Diversity
Cetartiodactyla is a group comprised of two orders of mammals that are superficially
quite different and that, until recently, were recognized as two separate monophyletic
clades. These orders are
Artiodactyla
, even-toed ungulates, including animals such as cows (
Bovidae
), camels (
Camelidae
), and deer (
Cervidae
), and
Cetacea
, a group of mammals that are highly specialized for an aquatic lifestyle, including
baleen whales
and
toothed whales
. Recent molecular evidence suggests that
Cetacea
evolved from artiodactyl ancestors, making
Artiodactyla
non-monophyletic unless
Cetacea
is included. Experts suggest the monophyletic clade representing artiodactyls and
cetaceans be called Cetartiodactyla.
Geographic Range
Cetartiodactyls are found the world over, from north of the Arctic Circle to the waters
surrounding Antarctica.
Artiodactyls
are native to all continents except Antarctica and Australia, and some
artiodactyls
are domesticated and have been introduced around the world by humans.
Cetaceans
inhabit all of the world's oceans and some freshwater lakes and rivers in South America,
North America, and Asia.
- Biogeographic Regions
- nearctic
- palearctic
- oriental
- ethiopian
- neotropical
- australian
- arctic ocean
- indian ocean
- atlantic ocean
- pacific ocean
- mediterranean sea
- Other Geographic Terms
- holarctic
- cosmopolitan
Habitat
Most
artiodactyls
live entirely on land and reside in a range of terrestrial habitats, such as savannah,
forest, mountains, desert, and farmland. One
artiodactyl
family,
Hippopotamidae
, is semi-aquatic and can be found in freshwater lakes, ponds, streams, and rivers.
Cetaceans
, on the other hand, are exclusively aquatic and inhabit the world's oceans, as well
as some freshwater rivers and streams.
- Habitat Regions
- temperate
- tropical
- polar
- terrestrial
- saltwater or marine
- freshwater
- Terrestrial Biomes
- tundra
- taiga
- desert or dune
- savanna or grassland
- chaparral
- forest
- rainforest
- scrub forest
- mountains
- Other Habitat Features
- agricultural
- riparian
- estuarine
Physical Description
Because
cetaceans
are so highly specialized for their aquatic lifestyle, they bear little resemblance
to their
artiodactyl
ancestors. They have nearly hairless, fusiform bodies. They lack hind limbs except
for tiny internal pelvic vestiges, and the forelimbs are modified into streamlined
flippers. The tail bears a flattened fluke. In addition, cetacean skulls are highly
modified so that the nares are located on the top of the head. On the other hand,
most
artiodactyls
are specialized for cursorial locomotion, with long, hoofed limbs, and they lack
the extreme aquatic specializations found in
cetaceans
. Most cetartiodactyls are relatively large animals, but there is an enormous range
of body sizes in this group. Blue whales,
Balaenoptera musculus
, are the largest animals on earth, growing over 27 meters in length and weighing
over 190,000 kg, whereas the smallest
artiodactyl
, the lesser mouse deer (
Tragulus javanicus
), is just 45 cm long and weighs 2 kg. Many species of cetartiodactyls exhibit sexual
dimorphism, with males larger than females or vice versa. Also, male
artiodactyls
often bear antlers or large horns, and some male cetartiodactyls (narwhals (
Monodon monoceros
),
tragulids
, and
suids
) bear large tusks.
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes alike
- female larger
- male larger
- ornamentation
Reproduction
Cetartiodactyls are monogamous, polyandrous, polygynandrous, or polygynous. Polygyny,
in which social groupings consist of adult females and their young and one or a few
adult males, is a common cetartiodactyl strategy. It occurs in species as different
as elk (
Cervus elaphus
) and killer whales (
Orcinus orca
). Sexual dimorphism in ornamentation (such as antlers) and body size indicates intense
male-male competition for mates in many species.
- Mating System
- monogamous
- polyandrous
- polygynous
- polygynandrous (promiscuous)
In general, cetartiodactyls are not highly prolific, giving birth to just one or two
young every one or two years. However, some members of the family
Suidae
may have 12 or more young at a time. Breeding may be either seasonal or year-round.
At least one species, the minke whale (
Balaenoptera acutorostrata
) regularly experiences a postpartum estrus. Gestation periods are as short as four
months in small
artiodactyls
to as long as 17 months in Baird's beaked whales (
Berardius bairdii
), and youngsters are weaned between 2 1/2 and 24 months of age. Age at sexual maturity
varies widely as well, from 5 months to more than 10 years.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- year-round breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- viviparous
- post-partum estrous
Most cetartiodactyls have precocial young, as it is necessary for them to be able
to walk or swim from the moment of birth. Young stay with their mothers for at least
four months. In some species, such as bottlenosed dolphins (
Tursiops truncatus
), young remain with their mothers for up to five years, long after they are weaned.
Males may care for their offspring indirectly by defending family groups, but they
generally do not help females raise their young.
- Parental Investment
- precocial
-
pre-fertilization
- provisioning
-
protecting
- female
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
-
pre-independence
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
- post-independence association with parents
- extended period of juvenile learning
- maternal position in the dominance hierarchy affects status of young
Lifespan/Longevity
Cetartiodactyls are relatively long-lived mammals. Most species live for at least
a decade in the wild, and captivity usually prolongs life expectancy by several years.
Cetaceans
are especially long-lived; 116-year-old fin whales (
Balaenoptera physalus
) have been reported from the wild and bowhead whales (
Balaena mysticetus
) may live up to 200 years.
Behavior
Most cetartiodactyls are highly social animals. Some live in large herds or pods
numbering hundreds or even thousands of individuals. Group living commonly results
in the formation of dominance hierarchies among both male and female cetartiodactyls.
A few species, such as greater mouse-deer (
Tragulus napu
) and river dolphins (
Platanista
) are solitary. Many cetartiodactyl species migrate seasonally or are nomadic, and
some bovid species (
Bovidae
) are territorial. Different species vary in their timing of daily activities: some
are diurnal, some are nocturnal, some are crepuscular, and some are active at any
time of the day or night.
- Key Behaviors
- cursorial
- terricolous
- natatorial
- diurnal
- nocturnal
- crepuscular
- motile
- nomadic
- migratory
- sedentary
- solitary
- territorial
- social
- dominance hierarchies
Communication and Perception
Cetartiodactyls perceive the world through visual, tactile, auditory, and chemical
means. Some
cetaceans
, the
Odontoceti
, navigate and hunt using echolocation. Terrestrial cetartiodactyls often communicate
with scent; many species, such as those in the family
Cervidae
, have specialized glands for doing so. Communication in
cetaceans
is accomplished largely by sound, as sound waves travel well in water. Low frequency
sounds produced by baleen whales (
Mysticeti
) may travel for hundreds of kilometers, allowing individuals to communicate with
one another over great distances.
- Other Communication Modes
- scent marks
- Perception Channels
- visual
- tactile
- acoustic
- echolocation
- chemical
Food Habits
As a group, cetartiodactyls consume a wide array of terrestrial and aquatic food items.
Most
artiodactyls
are herbivores, consuming grass, leaves, bark, and other plant parts. Those in the
family
Suidae
are omnivorous, and eat small
mammals
,
reptiles
,
amphibians
,
insects
and other invertebrates, fruit, bulbs, rhizomes, fungi, carrion, and
bird
eggs.
Cetaceans
consume plankton, fish,
squid
,
crustaceans
, and aquatic
birds
and
mammals
(including other
cetaceans
).
- Primary Diet
-
carnivore
- eats terrestrial vertebrates
- piscivore
- eats eggs
- insectivore
- eats non-insect arthropods
- molluscivore
- scavenger
- herbivore
- omnivore
- planktivore
- mycophage
- Foraging Behavior
- filter-feeding
Predation
Artiodactyls
are an important food source for many large mammalian carnivores, notably
felids
,
canids
, and
ursids
.
Cetaceans
, on the other hand, have few natural predators, save other
cetaceans
(killer whales,
Orcinus orca
),
sharks
, and occasionally walruses (
Odobenus rosmarus
). Group living ("safety in numbers") and camouflage are two defenses often employed
by cetartiodactyls against predation.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Cetartiodactyls are primary, secondary, and higher-level consumers, filling roles
of predator (most
cetaceans
) and prey (most
artiodactyls
). Terrestrial cetartiodactyls are plagued by ectoparasites such as
fleas
,
lice
, and
bot flies
.
Cetaceans
, though aquatic, are not free from external parasites either, and are host to
barnacles
,
copepods
, and
whale lice
. Both terrestrial and aquatic species host internal parasites as well, such as
tapeworms
,
flukes
, and
nematodes
. Interestingly, birds have evolved commensal relationships with both aquatic and
terrestrial cetartiodactyls.
Seagulls
follow schools of
dolphins
and consume small fish stirred up by the
cetaceans
, and
cowbirds
follow herds of
cattle
and consume insects stirred up by the hooves of the
artiodactyls
. Also, some cetartiodactyl species are mutualists with animals that feed on their
ectoparasites: topsmelt (
Atherinops affinis
) consume whale lice that live on the skin of gray whales (
Eschrichtius robustus
), while oxpeckers (
Buphagus
) remove fleas and other parasites from the skin of various African
artiodactyls
.
- topsmelt Atherinops affinis
- oxpeckers Buphagus
- fleas Siphonaptera
- lice Anoplura
- bot flies Oestridae
- barnacles Cirripedia
- copepods Copepoda
- whale lice Cyamidae
- tapeworms Cestoda
- flukes Trematoda
- nematodes Nematoda
- birds Aves
Economic Importance for Humans: Positive
Cetartiodactyls are of immense economic importance to humans. They have been hunted
for thousands of years, for food, for sport, and for various body parts. They are
important for ecotourism, be it a whale-watching boat off the coast of Maine or a
safari on the African savannah. Several
artiodactyl
species have been domesticated for thousands of years and are used to produce meat,
milk, leather and wool, and their dung is used as a fertilizer.
Cetaceans
are sometimes kept in captivity and taught to perform tricks. Many cetartiodactyls
are the focus of research programs that help us to better understand many aspects
of evolution, physiology, and behavior.
- Positive Impacts
- food
- body parts are source of valuable material
- ecotourism
- research and education
- produces fertilizer
Economic Importance for Humans: Negative
Artiodactyls
, especially domesticated species, sometimes carry diseases that are transmissible
to humans or other domestic animals. Wild
artiodactyls
sometimes interfere with farming operations by eating crops.
- Negative Impacts
-
injures humans
- carries human disease
- crop pest
- causes or carries domestic animal disease
Conservation Status
Currently, the IUCN classifies 54 cetartiodactyl species as data deficient, 146 as
lower risk, 40 as vulnerable, 32 as endangered, 14 as critically endangered, 7 as
extinct, and 2 as extinct in the wild. The biggest threats to many terrestrial species
are habitat loss and fragmentation and overhunting. Several large
cetacean
species were hunted nearly to extinction until an international treaty banned commercial
whaling in the 1980s. Populations of many large, commercially-valuable whales remain
severely depleted. Today,
cetaceans
face threats associated with global climate change, which could have widespread impacts
on their food supply in the near future.
Additional Links
Contributors
Tanya Dewey (editor), Animal Diversity Web.
Allison Poor (author), University of Michigan-Ann Arbor.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Palearctic
-
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oriental
-
found in the oriental region of the world. In other words, India and southeast Asia.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Ethiopian
-
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Neotropical
-
living in the southern part of the New World. In other words, Central and South America.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Australian
-
Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.
- introduced
-
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
- Arctic Ocean
-
the body of water between Europe, Asia, and North America which occurs mostly north of the Arctic circle.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Atlantic Ocean
-
the body of water between Africa, Europe, the southern ocean (above 60 degrees south latitude), and the western hemisphere. It is the second largest ocean in the world after the Pacific Ocean.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Pacific Ocean
-
body of water between the southern ocean (above 60 degrees south latitude), Australia, Asia, and the western hemisphere. This is the world's largest ocean, covering about 28% of the world's surface.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- holarctic
-
a distribution that more or less circles the Arctic, so occurring in both the Nearctic and Palearctic biogeographic regions.
Found in northern North America and northern Europe or Asia.
- cosmopolitan
-
having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- polar
-
the regions of the earth that surround the north and south poles, from the north pole to 60 degrees north and from the south pole to 60 degrees south.
- terrestrial
-
Living on the ground.
- saltwater or marine
-
mainly lives in oceans, seas, or other bodies of salt water.
- freshwater
-
mainly lives in water that is not salty.
- tundra
-
A terrestrial biome with low, shrubby or mat-like vegetation found at extremely high latitudes or elevations, near the limit of plant growth. Soils usually subject to permafrost. Plant diversity is typically low and the growing season is short.
- taiga
-
Coniferous or boreal forest, located in a band across northern North America, Europe, and Asia. This terrestrial biome also occurs at high elevations. Long, cold winters and short, wet summers. Few species of trees are present; these are primarily conifers that grow in dense stands with little undergrowth. Some deciduous trees also may be present.
- desert or dunes
-
in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- chaparral
-
Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- rainforest
-
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
- scrub forest
-
scrub forests develop in areas that experience dry seasons.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- pelagic
-
An aquatic biome consisting of the open ocean, far from land, does not include sea bottom (benthic zone).
- coastal
-
the nearshore aquatic habitats near a coast, or shoreline.
- agricultural
-
living in landscapes dominated by human agriculture.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- estuarine
-
an area where a freshwater river meets the ocean and tidal influences result in fluctuations in salinity.
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- sexual ornamentation
-
one of the sexes (usually males) has special physical structures used in courting the other sex or fighting the same sex. For example: antlers, elongated tails, special spurs.
- monogamous
-
Having one mate at a time.
- polyandrous
-
Referring to a mating system in which a female mates with several males during one breeding season (compare polygynous).
- polygynous
-
having more than one female as a mate at one time
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- year-round breeding
-
breeding takes place throughout the year
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- internal fertilization
-
fertilization takes place within the female's body
- viviparous
-
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
- young precocial
-
young are relatively well-developed when born
- natatorial
-
specialized for swimming
- diurnal
-
- active during the day, 2. lasting for one day.
- nocturnal
-
active during the night
- crepuscular
-
active at dawn and dusk
- motile
-
having the capacity to move from one place to another.
- nomadic
-
generally wanders from place to place, usually within a well-defined range.
- migratory
-
makes seasonal movements between breeding and wintering grounds
- sedentary
-
remains in the same area
- solitary
-
lives alone
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- dominance hierarchies
-
ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- scent marks
-
communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- echolocation
-
The process by which an animal locates itself with respect to other animals and objects by emitting sound waves and sensing the pattern of the reflected sound waves.
- chemical
-
uses smells or other chemicals to communicate
- filter-feeding
-
a method of feeding where small food particles are filtered from the surrounding water by various mechanisms. Used mainly by aquatic invertebrates, especially plankton, but also by baleen whales.
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- food
-
A substance that provides both nutrients and energy to a living thing.
- ecotourism
-
humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.
- causes or carries domestic animal disease
-
either directly causes, or indirectly transmits, a disease to a domestic animal
- carnivore
-
an animal that mainly eats meat
- piscivore
-
an animal that mainly eats fish
- insectivore
-
An animal that eats mainly insects or spiders.
- molluscivore
-
eats mollusks, members of Phylum Mollusca
- scavenger
-
an animal that mainly eats dead animals
- herbivore
-
An animal that eats mainly plants or parts of plants.
- folivore
-
an animal that mainly eats leaves.
- frugivore
-
an animal that mainly eats fruit
- omnivore
-
an animal that mainly eats all kinds of things, including plants and animals
- planktivore
-
an animal that mainly eats plankton
- mycophage
-
an animal that mainly eats fungus
References
Boisserie, J., F. Lihoreau, M. Brunet. 2005. The position of Hippopotamidae within Cetartiodactyla. Proceedings of the National Academy of Sciences , 102: 1537-1541.
Carey, J., D. Judge. 2002. "Longevity Records: Life Spans of Mammals, Birds, Amphibians, Reptiles, and Fish" (On-line). Accessed September 14, 2005 at http://www.demogr.mpg.de/ .
Gatesy, J., C. Hayashi, M. Cronin, P. Arctander. 1996. Evidence from milk casein genes that cetaceans are close relatives of hippopotamid artiodactyls. Molecular Biology and Evolution , 13(7): 954-963.
Gatesy, J., M. Milinkovitch, V. Waddell, M. Stanhope. 1999. Stability of cladistic relationships between Cetacea and higher-level artiodactyl taxa. Systematic Biology , 48(1): 6-20.
Gatesy, J. 1997. More DNA support for a Cetacea/Hippopotamidae clade: The blood-clotting protein gene Y-Fibrinogen. Molecular Biology and Evolution , 14(5): 537-543.
George, J., J. Bada, J. Zeh, L. Scott, S. Brown. 1999. ge and Growth Estimates of Bowhead Whales (Balaena mysticetus) Via Aspartic Acid Racemization. Canadian Journal of Zoology , 78: 1182-1198.
Graur, D., D. Higgins. 1994. Molecular evidence for the inclusion of cetaceans within the order Artiodactyla. Molecular Biology and Evolution , 11(3): 357-364.
IUCN, 2004. "2004 IUCN Red List of Threatened Species" (On-line). Accessed September 14, 2005 at www.redlist.org .
Milinkovitch, M., J. Thewissan. 1997. Even-toed fingerprints on whale ancestry. Nature , 388: 622-624.
Montgelard, C., F. Catzeflis, E. Douzery. 1997. Phylogenetic relationships of artiodactyls and cetaceans as deduced from the comparison of cytochrome b and 12S rRNA mitochondrial sequences. Molecular Biology and Evolution , 14(5): 550-559.
Naylor, G., D. Adams. 2001. Are the fossil data really at odds with the molecular data? Morphological evidence for Cetartiodactyla phylogeny reexamined. Systematic Biology , 50(3): 444-453.
Nowak, R. 1999. Walker's Mammals of the World, vol. II . Baltimore and London: The Johns Hopkins University Press.
O'Leary, M., J. Geisler. 1999. The position of Cetacea within Mammalia: Phylogenetic analysis of morphological data from extinct and extant taxa. Systematic Biology , 48(3): 455-490.
Rice, D. 1984. Cetaceans. Pp. 447-490 in Orders and Families of Recent Mammals of the World . New York: John Wiley and Sons.
Roberts, L., J. Janovy, Jr.. 2000. Foundations of Parasitology . New York: McGraw-Hill.
Shimamura, M., H. Yasue, K. Ohshima, H. Abe, H. Kato, T. Kishiro, M. Goto, I. Munechika, N. Okada. 1997. Molecular evidence from retroposons that whales form a clade within even-toed ungulates. Nature , 388: 666-670.
Simpson, C. 1984. Artiodactyls. Pp. 563-587 in Orders and Families of Recent Mammals of the World . New York: John Wiley and Sons.
Thewissen, J., E. Williams, S. Hussain. 2001. Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls. Nature , 413: 277-281.