Evolution

Diacodexisearly Eocene of North America,

Europe, Asia
IndohyusEocene of Asia
Shares

ear, tooth characters with fossil whales

Hippopotamidae now thought to be sister group

of Cetaceans (e.g. Cetartiodactyla)

Today artiodactyls far more diverse than

perissodactyls

Evolution

FIGURE 02: Traditional

phylogenetic hypothesis
of the orders Artiodactyla
and Cetacea

Evolution

FIGURE 01: A new

phylogenetic
hypothesis of the
Cetartiodactyla based
on molecular data
from Price et al.
(2005) and OLeary
and Gatesy (2008)

Morphology
Ruminant artiodactylsmultichambered

stomachs (foregut fermentation)

Rumen

(large fermentation chamber)

Reticulum
Omasum
Abomasum (true stomach)
Complex recycling of nutrientsincreased

efficiency
Rate of passage of food is slow

Morphology
Foot structure

paraxonic
Plane of
symmetry
through
digits 3
and 4

FIGURE 03: Right front

feet of four artiodactyls:
A hippopotamus, swine,
elk, and camel

Adapted from Howell, A. B. Speed in

Animals. University of Chicago
Press, 1965.

Morphology
Cannon bone (fused 3rd and 4th metapodials) in

some families
Limbs have springing ligaments
Astragalus with double-trochlea
Restricts

lateral movement

Limbs usually elongate and slim

Postorbital bar or process present
Horns with bone core present

Morphology

FIGURES 04 & 05: Left hind foot

of a pronghorn (Antilocapra
americana), showing the position
of the springing ligament

Camelidae (camels, llamas, etc.)

Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

3 genera / 4 species
northern Africa, central Asia, South America
Appear first in middle Eocene of N. America

plantigrade or digitigrade
3-chambered ruminant
Distinctive divergence of metapodials
60 to 650 kilograms
Long limbs and neck

Family Camelidae
FIGURE 07A: A one-humped
camel (Camelus dromedarius)
roaming the deserts of Qatar

Paul Cowan/ShutterStock, Inc.

FIGURE 07B: The vicugna

(Vicugna vicugna), a camelid that
inhabits the central Andes of
South America
Eduardo Rivero/ShutterStock, Inc.

Family Camelidae
Lower

incisors
retained
Broad
diastema
FIGURE 08: Skull
of an extinct
Pleistocene New
World giant llama
(Camelops); length
of skull 565
millimeters
Modified from Romer, A.S. Vertebrate Paleontology. University of Chicago
Press, 1966.

Suidae (hogs and pigs)

Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

8 genera / 16 species (5/18)

originally Eurasia and Africa, Pacific Islands to
Sulawesi & Phillipines
Appear in late Eocene of Asia
Thick bodied, relatively short limbs, up to 275 kg
4 digits with hooves

Long skullorbits posterior

Large, tusk-like canines (in some)

Family Suidae
5 genera and 18 species
Old World distribution
Appear in late Eocene of Asia
Thick bodied, with relatively short limbs
Weigh up to 275 kilograms
Long skullorbits posterior
Large, tusk-like canines (in some)

Family Suidae
Typically omnivorous
Bunodont molars

Family Suidae

EcoPrint/ShutterStock, Inc.

FIGURE 12: A warthog

(Phacochoerus africanus)
FIGURE 10: A male babirusa
(Babyrousa celebensis)
Teo Boon Keng Alvin/ShutterStock, Inc.

Tayassuidae (peccaries)
Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

3 genera / 3 species
southwestern United States to central

Argentina
4

digits with hooves

digits 3&4 fused proximally
scent gland on rump

Suborder Ruminantia

FIGURE 15: Two phylogenetic hypotheses of ruminant artiodactyls based

on the work of OLeary and Gatesy 2007 (left) and Price et al. 2005 (right)

Tragulidae (chevrotains and mouse deer)

Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

3 genera / 4 species
southeastern Asia & Africa
no horns or antler
hindlimb

Antilocapridae (pronghorn antelope)

Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

1 genus / 1 species
western North America
Both sexes with horns
External

sheath of horn shed annually

Fossil species with more complex horns
Cheek teeth high crowned
Orbits far posterior on skull

Family Antilocapridae

Courtesy of Terry Vaughan

FIGURE 17: The pronghorn (Antilocapra americana)

Family
Antilocapridae

Family Antilocapridae
Among fastest cursorial mammals
Speeds

up to 85 kilometers/hour (53 mph)

Extremely good at long-distance running
Speeds of 65 kph for over 10 kilometers
Evolutionary advantages:
enlarged airways
greater lung surface area
greater capillary density
higher concentration of hemoglobin

Giraffidae (giraffes and okapis)

Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

2 genera / 2 species
sub-Saharan Africa
Elongate neck and limbs

Unguligrade, cannon bone, ossicones

4-chambered ruminant
Short horns covered with fur on parietals
Ossified cartilage forms ossicones
Horns never shed, Horns on both sexes

Family Giraffidae

Family Giraffidae
FIGURE 20: A giraffe (Giraffa
camelopardalis) among umbrella
trees (Acacia tortillis) in Tsavo West
National Park, Kenya

Courtesy of Terry Vaughan

Cervidae (deer)
Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

19 genera / 51 species
Cosmopolitan except sub-Saharan Africa,

Australia, & Antarctica

Antlers,

F caribou too
Paraxonic, cannon bone
4-chambered ruminant

Family Cervidae
Some cervids have short antlers & enlarged

canines
Antlers grow from frontal pedicles
Antler growth under hormonal control
Triggered

by changes in day-length

Antlers

begin growth in April or May

Initially

covered with velvetfur-covered skin

Family
Cervidae
FIGURE 21
(clockwise from
left): Reeves
muntjac, North
American elk,
moose, caribou.
Courtesy of Terry Vaughan
Courtesy of Pam Wood

Ferenc Cegledi/Shutterstock, Inc.

Joe Ferrer/ShutterStock, Inc.

Family Cervidae

Family Cervidae
Cervids mostly browsers
7 to 800 kilograms
Some species gregarious, others solitary
Annual migrations in some species

Moschidae (musk deer)

Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

1 genus / 7 species
central and northeastern Asia
Long hindlimbs
lack

antlers
Saber-like canines
Agile

Bovidae (antelopes, cattle, gazelles,

goats, sheep, and relatives)
Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

50 genera / 143 species

Africa, Europe, Asia, & North America
First appear in Oligocene of Asia
derived from traguloid ancestor in Old World?
Reached New World in Pleistocene

Family Bovidae
Primarily grazers
Cheek teeth high crowned
Canines reduced or absent
Preorbital vacuities on skull in some
Lateral digits reduced or absent
Ulna reduced and fused with radius
Across

Bering land bridge

Unguligrade, paraxonic, cannon bone
4-chambered ruminant

Family Bovidae
FIGURE 23A: A male
Rocky Mountain
bighorn sheep

Courtesy of Terry Vaughan

Bambi L. Dingman/Dreamstime.com

FIGURE 23B: A
mountain goat
(Oreamnos
americanus) from
North America

Family Bovidae
Horns in males
Horns

never branched

Horns

never shed

Females

may bear horns (in heavier species)

Keratinized
Horns

sheath covers bony core

used for ritualized sparring and defense

from predators

Family Bovidae
FIGURE 24A: A group
of musk oxen

Photos.com

FIGURE 24B: A pair of

North American bison
(Bison bison) from
Wyoming
Courtesy of Terry Vaughan

Family Bovidae

Courtesy of Terry Vaughan

Courtesy of Terry Vaughan

Courtesy of Terry Vaughan

Courtesy of Terry Vaughan

Hippopotamidae (hippopotamuses)
Subclass Theria
Infraclass

Eutheria

Order Artiodactyla (even-toed ungulates)

2 genera / 2 species
sub-Saharan Africa
digitgrade

Cetacea
Fully adapted to aquatic life
Baleen whales (Mysticeti) include largest living

or fossil mammals
Remarkable swimming and diving ability
Complex social behaviors
Feed

at middle of marine food web

Toothed whales (Odontoceti)

Feed at top of marine food web
Echolocation

Paleontology
Eocene cetaceans from coastal Tethys Sea
Transitional forms from terrestrial to aquatic
Archaeoceti (fossil whales) believed to evolve

from ancestral artiodactyls

Hippos considered to be sister group to
ancestral cetaceans

Paleontology
FIGURE 01: One
phylogeny of
whales based on
morphological and
fossil evidence

Adapted from Zimmer, C. At the Waters Edge.

Simon & Schuster, 1998.

Paleontology
Early mysticetes (baleen whales) appear in

early Oligocene
Lacked

baleen, had teeth

Large eyessuggests they hunted fish
Later forms had both teeth and baleen

Limited bulk filter feeding possible

Middle

Miocene mysticetes lost teeth

Elongate rostrums to support more baleen

Bulk filter feeding

Paleontology
Transition to filter

feeding
corresponds to
opening of circumAntarctic currents
Explosion

in
marine plankton
productivity

FIGURE 02: Skull of a primitive

cetacean (Dorudon), a fossil
archaeocete from the Eocene
Adapted from Romer, A.S. Vertebrate Paleontology.
University of Chicago Press, 1966.

Paleontology
Odontocetes (toothed whales) probably

evolved from a basilosaurid ancestor

Highly telescoped skulls
Homodont dentition
Evolution of echolocation

Paleontology

FIGURE 03: Skull of

Prosqualodon, a fossil porpoise
from the Miocene
Adapted from Romer, A.S. Vertebrate Paleontology.
University of Chicago Press, 1966.

Morphology
Body fusiform
Nearly hairless
Thick layer of subcutaneous blubber
Teats enclosed within slits next to urogenital

opening
Testes remain abdominal
Vertebrae with high neural arches

Morphology

FIGURE 04: The skeleton of the

Tasmanian beaked whale

Morphology
Cervical vertebrae

compressed
Clavicle absent

FIGURE 05: Cervical

vertebrae of a dolphin
(Delphinus delphis,
Delphinidae).

Morphology
Forelimbs (flippers) paddle-shaped

Digits long and with additional phalanges

Hind limbs vestigial (not visible externally)
FIGURE 06: Dorsal view of the right forelimb of the bottlenose dolphin
(Tursiops truncatus, Delphinidae)

Morphology
Flukes horizontal
Skull highly modified
Posterior
Nasals

migration of nares

and parietals telescoped

Tympanic

bones not braced against skull

Surrounded by air sinuses

Morphology

Cetacean Adaptations
Must breathe air
Able

to alternate between periods of eupnea

and apnea
Rapid rates of gas exchange in lungs
Twice the number of red blood cells
29 times as much myoglobin
Heart rate drops during dives, ~1/2
Blood bypasses certain muscles during diving

Blood flow to brain maintained

Tolerate

high levels of lactic acid

Cetacean Adaptations

Data from Kooyman and Andersen (1969), who cite the sources of the
observations. *Data from Tyack et al. (2006).

Cetacean Adaptations
Deep diving adaptations:
Many

ribs lack connection to sternumlungs

collapse at depth
Lungs dorsal to diaphragm
Volume of non-vascular air spaces is large
Trachea short and large in diameter
Bronchioles braced by cartilage rings
Lungs highly elastic
Myoelastic sphincters in bronchioles of some
odontocetes

Cetacean Adaptations
Swimming Adaptations
Sub-dermal

springs in tailstock store and

release elastic energy
Drag reduced on body surfaces

Compliant spongy layer in outer skin dampens

pressure and turbulence

Larger

body mass increases speed

Also more favorable for thermoregulation

Suborder Mysticeti
4 families of baleen whales
Live in all oceans
Populations decimated by whaling

Suborder Mysticeti
Three distinct feeding modes

Grazing near surface right whales

Swim through swarms of plankton
FIGURE 10A: Skull
of the Atlantic right
whale; length of
skull roughly 4
meters. Note the
baleen plates
attached to the
maxilla.

Suborder Mysticeti
Lunge feedingrorquals
Huge

mouth and head

Furrows on throat allow vast expansion of
throat during feeding
Throat pouch contractedwater and food
pass through baleen plates
Mandible braced by frontomandibular stay
apparatus

Suborder Mysticeti

FIGURE 13: Drawing of the

frontomandibular stay apparatus of
Balaenopteridae
Adapted from Lambertsen, et al., J. Mammalogy 76 (1995): 877-899.

Suborder Mysticeti
Rorquals may also use bubble nets
Swim

below school of prey

Circle prey while emitting constant stream of
bubbles
Swim rapidly up through middle of bubble net
Engulf prey at surface

Suborder Mysticeti

Courtesy of Kirsten Graham

FIGURE 09A: A spiral shaped wall of bubbles is produced by

a submerged whale, which traps a school of fish

Suborder Mysticeti
Gray whales exhibit third style of feeding

Plow head through mud at bottom

Scoop or suck up sediments

Filter out marine organisms from bottom

Family Balaenidae (bowhead whales and

right whales)
Subclass Theria
Infraclass

Eutheria

Order Cetacea (dolphins, porpoises, and whales)

Suborder Mysticeti

2 genera / 4 species
temperate and polar oceans of the world
Large (18 m, up to 67,000 kg), thick-bodied
Huge head and tongue
350 long baleen plates per side
Flippers short and rounded

Balaenopteridae (rorquals)
Subclass Theria
Infraclass

Eutheria

Order Cetacea (dolphins, porpoises, and whales)

Suborder Mysticeti

2 genera / 7 species
temperate and polar oceans of the world
Vary widely in size:

Minke whale 11 m / 4,000 Kg; Blue whale 31 m / 160,000 Kg

Baleen plates short

Longitudinal furrows on throat aid feeding
May migrate long distances

Family Balaenopteridae

FIGURE 15: A common minke whale (Balaenoptera acutorostrata)

Kelvin Aitken/Peter Arnold, Inc.

Family Balaenopteridae

FIGURE 16: A mother and young humpback whale (Megaptera

Melissaf84/Dreamstime.com
novaeangliae; Balaenopteridae)

Family Eschrichtiidae (gray whale)

Subclass Theria
Infraclass

Eutheria

Order Cetacea (dolphins, porpoises, and whales)

Suborder Mysticeti

1 genus / 1 species
North Pacific (east & west coasts)
Up to 15 meters and 31,500 kg
Throat with 2-3 short grooves
Small head, short baleen plates
Migrate up to 22,000 kilometers

Family Neobalaenidae (pygmy right whale)

Subclass Theria
Infraclass

Eutheria

Order Cetacea (dolphins, porpoises, and whales)

Suborder Mysticeti

1 genus / 1 species
Cold waters of the southern hemisphere
Baleen
Only

to 6 m
Gray coloration, small falcate dorsal fin

Suborder Odontoceti
Toothed whales, porpoises, and dolphins
Seven families
All oceans and seas, and some river systems
Homodont teeth and piston-like tongue used

to capture prey
Some may stun prey acoustically

Suborder Odontoceti
Phylogeny of odontocetes
Sperm
River

whales considered basal lineage

dolphins not a monophyletic group

Amazon, LaPlata, and Yangtze River dolphins in one

group
Ganges and Indus River dolphins in another group
River

dolphins isolated in shallow seas as sea

levels dropped in Miocene

Family Delphinidae (dolphins, killer &

pilot whales)
Subclass Theria
Infraclass

Eutheria

Order Cetacea (dolphins, porpoises, and whales)

Suborder Odontoceti

17 genera / 34 species
all oceans, & some rivers (Ganges, Amazon)
Appear in Oligocene of Europe
Homodont dentition, 0/2 to 65/58
1.5-10 m long, 50 to 7,000 kg
Fatty-deposit melon above & behind snout

Family Delphinidae
Rapid swimmers, regular leaps
Highly gregarious typically
Form

schools (=pods)

Highly

vocal

Cooperative

behaviors in some species

Remarkable

intelligence

Family Delphinidae
FIGURE 18A: A killer whale

Courtesy of S. Mizroch, NOAA, NMFS,

National Marine Mammal Laboratory

FIGURE 18B: A Pacific

white-sided dolphin

Ferdericb/Dreamstime.com

Family Iniidae (river dolphins)

Subclass Theria
Infraclass

Eutheria

Order Cetacea (dolphins, porpoises, and whales)

Suborder Odontoceti

3 genera / 3 species
Freshwater rivers (Yangtze, Amazon, Doce

rivers)
Chinese river dolphin now extinct
Homodont

dentition

Family Monodontidae (beluga & narwhal)

Subclass Theria
Infraclass

Eutheria

Order Cetacea (dolphins, porpoises, and whales)

Suborder Odontoceti

2 genera / 2 species
High latitude northern hemisphere oceans &

major rivers
Almost

homodont dentition, 5/2 to 11/11

Narwhal I on left spiral up 2.7 m

Family Monodontidae
Both species:
56

m long &
2,000 kg
Gregarious

FIGURE 19: A male

(foreground) and female
(background) narwhal
(Monodon monoceros)
in the Arctic Ocean

David Fleetham/Visuals Unlimited

Family Phocoenidae (porpoises)

Subclass Theria
Infraclass Eutheria
Order Cetacea (dolphins, porpoises, and whales)
Suborder Odontoceti

4 genera / 6 species (3/6)

Coastal waters of all oceans & seas of northern

hemisphere, coast of South america & southeastern

Asia

1.5 2.1 m, short jaws

Slow, inconspicuous, small groups
15/15 to 30/30

Family Physeteridae (sperm whales)

Subclass Theria
Infraclass Eutheria
Order Cetacea (dolphins, porpoises, and whales)
Suborder Odontoceti

2 genera / 3 species
Oceans of the world except Arctic Ocean

Largest toothed whale, 18 m, >53,000 kg

Physeter - head 35% body, spermaceti organ, oil

Family Physeteridae
Physeter:

Socialgroups of up to 1,000 individuals

Schools usually contain one large male

Younger

males form bachelor herds

Use

powerful echolocation signals at great

depth

Feed

on large squid and fish

Family Platanistidae (Indian river dolphins)

Subclass Theria
Infraclass Eutheria
Order Cetacea (dolphins, porpoises, and whales)
Suborder Odontoceti

2 genera / 2 species (1/2)

Major rivers of India: Ganges, Indus,
1 - 3 m
Long slender beaks, fish & invertebrate eaters
25/24 to 61/61
Eyes educed
Side swimmers
echolocation

Family Ziphiidae (beaked whales)

Subclass Theria

Infraclass Eutheria

Order Cetacea (dolphins, porpoises, and whales)

Suborder Odontoceti

6 genera / 19 species
Coastal waters of all oceans & seas of northern hemisphere,

coast of South america & southeastern Asia

Up to 13 m
Inconspicuous
0/1-2 to 19/27
Flukes unnotched, small falcate dorsal fin

Family Ziphiidae
Stomach divided into 414 chambers
Deep diversstay submerged for long times
Most species social
Open ocean habitats make these cetaceans

difficult to study

Perissodactyla
Monophyletic group
3 families, 6 genera, and 16 species
Formerly much more diverse
Axis of symmetry of footmiddle digit
Mesaxonic

condition

Elongate rostrums with molariform teeth

Simple stomach but large ceacum

Perissodactyla

FIGURE 01: One

phylogenetic hypothesis of
the Perissodactyla and their
sister groups

Perissodactyla

Adapted from Howell, A. B. Speed in Animals.

University of Chicago Press, 1965.

Evolution
Arose in late Paleocene in Asia
Diversified and spread to Europe and North

America by early Eocene

Dominant browsers of the Eocene
Began to decline in Miocene
Climatic

changes
Competition with Artiodactyls

Evolution
Tapirs evolved in North America in Eocene
Rhinoceroses evolved in Eurasia and North

America in early Eocene

Paraceratherium

(Oligocene)largest known
land mammal (5 meters tall, 11,000 kilograms

Horses evolved in North America in early

Eocene
Evolution

of horses more complex than

previously thought

Horse Evolution
FIGURE 03: Skull of Hyracotherium
(Eohippus), one of the first known equids

FIGURE 04: Right upper molars of

four fossil equids and the extant
Equus. These teeth illustrate stages
in the evolution of the equid molars

Both illustrations modified from Romer, A.S. Vertebrate Paleontology.

University of Chicago Press, 1966.

Horse Evolution

Adapted from MacFadden, B. J., Science. 307 (2005): 1728-1730.

FIGURE 05: A time chart of the North

American equids over the past 55 my

Cursorial Adaptations
Integrated locomotor and respiratory

functions
Synchronize

breathing with stride cycles

Tracheal valving shunts air to lungs

Slim, elongate legs
Running speed determined by
Stride

length
Stride rate

Cursorial Adaptations
Stride length
Longer

legs
Metacarpals and metatarsals elongate
Loss or reduction of clavicle
Flexion and extension of spine
Stride rate
Increase

the number of moveable joints in limb

Only hoof-bearing tips of digits contact ground

Muscles

positioned near bodytendons distally

Cursorial Adaptations

Cursorial Adaptations
FIGURE 07: A running pronghorn, showing the flexion and extension of
the vertebral column and the changing position of the scapula

Cursorial Adaptations
Ankle joint
Astragulus

rests on distal tarsal bones

Tarsals

modified by loss or fusion of elements

Weight

transferred to central digits

Nuchal ligament supports weight of head

Springing ligament in feet

Cursorial Adaptations

FIGURE 08: The tarsus of

the horse (Equus caballus)

FIGURE 09: The nuchal

ligament of the horse (Equus
caballus) illustrating the
extent of the insertion on the
skull and vertebral spines

Dietary Adaptations
Herbivorous diet
Molariform cheek teeth
Large complex occlusal surfaces
Plant material difficult to digest
Plant

defensive compounds
Cellulose in plant cell walls
Complex chambers for housing microbes
Microbes needed to break down cellulose

Dietary Adaptations
Microbial fermentation
Slow

process
Requires fermentation chamber
Perissodactyla use hindgut fermentation
Cecum

(or caecum)expanded colon

Ruminant Artiodactyls use foregut

fermentation
Multichambered

stomach

Dietary Adaptations

FIGURE 10: A comparison of the anatomy of the gastroinstestinal

tract of a cow and a white rhino

Family Equidae (asses, horses, and zebras)

Subclass Theria
Infraclass Eutheria
Order Perissodactyla (horses, rhinoceroses, and
tapirs)
1 genus / 9 species (1/8)
Wild: East Africa, Near East to Mongolia, domestic: nearly

cosmopolitan
mesaxonic
Unguligrade, radius & ulna fused

3/3, 1/1, 3-4/3, 3/3 = 40-42, hypsodont

Only digit three bears weight

Family Equidae
Equus evolved in North America
Crossed into Asia in late Pliocene
Horse domestication may date as far back as

4,500 years ago in Central Asia

Horses returned to North America with

Spanish conquistadors in 1500s

Family Equidae

ARCO/I. Schulz/age fotostock

Courtesy of Terry Vaughan

FIGURE 12A: A mountain

zebra (Equus zebra) in
Mountain Zebra National
Park, South Africa

FIGURE 12B: A Przewalskis

horse (Equus caballus przewalskii)
stallion from Mongolia

Family Equidae
Most species are highly social
Form

herds and/or clans

Polygynous mating system
Social hierarchyled by dominant stallion
Harems formed in some species
Bachelor herds

Complex

behavior and vocal communication

Fission-fusion social system common

Family Equidae

ARCO/I. Schulz/age fotostock

Courtesy of Terry Vaughan

FIGURE 12A: A mountain

zebra (Equus zebra) in
Mountain Zebra National
Park, South Africa

FIGURE 12B: A Przewalskis

horse (Equus caballus przewalskii)
stallion from Mongolia

Family Equidae

Family Rhinocerotidae (rhinoceroses)

Subclass Theria
Infraclass Eutheria

Order Perissodactyla (horses, rhinoceroses, and

tapirs)
4 genera / 5 species
Sub-Saharan Africa, Southeast Asia
Mesaxonic, 3 digits hind, 3-4 digits fore
1-2/0-1,

0/1-1, 3-4/3-4, 3/3, hypsodont

Graviportal limbs
Weigh up to 2,800 kilograms
Nasal bones thickened and enlarged
Support

keratinous horn

Family Rhinocerotidae
Solitary to social
Territorialuse scent marking (dung piles)
Sumatran rhino (Dicerorhinus) smallest
Coarse hairs
Solitarytropical

forests

Vocal
Less

than 300 living today

Use of horns in traditional medicine

All 5 species listed as endangered

Family Rhinocerotidae
FIGURE 16A: The white rhinoceros
(Ceratotherium simum) of South
Africa
Carlos Arguelles/ShutterStock, Inc.

Courtesy of Terry Vaughan

FIGURE 16B: An Indian rhino

(Rhinoceros unicornis). Note the single
horn and the thick, plate-like skin in this
species

Family Rhinocerotidae

FIGURE 17: Skull

of a black rhinoceros
(Dinoceros bicornis)

Family Tapiridae (tapirs)

Subclass Theria
Infraclass

Eutheria

Order Perissodactyla (horses, rhinoceroses, and

tapirs)
1 genus / 4 species
Central to South America, Malaya
Mesaxonic, 3 digits hind, 4 digits fore
Short fleshy proboscis
3/3,

1/1, 4/3-4, and 3/3 = 42-44, low-crowned

Family Tapiridae
FIGURE 13: A Brazilian tapir
(Tapirus terrestris) from South
America. The upper lip is
elongate, forming a downcurved proboscis.
Ferenc Cegledi/ShutterStock, Inc.

Jan Gottwald/Dreamstime.com

FIGURE 14: A Malayan tapir

(Tapirus indicus) showing the
flexibility of the proboscis

Family Tapiridae
Nasal bones of skull retractedabove orbits

FIGURE 15: Skull of

a Bairds tapir
(Tapirus bairdii)

Introduction
Creodonts appear in the Paleocene
Carnivorans evolved in response to increasing

radiation of herbivores (prey)

Carnivora phylogeny controversial
Two major clades
Feliformia
Caniformia

Carnivora

Adapted from Flynn, J. J., et al., Syst. Biol. 54 (2005): 317-337.

Carnivora
Morphological characters
Expanded

braincase
Flange on lateral basioccipital
Loss of third molar
Fused scaphoid
and lunar bones
in carpals

FIGURE 02: Anterior view of

the left carpus of the gray fox
(Urocyon cinereoargenteus).

Carnivora
Major Patterns
Two cladesFeliformia and Caniformia

Nandiniidae (African palm civet) basal to Feliformia

Eupleridae (Malagasy carnivorans) radiated in

isolation on Madagascar

Seals, sea lions, and walrus for a clade (Pinnipedia)

Red panda (Ailuridae) is basal member of the

superfamily Musteloidea

Mephitidae (skunks) form a distinct clade within

Musteloidea

Carnivora
Mostly predaceous
Canines

are usually large

Acute sense of smell

Cursorial ability strongly developed in some

groups (e.g. Canidae and Felidae)

In many, 4th upper premolar and 1st lower
molar are carnassials
Temporomandibular joint is tight

Feliformia
Nandiniidae (African palm civet)
Felidae (cats)
Viverridae (civets and genets)
Hyaenidae (hyenas and aardwolf)
Herpestidae (mongooses)
Eupleridae (Malagasy civets and mongooses)

Nandiniidae (African palm civet)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

1 genus / 1 species
sub-saharan Africa
Nandinia Omnivore/herbivore
Good

climber, headfirst down

Family Nandiniidae
Single species (Nandinia binotata), African

palm civet
Genetically distinct from other feliforms
Sub-Saharan Africa south to Zimbabwe
2 to 5 kilograms
Short, woolly fur
Omnivorous

Felidae (cats)
Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

18 genera / 36 species (14/40)

Cosmopolitan except on Antarctica, Australis,

& Madagascar
3/3 1/1 3/2 1/1 = 30
Most highly specialized for predation

Family Felidae
Digitigrade foot posture
Claws recurved and retractile (except in

cheetah)
Spotted or striped pelage common
Weights range from 1 to 275 kilograms

Family Felidae
Excellent senses of smell, sight, and hearing
Some species nocturnal
Tapetum lucidum present in eye
Some species are agile climbers
Cheetahs sprint to 90 kph (60 mph) over short

distances
Many species solitary
Lions

are sociallive in large prides

Family Felidae
Cat-like mammals appear in North America in Eocene

Adapted from Martin, L.D., Transactions of the Nebraska

Academy of Sciences, 7 (1980):141154. 1980.

Family Felidae
Modern cats began radiation in Miocene
Panthera lineage arose in Eurasia 11 million

years ago
Eight felid lineages evolved
Spread across land bridges as sea levels
changed
Roughly 6 million years ago lineage of small
cats evolved (ancestors of domestic cats)

FIGURE 03A: Caracal (Caracal caracal)

FIGURE 03B: A margay (Leopardus wiedii)

Purestock/age fotostock
Courtesy of Terry Vaughan

FIGURE 03D: A mountain lion (Puma concolor)

FIGURE 03C: A leopard (Panthera pardus)

Philip Lane/Dreamstime.com
Courtesy of Terry Vaughan

Family Felidae

Viverridae (civets, genets, linsangs, etc.)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

20 genera / 34 species (15/35)

Africa, southern Europe, Asia, Indonesia &

Phillipine
3/3 1/1 3-4/3-4 2/2 = 36-40
Diverse niches

Family Viverridae
Small to medium size (600 grams to 20

kilograms)
Long rostrum
Short legs (relatively)
Tail long and bushy
Well-developed perineal (anal) glands

Family Viverridae
Some are agile climbers, some semiaquatic,

others terrestrial
Mostly carnivorous and nocturnal

FIGURE 08B: Largespotted genets (Genetta

tigrina), agile climbers
that often feed on birds

Courtesy of K. Wells

FIGURE 08A: A Maylayan civet

(Viverra tangalunga)

Courtesy of T. R. Huels

Family Viverridae

FIGURE 10:
Occlusal view of
the cheek teeth of
a large-spotted
genet

Hyaenidae (aardwolf and hyenas)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

4 genera / 4 species (3/4)

Africa, southwestern Asia, & India
3/3 1/1 4/3 0-1/1 =32-34 (except Proteles)

Family Hyaenidae
Both scavengers and predators
Spotted hyenas form large packs or clans
Highly

social

Mostly

nocturnal

Striped hyenas more omnivorous

Aardwolf has reduced dentition
Feeds

on termites

Family Hyaenidae

Courtesy of Terry Vaughan

FIGURE 13: Spotted hyena (Crocuta crocuta) in

Etosha National Park, Namibia

Family Hyaenidae

FIGURE 14: Skulls of a spotted hyena and an aardwolf

Herpestidae (mongooses)
Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

20 genera / 34 species (14/33)

Africa, Asia, southern Europe & Madagascar,

introduced widely
3/3 1/1 3-4/3-4 2/2 = 36-40
Highly social, scent marking

Family Herpestidae
Small, long-bodied carnivores
270 grams to 5 kilograms
Large auditory bullae
Anal scent glands well-developed
Some species have complex social lives

Family Herpestidae

EcoPrint/ShutterStock, Inc.

Courtesy of Terry Vaughan

FIGURE 11: Left: Yellow mongoose (Cynictis

penicillata) in the Kalahari Desert.
Above: A group of meerkats (Suricata suricatta) on
lookout duty.

Family Herpestidae

Eupleridae (Malagasy carnivores)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

7 genera / 8 species
Madagascar
Nocturnal / terrestrial (most)

Family Eupleridae

Courtesy of M. Korinek

FIGURE 12: Above: The fossa, Cryptoprocta ferox, is

the largest living carnivore in Madagascar. Top right:
The Galidia elegans, the Malagasy ring-tailed
mongoose, is diurnal and lives in small family
groups. Right: Galidictis fasciata, the Malagasy
broad-striped mongoose, is nocturnal and solitary.

Courtesy of James Ryan

Courtesy of James Ryan

Eupleridae

Caniformia
Family Canidae (wolves, foxes, and allies)
Family Ursidae (bears)
Family Odobenidae (walrus)
Family Otariidae (eared seals and sea lions)
Family Phocidae (seals)
Family Ailuridae (red panda)
Family Mephitidae (skunks and allies)
Family Procyonidae (raccoons and allies)
Family Mustelidae (weasels, badgers, & allies)

Canidae (coyotes, dogs, foxes, jackals, etc.)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

14 genera / 34 species (13/35)

Cosmopolitan except Antarctica
3/3 1/1 4/4 2/2 = 42

Family Canidae
Fossils date to Eocene of North America
Broadly adapted carnivores
Large nasal chamberexcellent sense of smell
Limbs usually long, Highly cursorial, Feet digitigrade
Blunt, non-retractile claws
Clavicle absent
Robust canines, carnassials present
Molars retain crushing surfaces
More flexible diet

Family Canidae
Robust canines, carnassials present
Molars retain crushing surfaces
More flexible diet
Limbs generally long
Clavicle absent
Feet digitigrade
Blunt, non-retractile claws

Family Canidae
Larger species hunt with lengthy pursuit
Endurance
Some

more important than speed

species hunt in packs

Smaller species opportunistic hunters/

scavengers
Smaller species more solitary

Family Canidae

Johan Reineke/Dreamstime.com

Courtesy of Terry Vaughan

FIGURE 15:
Clockwise, from
top left: African
wild dog (Lycaon
pictus); Gray
wolf (Canis
lupus); bat-eared
fox (Otocyon
megalotis); and
black-backed
jackel (Canis
mesomelas).

Courtesy of Terry Vaughan

Courtesy of Terry Vaughan

Family Canidae

Ursidae (bears)
Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

5 genera / 8 species
Primarily northern hemisphere, but on all

continents except Antarctica & Australia

Family Ursidae
Large body size
More omnivorous diet (except polar bears)
Long rostrum
Post-carnassial teeth robust and adapted to

crushing
Limbs powerful, feet plantigrade
Claws non-retractile
Tails very short

Family Ursidae
Omnivorous feeding habits
Polar

bear diet restricted to seals

Giant

panda diet mostly bamboo

Northern species may hibernate

Heart
Live

rate and metabolism slows

off accumulated fat reserves

Arouse

periodically

Family Ursidae

Karel Bro/ShutterStock, Inc.

FIGURE 18: Above: A Kodiak brown bear (Ursus arctos), also

known as the grizzly bear, in Alaska. Right: A giant panda
(Ailuropoda melanoleuca) from China.

Mike Flippo/ShutterStock, Inc.

Family Ursidae

Pinnipedia
Monophyletic group
Family

Odobenidae (walrus)

Family

Otariidae (eared seals)

Family

Phocidae (earless seals)

Probably evolved from a bear-like ancestor in

late Eocene
Highly adapted to marine lifestyle

Pinnipedia
Large (45 to 3,600 kilograms)
Favorable

ratio of surface area to volume

Body insulated with blubber (and fur in some)

Pinnae small or absent
External genitalia and mammary nipples withdrawn

beneath body surface

Tail rudimentary
Forelimbs flipper-like

Pinnipedia
Only distal limbs protrude from body
Fusiform body shape
Skull partially telescoped
Orbits large
Manus and pes fully webbed
Vertebral column flexible
Diving and swimming ability highly advanced

Pinnipedia

Adapted from King, J. E. Seals of the World. Cornell University Press, 1983.

Phocidae (seals)
Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

10 genera / 19 species (13/19)

usually along coasts above 30N and below

50S latitudes, exceptions Galapagos Islands

No

external ear

Hind

flippers useless on land

Foreflippers

short and well furred

Family Phocidae
Pelage spotted, banded, or mottled
Heavy layers of blubber
Cheek teeth more homodont
Fusiform body shape reduces drag
80 to 3,600 kilograms

Family Phocidae

Galina Barskaya/Dreamstime.com

FIGURE 24A: Harbor seal (Phoca vitulina; Phocidae) on the California coast

Family Phocidae
Many species are monogamous
Some gregarious and polygynous
Elephant seals
Form dominance hierarchies at rookeries
Feed on fish, cephalopods, and mollusks
Leopard seal eats penguins
Crab-eater

seal is a filter-feeder

Otariidae (fur seals and sea lions)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

7 genera / 14 species (7/16)

Pacific Ocean (N & S) and coasts, Atlantic

coast of South America, SW Australian coast

of Indian Ocean
Eared seals and sea lions
Better able to move on land than phocid seals

Family Otariidae
Small ear pinna
Nails on three middle digits
Males larger than females
Body covered with fur
60 to 1,000 kilograms

Family Otariidae

Pichugin Dmitry/ShutterStock, Inc.

FIGURE 24B: A New Zealand sealion (Phocaractos hookeri; Otariidae)

Family Otariidae

Gregarious and highly vocal on land

Courtesy of Terry Vaughan

Seasonal breeding rookeries

Family Otariidae

Family Odobenidae (walruses)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

1 genus / 1 species
northern oceans and coasts, usually above

58 N latitude
Largeup to 1,270 kilograms
Nearly hairless, stout vibrissae
Lack ear pinna
Hind flippers can be brought under body

Family Odobenidae
Both sexes have tusk-like upper canines
Tusks

lack enamel

Feed on mollusks taken from sea floor

Gregarious and polygynous
Groups
Vocal

may reach 1,000 individuals

in and out of water

Family Odobenidae

Morten Hilmer/ShutterStock, Inc.

Superfamily Musteloidea
Family Ailuridae
(red

panda)

Family Mephitidae
(skunks and stink badger)
Family Procyonidae
(raccoon, ringtail, and allies)
Family Mustelidae
(weasels, badgers, otters, and allies)

Ailuridae (red panda)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

1 genus / 1 species
southeastern Asia
Not

related to giant pandas

Enlarged radial sesmoid for arboreal grasping
Arboreal and largely crepuscular, Mostly solitary
Eat bamboo leaves, berries, flowers etc.
Endangered throughout their range

Family Ailuridae

Greg Payan/Dreamstime.com

FIGURE 27A: A red panda (Ailurus fulgens)

Mephitidae (skunks and stink badgers)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

4 genera / 13 species
North America to central South America &

Indonesia and Phillipines

Skunks

are moderately small (0.5 to 4 kilograms)

Conspicuously colored (black and white)
Adapted for digging (Muscular legs, long claws)
Nocturnal, solitary, omnivorous
Anal glands spray noxious odor

Family Mephitidae

Courtesy of Terry Vaughan

FIGURE 27B: Striped skunk (Mephitis mephitis) in Arizona

Family Procyonidae (coatis, raccoons, etc.)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

6 genera / 18 species (6/14)

North and South America (Canada to

Argentina)
Omnivorous

and agile climbers

Plantigrade, non-retractile/semi-retractile claws
Dexterous forefeet in some species
Tail longprehensile in kinkajous

Family Procyonidae
FIGURE 28A: A raccoon (Procyon lotor)
from North America

L.L. Masseth/ShutterStock, Inc.

FIGURE 28B: A South American

coati (Nasua nasua)
Musat Christian/Dreamstime.com

Family Procyonidae

Family Mustelidae (badgers, otters, weasels)

Subclass Theria
Infraclass

Eutheria

Order Carnivora (carnivores)

25 genera / 65 species
Cosmopolitan except Antarctica, Australia,

Madagascar, & oceanic islands

Short

limbs and elongate bodies (in some)

Skull with long braincase and short rostrum
Carnassials blade-like in many species

Family Mustelidae
Anal scent glands well developed
Tail long
Smallest species weigh 35250 grams
Largest species 3245 kilograms
Carnivorousaggressive hunters
Sea otters feed on invertebrates and
vertebrates
Mostly terrestrial (except otters and mink)

Family Mustelidae
Badgers adapted for digging
Induced ovulation common
Typically

require prolonged copulation

Delayed implantation of embryo common

May

be delayed up to 10 months (badgers)

Family Mustelidae

Photodisc

Serg Zastavkin/ShutterStock, Inc.

FIGURE 30: Left, a wolverine (Gulo gulo) and, right, a North American river
otter (Lontra canadensis).

Family
Mustelidae

Family Mustelidae

FIGURE 32: Occlusal views of the cheek teeth of two carnivores

Family Manidae (pangolins)

Subclass Theria
Infraclass

Eutheria

Order Pholidota (pangolins)

1 genus / 7 species
S Africa & SE Asia