the
Systematics, Evolution, and Genetics of Bears
Ailuropodini. The former includes the early genus Miomaci
of the late Miocene in Europe and Indarctos, a genus known
from the late Miocene of Eurasia and North America. The
Ailuropodini comprises several genera, the oldest and most
primitive of which are the European Kretzoiarctos and Agriarc-
tos of late middle and early late Miocene age. Together with the
Chinese Ailurarctos, these genera form a clade that is sister to
Ailuropoda, culminating in the extant giant panda. This lin-
eage shows a strong trend toward increased size and special-
ization to herbivory.
Molecular Phylogenies
The relationships among today’s bear species have been
unclear or contradictory depending on which methods were
used. In particular, the relationships between the six ursine
species remained unresolved until recently, because early
molecular studies contradicted each other, mostly because of
insufficient data (Talbot & Shields 1996; Yu et al. 2004). How-
ever, even larger molecular data sets that included nuclear
genes did not confidently resolve the bears’ phylogeny (Pages
et al. 2008; Kutschera et al. 2014), resulting in a “forest of gene
trees” with contradicting phylogenies. This suggested that bear
evolution was either shaped by incomplete lineage sorting or
that ancient hybridizations complicated phylogenetic recon-
struction (Kutschera et al. 2014).
The first analyses of whole-genome data from all ursine
and arctotheriine species have recently resolved relationships
among bears (Figure 1.3; Kumar et al. 2017). Previously, the
position of the American and Asian black bears with respect to
their relatives was much debated (Pages et al. 2008; Kutschera
et al. 2014). In the bifurcating tree (Figure 1.3), the American
black bear is the sister species to polar and brown bears, while
Figure 1.3 Phylogenomic analyses of 18,621 maximum likelihood trees
estimated from non-overlapping 100-kb genome fragments from whole
genome data of seven bear species (Kumar et al. 2017). (A) Schematic of
coalescent species tree with 100% bootstrap support of all branches.
Divergence time estimates in million years ago (Ma) from Kumar et al. (2017)
were mapped onto the tree that was obtained from an analysis of 5.2 million bp
coding sequences in MCMC tree (PAML) with multiple calibration points. (B)
Split network analysis at a 7% threshold level. Ma, million years ago; ABC, brown
bear from the Admiralty, Baranof, and Chichagof (ABC) Islands. Paintings by Jon
Baldur Hlidberg (www.fauna.is). Modified from Kumar et al. (2017), figure 1.3,
licensed under CC BY 4.0.
7
 Systematics, Ecology, and Behavior
sun and sloth bears are grouped with the Asian black bear. The
Arctotheriinae forms the sister group to all ursine bears.
The giant panda was consistently placed as sister group to
the Ursinae/Arctotheriinae in previous phylogenetic analyses
(e.g. Kutschera et al. 2014).
However, analyses of bear genomes have shown that gene
flow among species typifies the bears’ evolutionary history
(Kumar et al. 2017). Although harder to interpret, the bears’
evolutionary radiation is actually a network, where signals
from deviating phylogenetic trees become evident, with the
American black bear placed between Asian bears and polar/
brown bears (Figure 1.3B; Kumar et al. 2017). A brown bear
from the Admiralty, Baranof, and Chichagof (ABC) Islands,
off the western Canadian coast, is positioned between polar
and brown bears, consistent with gene flow between ABC
brown bears and polar bears (Miller et al. 2012; Cahill et al.
2015).
This network represents best the genome-wide evolution-
ary signal among extant ursids (Bapteste et al. 2013), because
frequent hybridization among bears means their genomes
are mosaics of evolutionary histories. A similar mosaic can
result from incomplete lineage sorting, where ancestral gene
lineages are randomly sorted along the species tree, and where
phylogenetic signals may also differ from the species tree
(Pamilo & Nei 1988). Only if all phylogenetic information in
the evolutionary network is forced into one bifurcating tree is
the more conventional view of bear evolution recovered
(Figure 1.3A). While this tree may serve many purposes, such
as taxonomy, conservation genetics, or molecular clock ana-
lyses, it provides an incomplete understanding of the bears’
evolutionary history and it may also explain why some
morphological characters are difficult to place on a simple,
bifurcating tree.
Genetic differences among the bears can be converted into
time, assuming that substitutions accumulate relatively con-
stantly between species, as in a molecular clock (Zuckerkandl
& Pauling 1962). Molecular divergence times among bears
have been studied extensively (e.g. Kutschera et al. 2014;
Kumar et al. 2017), with particular interest in the divergence
of polar and brown bears (e.g. Hailer et al. 2012; Miller et al.
2012). Genome-wide estimates suggest the deepest split in the
Ursinae (Ursus versus Melursus) was 5.0 (4.5–6.0) million
years ago (Ma), and the divergence of polar and brown bears
was 0.9 (0.6–1.1) Ma (Kumar et al. 2017).
Phylogenetic reconstruction of bears exemplifies the com-
plications that gene flow creates for understanding evolution-
ary relationships. One of the first molecular studies on bear
species used mitochondrial DNA (mtDNA) to study the rela-
tionship between polar and brown bears (Cronin et al. 1991).
This analysis suggested that polar bears originated from brown
bears from the ABC Islands, questioning their validity as a
distinct species and leaving Ursus arctos paraphyletic. How-
ever, some 20 years later, a comprehensive analysis of nuclear
gene loci demonstrated that polar bears are a separate, ancient
lineage that diverged from the common ancestor of polar and
8
brown bears about 600,000 years ago (ya) (Hailer et al. 2012).
Therefore, the mtDNA of ABC Islands brown bears probably
transferred to polar bears by hybridization, making it the first
example of gene flow among bear species. For this transfer of
mtDNA, female brown bears and male polar bears must have
hybridized some 150,000 ya, coinciding with the penultimate
glacial period (c.185,000–135,000 ya; Marine Isotope Stage 6;
Margari et al. 2010). The brown bear mitochondrial haplotype
has been identified in hundreds of polar bears studied for this
locus so far (e.g. Cronin et al. 1991; Hailer et al. 2012). Under
this scenario, the brown bear mitochondrial haplotype spread
to all polar bears in a comparatively short time, also implying
that brown bear mtDNA was selectively advantageous for
polar bear survival, or the population bottleneck was very
severe. An alternative explanation for high genetic similarity
of polar and brown bear mtDNA is that gene flow occurred
from polar bears into some brown bear populations (Cahill
et al. 2013, 2018; Liu et al. 2014; Hassanin 2015). Indeed, gene
flow between polar and brown bears was repeatedly detected in
analyses of entire genomes (e.g. Miller et al. 2012; Cahill et al.
2015; Kumar et al. 2017). Up to 8.8% of the brown bear
genome is of polar bear ancestry, but polar bear genomes
harbor no significant genetic introgression from brown bears
(Cahill et al. 2015). Cahill et al. (2018) suggested that brown
bears carried some polar bear alleles away from where admix-
ture occurred through male-dominated dispersal. It is unclear
if this gene-flow asymmetry reflects continuing hybridization
between the two species, is a remnant from the speciation
event, or results from strong selection pressure on the highly
specialized polar bear.
Genome data from all bear species show that gene flow was
much more common than assumed for mammalian genera
(Kumar et al. 2017). Whether hybridization is current or
ancestral among bears, different parts of their genomes have
different evolutionary histories. This complicates our under-
standing of bear evolution. For example, some parts of the
American black bear’s genome support a close relationship
with the polar and brown bears, while other parts place the
American black bear closest to the Asian black bear
(Figure 1.3B). Phylogenetic reconstruction among the Asian
bears is even more complicated and consistent with natural
hybridization between Asian black bears and sun bears (Gal-
breath et al. 2008). Technical advances in analyses of ancient
DNA allow studies of entire genomes of extinct species.
Recently, a study found gene flow from extinct cave bears,
Ursus spelaeus, into brown bears (Barlow et al. 2018), provid-
ing direct evidence for ancestral hybridization. Such interspe-
cific gene flow might have been advantageous for adapting to
changing environments.
Phylogeography and Intraspecific Taxonomy
Although relationships between species are now better under-
stood, relationships between populations within species and the
taxonomic status of those populations are often unresolved in

the Ursidae despite descriptions of many different subspecies
and even species. For example, 232 recent and 39 fossil species
and subspecies of today’s Ursus arctos have been described (see
Erdbrink 1953; Hall 1981), which Kurtén and Anderson (1980)
considered “a waste of taxonomic effort, which, as far as we
know, is unparalleled.” Phylogeographical studies allow better
understanding of how populations are related to each other,
where refugia existed during glaciations, and how species have
recolonized continental ranges. However, lack of genetic and
morphological data and their frequent apparent incongruence,
along with high intraspecific variability, have led to poor reso-
lution of intraspecific taxonomy and relationships in some
species. For example, polar and Andean bears are considered
monotypic, despite geographical variation in size and some
genetic structuring of populations, although ESUs (evolution-
ary significant units) or MUs (management units) are recog-
nized owing to genetically discrete populations, which occupy
different habitats or experience different climatic conditions. In
contrast, 8–12 extant brown bear subspecies are described;
some are historically and widely recognized, but recent ones
are not fully accepted, and for some, their geographical ranges
are poorly known. Below, we briefly review the phylogeography
and current status of subspecies within each bear species.
Ailuropoda melanoleuca
The giant panda was widely regarded as monotypic until Wan
et al. (2003) described A. m. qinlingensis, from the Qinling
Mountains, Shaanxi Province, based on genetic differentiation
from A. m. melanoleuca in Sichuan. This proposed new sub-
species appears to have a browner pelage and smaller skull
with larger molars (Wan et al. 2005). However, the distribution
gap between the two putative subspecies is caused by human
impacts, so genetic differences may represent genetic drift in
now separate populations or clinal variation in a previously
continuous range.
The giant panda’s phylogeography shows a mismatch
between mitochondrial and nuclear genetic studies. Early stud-
ies of short mtDNA sequences showed no haplogroups, even
using longer sequence lengths up to 680 base pairs (bp) (Zhang
et al. 2007; Hu et al. 2010a), which distinguish brown bear
subclades. Recent studies, using complete mitochondrial
genomes, revealed at least three extant and two extinct clades
in giant pandas (Barlow et al. 2019). However, although mito-
chondrial clades are well defined and calibrated, they are not
restricted to any population and show no geographical subdiv-
ision. In contrast, nuclear DNA reveals quite finely resolved
geographical patterns with the Qinling Mountains, home to
the most divergent population (Zhao et al. 2013), and Xiaox-
iangling as a probable refugium (Chen et al. 2013). This
discordance in genetic markers probably results from female-
biased dispersal (Hu et al. 2010b). Also, considering drastic
range decreases and population fragmentation, it is probable
that many other genetic lineages are extinct, which could have
highlighted the species’ history and phylogeography.
Systematics, Evolution, and Genetics of Bears
Tremarctos ornatus
Although subspecies were proposed previously (García-Rangel
2012), today the Andean bear is regarded as monotypic. How-
ever, genetic studies show that Andean bears comprise two
ESUs: the Northern Andean clade (NAC; Venezuela,
Colombia, Ecuador, and north-central Peru) and the Southern
Andean clade (SAC; southern Peru and the northern and
central Bolivian Andes), which split around 500,000 ya
(Ruiz-García 2013; Ruiz-García et al. 2020a). Additionally, in
Bolivia, some Andean bears from Santa Cruz Department were
more related to the NAC than the SAC. These results and the
slightly higher-level genetic diversity in the SAC show that the
latter was the ancestral Andean bear population, contradicting
palaeontological finds, which support the most northern
Andean bear population as ancestral. Haplotypes vary in how
they spread through the Colombian Andean Cordilleras.
Gene-flow estimates were relatively high; therefore, geograph-
ical barriers have not prevented dispersal of Andean bears
there. However, significant genetic heterogeneity was found
in northern Colombian populations, with significant spatial
autocorrelation for the Andean bear in Colombia (Ruiz-García
et al. 2020b). In contrast, no genetic differentiation was found
between different Cordillera Provinces, and northern–
southern regions in Ecuador (Ruiz-García et al. 2020c).
The first molecular genetic studies of Andean bears used
nuclear markers (microsatellites; Ruiz-García 2003, 2007;
Ruiz-García et al. 2005) and estimated low to medium genetic
diversities. However, mitochondrial markers yielded high
levels of genetic diversity. This paradox is probably explained
by “ascertainment bias” (Ellegren et al. 1995), whereby the
species for which microsatellites were developed shows higher
genetic diversity than that of the non-target species. In this
case, the microsatellites used for Andean bears were originally
designed for American black bears (Kumar et al. 2017).
Melursus malayanus
Usually two subspecies of sun bears are recognized: M.
m. malayanus on the Asian mainland and Sumatra, and M.
m. euryspilus from Borneo (Corbet & Hill 1992). Meijaard
(2004) analyzed skull morphometrics of sun bears and con-
firmed that the Bornean population, which has a smaller skull
and relatively longer upper tooth row, is distinct from Suma-
tran and mainland populations.
Very little is known about sun bear genetics. Meijaard
(2004) cited an unpublished mtDNA study by L. Waits on
sun bears from Sumatra and Borneo that yielded five clades
with no geographical structure, suggesting some gene flow
between islands during the Last Glacial Maximum (LGM).
From published data, two mtDNA lineages are reported from
Borneo (divergence 5.6 ± 1.9 base pairs (bp)), but no geo-
graphical separation (Onuma et al. 2006). Yu et al. (2007) and
Krause et al. (2008) obtained mtDNA sequences from three
putative continental Asian sun bears that could represent
another subspecies, and which yielded two additional
9
 Systematics, Ecology, and Behavior
haplotypes that diverged from the two Bornean lineages by
14.39 ± 3.39 bp and 16.26 ± 3.57 bp, respectively. The differ-
ences between these mtDNA clades suggest a robust structure
exists, dividing continental and island bears, but this is not yet
fully resolved.
Melursus ursinus
Usually two subspecies of sloth bear are recognized: M.
u. ursinus from peninsular India and M. u. inornatus, which
has a smaller skull, from Sri Lanka (Corbet & Hill 1992).
However, sample sizes for cranial lengths were very small.
There may be clinal variation in skull size with latitude from
larger northern animals to smaller southern animals, but this
has not yet been studied.
The sloth bear’s phylogeography has not been studied.
However, in GenBank there are seven sequences of mtDNA
fragments, including Zhang and Ryder 1993 (captive); Yu et al.
2007 (no locality); Talbot and Shields 1996 (captive); Mohan
et al. unpublished (India); and Krause et al. 2008 (no locality).
Not all these sequences could be aligned, although a compari-
son reveals at least three highly divergent haplogroups, lacking
any geographical information.
Melursus thibetanus
Asian black bears are widely distributed in southern and east-
ern Asia; several subspecies are usually recognized, including
M. t. thibetanus from Nepal to SE Asia, M. t. japonicus from
Japan, M. t. formosanus from Taiwan, M. t. gedrosianus from
Pakistan, M. t. ussuricus from the Russian Far East, Korea and
northern China, M. t. mupinensis from southern and central
China, and M. t. laniger from Kashmir and Punjab (Ellerman
& Morrison-Scott 1953).
The Asian black bear’s phylogeographical structure, based
on mtDNA, is robust, but not yet fully resolved. It includes
several haplogroups roughly congruent with seven putative
subspecies (Hwang et al. 2008; Kadariya et al. 2018). The
Japanese subspecies forms a distinct basal mitochondrial clade
(Wu et al. 2015; Kadariya et al. 2018), which probably colon-
ized Japan in the Middle Pleistocene, and is further substruc-
tured among the Honshu bears (Ohnishi et al. 2009; Kim et al.
2011). The mainland Asian clade probably represents a latitu-
dinal cline, with increasing adaptation to more tropical cli-
mates from north to south, including the Taiwanese
population, but the exact phylogenetic order of the matrilines
is uncertain (Ohnishi et al. 2009; Kim et al. 2011; Wu et al.
2015; Kadariya et al. 2018). Recently, Kadariya et al. (2018)
showed that the Himalayan population is a distinct basal
lineage to mainland Asian populations. This suggests that
three subspecies are recognizable: M. t. thibetanus from
mainland Asia, M. t. laniger from the Himalayas, and
M. t. japonicus from Japan. This species’ phylogeographical
structure is consistent with female philopatry and it seems very
likely that it was influenced by the Pleistocene and Holocene
glaciations throughout its range.
10
Although the Ussuri black bear (known as M. t. ussuricus)
is separated by more than 1500 km from other populations, it
is not genetically distinct. The Ussuri population arose very
recently, separating c.20,000 years ago (peak LGM), and
adapting to colder climates. The distribution gap, cf. the tiger,
Panthera tigris, arose even later, owing to forest loss since the
early Neolithic, c.9000 years ago, and the exploitation of bears
for traditional medicines by early civilizations in northern
China since at least the Shang dynasty c.3500 years ago (Barnes
1999; Ren 2000). Owing to its ecological distinctiveness and
geographical isolation, the Ussuri population should be con-
sidered as a separate MU.
Ursus americanus
Owing to its widespread distribution, coupled with polychro-
matism (Rounds 1987), many subspecies of U. americanus (16
listed by Hall 1981) were recognized, with U. a. americanus
occupying most of the species range.
For American black bears phylogeographical structures are
described for both mitochondrial and nuclear DNA (Wooding
& Ward 1997; Pelletier et al 2011; Puckett et al. 2015). Analysis
of mtDNA revealed two lineages; the continental lineage is
subdivided into eastern and western subclades, while the
coastal lineage has a much simpler structure along the Pacific
coast of North America. Nuclear SNP (single nucleotide poly-
morphism) data also showed quite clear geographical struc-
ture, including three main continental nuclear clusters (Alaska,
Eastern, and Western), which were divided into nine subdiv-
isions (Puckett et al. 2015). However, nuclear and mitochon-
drial clusters mostly did not coincide. The current
phylogeographical structure is believed to have originated
before the LGM [31–67 kya (thousand years ago) for nuclear
clusters and 169 kya to 1.07 Ma for the mitochondrial ones] in
several refugia in northern and southern North America,
including Beringia, the north-west Pacific, the Southwest, and
the Southeast (Puckett et al. 2015).
Neither mitochondrial nor nuclear structures match sub-
species delineation very well (Puckett et al. 2015). Indeed, it is
difficult to discern subspecies owing to complex admixture
following expansion from these refugia, although Puckett
et al. (2015) suggested that the nine subclusters could be
associated with subspecies, which still seems too many for a
highly mobile species showing extensive admixture. Perhaps
the three mtDNA clusters, which appear mostly geographically
discrete, would be better recognized as subspecies, including
U. a. americanus (clade A-east) in the east, U. a. emmonsii in
the west (clade B) and U. a. cinnamomum (clade A-west) from
the central range.
Ursus maritimus
The polar bear is a monotypic species, although subspecies
were previously proposed. Wilson (1976) found clinal vari-
ation in skull morphometrics across North America, with the
possible exception of a distinct population in South Alaska,

which was proposed as a subspecies. Currently, 19 MUs are
recognized (Durner et al. 2018).
For the polar bear, mtDNA reveals no clear genetic struc-
ture, although particular locations may differentiate in haplo-
type frequencies (Campagna et al. 2013; Kutschera et al. 2016).
This lack of structure is partially attributed to long-distance
movements of individuals and partially to the relatively recent
origin/divergence of the species’ mtDNA lineages (see above).
In contrast, microsatellite and SNP data show four to six
geographical clusters, which are consistent between studies
(Kutschera et al. 2016; Malenfant et al. 2016) and show differ-
entiation between adjacent populations (Crompton et al. 2008;
Campagna et al. 2013; Viengkone 2016). Putative populations
are often identified as MUs, although microsatellite clusters
have broader geographical ranges and may include several
putative MUs. A comprehensive reconstruction of the species’
history, cf. brown bear and American black bear, is prevented
by current unavailability of sufficient genetic markers with
enough phylogenetic signal.
Ursus arctos
The brown bear is the most widespread ursid species and,
given the wide variety of habitats it occupies, it is unsurprising
that its morphology is so variable. As a result, many subspecies
have been recognized, but the principal extant ones are: Eur-
asian brown bear, U. a. arctos; Syrian bear, U. a. syriacus;
Isabelline bear, U. a. isabellinus; blue bear, U. a. pruinosus;
Kamchatkan bear, U. a. beringianus/piscator; Siberian brown
bear, U. a. collaris; Japanese brown bear, U. a. lasiotus; Kodiak
bear, U. a. middendorffii; and grizzly bear, U. a. horribilis
(Ellerman & Morrison-Scott 1953). However, brown bears
show much individual, seasonal, age, and sexual variation,
and thus it can be challenging to ascertain subspecies tax-
onomy based on morphology alone.
Although there have been many phylogeographical studies
on brown bears, very few used nuclear DNA, so phylogeogra-
phical patterns often reflect female natal philopatry. The phy-
logeographical structure of the brown bear is well resolved.
Several mitochondrial haplogroups are recognized, some of
which are widely distributed and sympatric, whereas others
are restricted to particular geographical regions. The picture is
further complicated (see above), because polar bears share a
mitochondrial DNA haplogroup with ABC Islands brown
bears and an extinct Irish population (Edwards et al. 2011).
Some studies found that current phylogeographical patterns
do not match distributions of mtDNA haplotypes from fossil
specimens (e.g. Paetkau et al. 1998; Barnes et al. 2002; Hofrei-
ter et al. 2004; Valdiosera et al. 2007; Ersmark et al. 2019), such
that today’s patterns result from female philopatry, population
expansion and fragmentation, and genetic drift. MtDNA
reveals at least six major clades grouped into two main
branches. The first branch includes Western European (clade
1) and the Iranian clade from south-west Asia and the ABC
Islands (clade 2; including the polar bear) (Davison et al. 2011;
Systematics, Evolution, and Genetics of Bears
Ashrafzadeh et al. 2016; Salomashkina et al. 2017). The second
branch includes bears from southern North America and
Hokkaido (clades 4 and 3d), Tibet (clade 5), and superclade
3, which occurs throughout most of the Eurasian and North-
ern American species range and is divided into two (3a and 3b)
to four (3a, 3b, and two Caucasian) subclades (Murtskhvaladze
et al. 2010; Çilingir et al. 2016; Salomashkina et al. 2017)
(Figure 1.4). There is also a recently discovered Himalayan
clade that appears to form a third main branch (Lan et al.
2017). Particular traits of the brown bear’s phylogeography
allow reconstruction of its complex colonization history,
which was heavily influenced by glacial cycles, involving mul-
tiple cycles of range contractions followed by lineage sorting
and subsequent range expansions (Davison et al. 2011; García-
Vázquez et al. 2017; Anijalg et al. 2018). Therefore, care should
be taken in making taxonomic decisions based on contempor-
ary genetic data alone without a deeper time perspective.
MtDNA reflects matrilinear history, but it is not directly con-
nected with environmental adaptation, and thus in some cases
animals inhabiting the same area may morphologically evolve
into a homogeneous population while still carrying very diver-
gent matrilines through different colonizations and subsequent
population introgression. For example, before data on their
mtDNA were available, the genetically distinct ABC bears were
not a recognized subspecies, while clearly morphologically
distinct bears from Kamchatka and Kodiak were recognized
as subspecies, but are just branches inside the 3a subclade. The
Syrian bear comprises three major mtDNA clades (and seven
subclades), and the Siberian brown bear seems to be a mixture
of two. The recognition of local populations in the Apennines
(U. a. marsicanus) and the Pyrenees (U. a. pyrenaicus) cannot be
supported based on current genetic evidence (Loy et al. 2008;
Colangelo et al. 2012).
Paternal phylogeography is a new genetic approach that
could be applied to this species, which is based on SNPs and
short tandem repeats (STRs) located in the non-recombining
fragment of the Y-chromosome (Bidon et al. 2014; Hirata et al.
2017). However, there are currently insufficient data for a
comprehensive coverage of the brown bear’s widespread range
to understand its geographical structure.
Population Genetic Assessment and
Management of Bear Populations
for Conservation
Conservation genetics based on non-invasive genetic sampling
has revolutionized how individuals, groups, and populations of
bears are characterized and monitored (Waits et al. 1999;
Schwartz et al. 2007). The majority of population genetic
analyses in bears used microsatellites or STRs. However,
markers based on genomic approaches, such as SNPs and
genome sequencing of individuals, have been increasingly
applied. Substantial improvements in methodology have led
to the extensive use of non-invasive genetic sampling to
11
 Systematics, Ecology, and Behavior

Figure 1.4 Bayesian phylogenetic tree of brown bears based on mitochondrial genomes (Anijalg et al. 2018). Numbers on nodes and after clades represent
posterior mean estimates of the ages of different lineages, with numbers in parentheses representing 95% credibility intervals. Vertical gray lines represent long
glacial periods. The recently described Himalayan brown bear clade is not shown (Lan et al. 2017), but is basal to clade 1, Western lineage.

identify and determine the numbers of bears in areas, coun-
tries, or even transborder regions (Bischof et al. 2016). Non-
invasive samples are used for genetic assessments of popula-
tions, including estimation of genetic diversity and substruc-
ture, inbreeding, effective population size and thus population
viability, and conservation status. Further, regular sampling
allows development of databases containing many individual
genotypes and also comprehensive ecological monitoring data
of the same individuals over several years [e.g. Norway (see
Figure 1.5) and the USA; see Chapter 13].
DNA-Based Monitoring of Bears
Today, many techniques are used in DNA research and moni-
toring of bears. Different genetic markers and assessments may
show different results owing to the resolution of genetic data.
Wildlife researchers and managers must define a specific gen-
etic sampling and monitoring scheme, and choose the genetic
assessment method which fits their needs and delivers the
information needed to achieve management or research goals.
Early studies of populations were based on high-quality DNA
samples from blood or tissues collected from captured or dead
bears. From such samples many genetic markers can be used
to identify individuals, ascertain sex and assess genetic diver-
sity, structure, kinship, and gene flow among individuals and
populations, or to obtain long fragments of mtDNA to study
phylogeny.
When studying large, small or endangered bear popula-
tions, or when budgets are limited, non-invasive sampling
(feces, hairs) provides opportunities to obtain genetic data
without disturbing or even observing individuals and may be
less expensive. The collected samples are usually of low-quality,
somewhat degraded DNA, which constrains the number of
markers that can be studied and the length of the studied
DNA fragment, and therefore options for population genetic
analyses (see Table 1.3). However, when samples are successfully
genotyped, individual identification allows estimation of home
ranges, population size through capture–mark–recapture

12
 Systematics, Evolution, and Genetics of Bears

Figure 1.5 The Norwegian Large Predator Monitoring Program operated by Rovdata. Rovdata is responsible for the analyses and reports the results and monitoring
data on brown bears in the country. Roughly 1000 non-invasive samples (mainly feces and hairs) are collected annually to assess the number of bears to guide the
management by estimating the annual reproductive output (Bischof & Swenson 2012). Results of individual identifications, and other data, are accessible to the public
at www.rovbase.no. From 2016 to 2019, for example, 2867 feces samples were collected and analyzed: successfully genotyped samples (n = 1770; black), negative
samples (n = 1097; gray); figure by Kopatz et al. (in preparation, modified).

13
 Systematics, Ecology, and Behavior

Table 1.3 Applications for DNA-based studies of bears depending on the type of sample obtained. x, commonly observed in the literature; (x), observed in
recent studies or potential application.

Tissue Fecal Ancient DNA eDNA samples/

samples samples/hairs samples (mixed genetic material)
Species (mitochondrial Species x x x x
DNA) identification
Lineage x x x
identification
Interspecific x (x)
phylogeny
Individual (nuclear DNA) Genotyping x x (x)
Sex identification x x (x)
Genetic diversity x x
Abundance x x
estimates
Structure/gene flow x (x)
Kinship analyses x (x)

methods, or population trends of identified groups or subpo-
pulations based on genetic parameters (e.g. Kendall et al. 2009).
Importantly for isolated and endangered species and popula-
tions, such genotyping data can also be used to identify MUs
or ESUs.
In scenarios where collection of tissues, feces or hairs is
impossible, species or even individuals can be identified from
environmental DNA (eDNA) samples, i.e. biological material
collected from substrates such as soil, water, or snow that bears
have encountered. Several recent studies applying eDNA tech-
nologies in various conditions and environments highlight the
potential for this technology. For example, Wheat et al. (2016)
collected saliva swabs from salmon carcasses and showed that
this is a more cost-effective, rapid method to identify individual
bears than the use of scats. Von Duyke et al. (2017) simulated
fresh tracks by pressing the foot of a subsistence-harvested polar
bear into snow. The genotype from the snow sample, based on
next-generation sequencing (De Barba et al. 2017), matched the
genotype obtained from this harvested bear. Further technical
developments might allow researchers to obtain bear DNA from
tracks in the field, from different substrates (such as soil around
the den), or indirectly through invertebrate- or parasite-derived
DNA (iDNA; Abrams et al. 2019). The possibility of identifying
individuals with these new approaches could revolutionize the
monitoring and management of bear populations.
Population Genetic and Genomic Assessments
Genetic diversity provides a metric of genetic health, an evalu-
ation of historical reductions in population size, and insights into
the potential for long-term viability through adaptive capacity.
Genetic diversity varies considerably across bear populations and
is affected by the number and variability of loci along with sample
size. For example, in brown bears, heterozygosity ranges from
0.27 to 0.81 (Skrbinšek et al. 2012). As expected, low diversity
occurs in small and isolated populations (e.g. brown bears in the
Gobi Desert, Cantabrian Mountains, and Kodiak Island; Andean
bears in Ecuador). Theoretical concepts and methodology to
estimate effective population sizes in natural populations have
been evaluated (Skrbinšek et al. 2012; Kamath et al. 2015; Kopatz
et al. 2017) to determine whether they would be practical to assess
and track the number of individuals effectively contributing to
populations (Pierson et al. 2018). A study on a population of
grizzly bears in Greater Yellowstone Ecosystem, USA demon-
strated that numbers of bears and effective population size have
increased, despite relatively low and stable genetic diversity
(Kamath et al. 2015). While limited evidence for deleterious
inbreeding effects exists in natural populations (but see Dunbar
et al., 1996), decreased litter sizes and albinism have been dem-
onstrated in captive brown bears (Laikre et al. 1996). Similarly,
male Scandinavian brown bears with high interrelatedness had
lower yearly reproductive success (Zedrosser et al. 2007), and, in
the Apennine mountains, alleles predicted to be deleterious were
fixed in brown bears, including several associated with health
problems in humans (Benazzo et al. 2017). Low diversity has also
been demonstrated in the captive-bred giant panda population
(Shen et al. 2009) and genetic bottlenecks have been illustrated for
American black bears (Murphy et al. 2017). However, genetic
diversity can increase rapidly when populations and genes mix
(e.g. Mikle et al. 2016).
The importance of isolation has led many studies to assess
population structure, often in combination with assignment
tests and coalescent approaches, to evaluate connectivity
between subpopulations. For example, Dutta et al. (2015)
identified migrants between populations of Indian sloth bears.

14
 Systematics, Evolution, and Genetics of Bears

Many studies use landscape–genetic methods to evaluate
resistance of various landscape features, with the aim of man-
aging landscapes to maintain or enhance connectivity between
separated subpopulations. Draheim et al. (2018) found that
American black bears showed a stronger signal of isolation by
resistance over time as forest fragmentation increased. Parent-
age approaches have been used to evaluate movement (e.g.
Graves et al. 2014), as well as to understand the impact of
interactions of behavior and relatedness on fitness, through
mechanisms such as multiple paternity, sexually selected
infanticide, and site fidelity (e.g. Norman et al. 2017). Analyses
of population structure are also used in forensics to identify
the source of illegally killed individuals or bear body parts
(Frosch et al. 2014). With the development of genomics, new
approaches to diversity, inbreeding, and other questions are
becoming more common, as well as the ability to evaluate the
role of selection and adaptive capacity in population survival
and evolution. Genetic information enables studies on related-
ness and kinship. For example, Mikle et al. (2016) used a
pedigree approach to document mechanisms behind changes
in diversity, where initial low diversity in a population resulted
from a few reproductively dominant individuals, but increases
in migration between subpopulations over time led directly to
increased diversity. Norman et al. (2017) developed a method
to identify fine-scale population genetic structure, which is
sensitive to spatial and temporal changes in the landscape.
Conclusions
Despite the few extant ursid species, the evolution of bears has
been relatively well studied. This is partly because of their large
size, so they are common in the fossil record, but also because
they compete for space and resources with people, and have
been and continue to be targets for direct exploitation, includ-
ing entertainment, sport hunting, and to provide fur and other
tissues for human use. In recent years genetics have provided a
better understanding of relationships among bear species,
which are complicated owing to ancient hybridizations, and
with other carnivorans, although some uncertainties remain.
Genetics have also improved our knowledge of variation
within species, some of which, such as the brown bear, have
very extensive geographical ranges, so that species’ histories
and colonizations can be resconstructed in deeper time by
comparison with the fossil record.
Genetics are vital for the future successful conservation and
management of bear populations, especially with the advent of
new techniques in recent years. Whereas the population gen-
etics of the European brown and grizzly bears, as well as the
American black bear to some extent, are quite well studied in
terms of population genetics, there are few genetic assessments
for other bear species, e.g. sun bears, sloth bears, and Asian
black bears. Increased human expansion and a changing cli-
mate will cause further fragmentation, stress populations, and
endanger bear species across the world, which will challenge
current conservation, management, and research. Therefore,
the genetic assessment of bear populations and their status and
connectivity will increase in importance. Attempts to rewild
certain areas may include the release of bears, which should
incorporate genetic monitoring programs. Bears will continue
to play an important role in genetic research because they
(especially brown bears) are ecologically among the most
intensively studied species, which enables comprehensive stud-
ies on the interplay of wildlife ecology and genomics. Specific-
ally, we expect future genetic studies to investigate further
adaptation and natural selection of bears, inbreeding, and
evaluate circumstances for which effective population size
might be appropriate for tracking abundance of bears. As
new tools that use genetic information for conservation ques-
tions emerge, and sampling, genotyping, and sequencing tech-
niques improve and become cheaper, genetics will become
even more essential for bear research, conservation, and
management.

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