Geographic Range
Alces alces
, commonly called moose in North America and Eurasian elk in Europe, have a circumpolar
distribution in the boreal forests of the Northern Hemisphere. (Eurasian elk should
not be confused with North American elk,
Cervus canadensis
, which are a different species.) In Eurasia,
Alces alces
have a range on the west from Scandinavia, Poland and southern Czech Republic to
Siberia (Russia) in the east. Its southern range extends to Ukraine, northern Kazakhstan,
northern China and northern Mongolia. In North America, moose are found throughout
much of Alaska and Canada and just south of the border between the contiguous United
States and Canada, but extend farther southward down the Rocky Mountains to Utah and
Colorado. Factors that likely limit their northern distribution are sufficient forage
and snow depths greater than 70 cm for long periods. Warm climates that have temperatures
above 27 C for long periods likely limit their southern distribution.
- Biogeographic Regions
- nearctic
- palearctic
- Other Geographic Terms
- holarctic
Habitat
Moose can be found in a range of habitats in the cold, northern regions of the globe
that have seasonal snow cover. They inhabit the taiga and temperate forest biomes,
including the tundra-subalpine zone. Forest types include boreal, broadleaf and mixed
(coniferous-deciduous). Within these forests, they prefer the early successional stage,
where forage is in abundance due to disturbance. Fire, logging, flooding, or glacial
action greatly increase the quality and quantity of forage for moose and, ultimately,
moose density. Besides forested habitats, moose will seek out areas near water, such
as ponds, lakes, rivers and swamps, which also have a concentration of their favorite
foods. Their large bodies, inability to sweat, and the heat produced by fermentation
in their guts mean they cannot tolerate temperatures exceeding 27 degrees Celsius
for long.
- Habitat Regions
- temperate
- terrestrial
- Terrestrial Biomes
- tundra
- taiga
- forest
- scrub forest
- mountains
- Aquatic Biomes
- lakes and ponds
- rivers and streams
- Other Habitat Features
- riparian
Physical Description
Moose are the largest species in the
deer
family, having long, slender legs that support a massive body, while a short, thick
neck and humped shoulders that support a large head. This horse-size species can be
as tall as 2.3 m at the shoulders with long ears (250 mm) and an inconspicuous tail
(80 to 120 mm). One of the most distinctive features of this species is its long,
bulbous, drooping muzzle. The upper lip overhangs the lower lip and between its nostrils
is a triangular patch of bare skin. Under the neck hangs a flap of furred skin called
the bell, which may or may not be present in females. With no upper incisors or canines,
moose must nip off plants between a bony upper palate and their lower incisors. They
have a dental formula of I 0/3, C 0/1, P 3/3, M 3/3 = 32.
Males are distinguished by carrying the largest antlers of any mammal, which can weigh
as much as 35 kg in North American moose. Antlers are grown in the spring and shed
in the winter each year. The widest antler spread recorded is 2048 mm. Antlers of
North American moose are palmate, having a main palm and brow palm in a butterfly
configuration. A cervine-shape, without palmation, is more common in European moose
(
Alces alces alces
) and Manchurian moose (
A. a. cameloides
).
Moose are sexually dimorphic, with males being more than 40% heavier than females.
Live weights of males range from 360 to 600 kg with lengths from 2.4 to 3.1 m. Females
range from 270 to 400 kg with lengths 2.3 to 3.0 m. The largest subspecies of moose,
Alces alces gigas
, occurs in Alaska with a maximum weight of 771 kg for a male and 573 kg for a female.
Pelage is generally dark, black to brown or grayish brown, with the lower legs being
lighter. An all-white color phase is rare. Their underfur and long guard hairs provide
excellent insulation from cold. Young have a reddish brown pelage and are not spotted
like other young in the deer family. Individual hairs are 15 to 25 cm long and hollow,
resulting in excellent insulation.
- Other Physical Features
- endothermic
- homoiothermic
- bilateral symmetry
- Sexual Dimorphism
- male larger
- ornamentation
Reproduction
Moose are polygynous. Males and females attract each other by making vocalizations
and scent marking trees. Moose exhibit two different types of breeding strategies:
tundra moose in Alaska form harems and taiga moose form transient pair bonds. In the
harem mating system, the largest, most dominant male attempts to herd a group of females
together, which he defends from all other males. Other bulls challenge the harem master
for the right to mate. Young bulls with smaller antlers typically retreat from the
dominant bull, whereas evenly sized bulls will fight. Bulls will engage their antlers,
pushing and twisting, while attempting to gore each other. In the pair bonding system,
a dominant bull stays with and defends just one cow until he can mate with her. Afterward,
he searches for other females who have not yet been bred by other bulls.
- Mating System
- polygynous
Moose breed in September and October of each year. The female estrous cycle lasts
24 to 25 days, with the length of the heat being 15 to 26 hours. If the female is
not bred within this time, she will recycle through estrus in about three weeks. Gestation
averages about 231 days with cows giving birth to one calf on average, although twins
are common. Calves are born at an average weight of 16.2 kg and gain approximately
1 kg per day while they are nursing. Males and females are sexually mature at two
years of age but full growth potential isn't reached until 4 or 5 years of age. At
that age females are at their reproductive peak and males have the largest antlers.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- viviparous
Only females take care of their young for a period of one year. Females seek secluded
sites to give birth to young and remain isolated until the calves are weaned. Calves
can browse and follow their mother at 3 weeks old and are weaned at 5 months. They
remain with their mother until about 1 year after their birth, when the mother's next
young is born.
- Parental Investment
- precocial
-
pre-fertilization
- provisioning
-
protecting
- female
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- female
-
protecting
- female
-
provisioning
-
pre-independence
-
protecting
- female
-
protecting
- inherits maternal/paternal territory
- maternal position in the dominance hierarchy affects status of young
Lifespan/Longevity
Many calves, as much as 50% or more, do not live beyond their first six weeks of life
due to predation by bears and wolves. Once they reach adulthood, their chances of
survival are high. Adult females have an average survival of 95%. Male survival is
more variable due to hunting and male-male competition. Adult moose are in their prime
from 5 to 12 years of age but begin to suffer from arthritis, dental diseases and
wear, and other factors after about 8 years. Few moose live past 15 years in the wild,
although one cow was recorded to have lived to the age of 22. Peak reproductive age
in females is 4 to 12 years of age and 4 to 8 years in males.
Behavior
Moose are the least social species among cervids, remaining fairly solitary except
during the mating season. They are not territorial. Outside of the rutting period,
males and females are sexually segregated: males and females are separated spatially,
temporally, and/or by habitat. It has been hypothesized that this is due to the differences
in nutritional needs of the sexes due to body size differences. Also, cows with calves
at heel seclude themselves from conspecifics to reduce the risk of being singled out
by predators. Moose are crepuscular by nature, being most active at sunrise and sunset.
Despite their ungainly appearance, moose are able to run silently through dense forests.
Maximum speeds have been clocked at 56 km/hr and sustained speeds at 9.6 km/hr. Moose
are also strong swimmers, being known to swim up to 20 km or up to 9.5 km per hour.
Most of their time centers around feeding. The daily pattern is traveling to a new
site to feed, avoiding predators, browsing on plants, standing, and lying down for
the rumination of their food. Moose mainly stay in the same general area, though some
populations migrate between sites favorable at different times of the year. These
migrations can exceed 300km in European populations.
- Key Behaviors
- cursorial
- terricolous
- diurnal
- crepuscular
- motile
- migratory
- sedentary
- solitary
- dominance hierarchies
Home Range
Home range size of moose varies between 3.6 to 92 km2. During their first year of
life, young moose occupy the same home range as their mother and do not establish
their own home range until the age of two. Home range size of males tend to be larger
than females. Some moose migrate seasonally, up to 179 km in North America and 300
km in northeastern Europe.
Communication and Perception
Moose are not as vocal as other members of the
deer
family, such as elk (
Cervus elaphus
). Most of their vocalizations occur during the rut. Females make a long, quavering
moaning call when in estrus, which attracts males and can be heard up to 3.2 km away.
Males make a grunt to court females or challenge other bulls. Both sexes are capable
of making a loud, guttural “roaring” sound as a threat. Moose will also communicate
chemically by scent-marking trees. They strip trees and shrubs of their bark and rub
their foreheads and preorbital glands into the bare spot to advertise their presence
to the opposite sex. Moose are not known to have good eye sight. Confirmation of this
is not available due to a lack of studies, although it is known that they depend less
on sight than their sense of hearing or smell. Moose have very acute hearing due in
part to the large external surface area of their ears. Their ears are capable of rotating
independently, giving them stereophonic hearing. Their sense of smell is exceptional
due to the large surface area of their nasal cavities, which are lined with millions
of sent-smelling cells.
- Other Communication Modes
- pheromones
- scent marks
Food Habits
The word “moose” comes from the Native American tribe, the Algonquins, which means
“twig eater” in their language. It is an appropriate name because moose primarily
browse upon the stems and twigs of woody plants in the winter and the leaves and shoots
of deciduous plants in the summer. For moose in Poland, 87% of their diet consisted
of trees and shrubs with the most important species being pine (
Pinus silvestris
), which represented 52% of their diet. In North America, moose have been observed
to consume as many as 221 plant species and genera, and in Russia 355, although only
a select few comprise a significant portion of their diet. Willows (
Salix
spp.) are the most preferred forage where available. In interior Alaska willows accounted
for 94% of the biomass consumed in the winter. Other species that are consumed are
paper birch (
Betula papyrifera
), quaking aspen (
Populus tremuloides
), and balsam fir (
Abies balsamea
). Moose also consume aquatic vegetation during the summer because it is highly digestible
and abundant in some areas. An adult requires about 20 kg food per day.
- Plant Foods
- leaves
- wood, bark, or stems
- flowers
Predation
Wolves, brown or grizzly bears, black bears, and cougars are major predators of moose,
and to a lesser extent coyotes and Amur tigers. Major predators can have such an impact
on moose populations that they can slow their population growth and hold them below
the carrying capacity of the habitat. Whether or not predators can hold (regulate)
a moose population at an equilibrium point is controversial. Predation by bears tends
to be the highest in the spring when calves are the most vulnerable. Predation by
wolves is higher in the winter when snow depths are high enough to impede the movement
of moose. Moose aggressively defend themselves and their young with their robust antlers
and sharp hooves.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Moose can have a significant impact on ecosystems because of their daily activities
of feeding, trampling, defecating, and urinating. One researcher estimated that the
Swedish moose population contributed 300,000 metric tons of feces each year to the
land. This equals about 5,600 tons of nitrogen, which is essential for plant growth.
Moose can affect the rate of nutrient cycling, floral composition, rate of forest
succession, and biological diversity of a forest. For this reason, they are considered
to be a keystone species.
Moose are affected by several diseases and parasites. "Moose disease", fatal to moose,
is caused by a brainworm which most commonly infects
white-tailed deer
. Moose can become severely infested with winter ticks and death can sometimes result
in winter as a result of blood loss and nutritional stress.
- Ecosystem Impact
- creates habitat
- keystone species
- winter ticks ( Dermacentor albipictus )
- brainworm ( Parelaphostrongylus tenuis )
- tapeworms ( Taenia krabbei )
- hydatid disease ( Echinococcus granulosus )
Economic Importance for Humans: Positive
Moose are hunted throughout much of their range and provide millions of pounds in
meat to humans each year. In 1983, 152,000 moose were harvested in Sweden, representing
14% of the total meat consumption of the country. Moose hunting also generates a considerable
amount of money to local economies. Moose hunters contribute $31 million annually
to Alaska’s economy and $50 million to Canada’s. Moose also draw many tourists for
wildlife viewing opportunities. Moose milk is harvested from captive moose in some
areas, including Russia and Sweden.
- Positive Impacts
- food
- body parts are source of valuable material
- ecotourism
- produces fertilizer
Economic Importance for Humans: Negative
Vehicle collisions with moose are a serious problem in North America and Europe. In
North America, nearly 3,000 occur each year. In Sweden alone, 4,500 occur per year
on average, as well as 10 to 15 human fatalities. Beyond the incalculable cost in
terms of human life, moose collisions cause significant property damage, escalate
insurance premiums, can cause a local moose population to decline and reduce recreational
opportunities. Moose can also be a pest to agriculture and forestry in some areas.
Moose foraging can inhibit the growth of young trees. In Russia, damage to the forestry
industry was estimated in millions of rubles, during the 1950's.
- Negative Impacts
- injures humans
- crop pest
Conservation Status
IUCN lists moose as a species of “Least Concern” because a majority of populations
are expanding and extremely abundant despite heavy hunting pressure in parts of their
range. The Eurasian populations are estimated to be 1.5 million individuals, one-third
of which consists of the European populations. In 2002, populations of the United
States and Canada were estimated at 1,000,000. Some exceptions to this favorable assessment
are moose in Manchuria and Mongolia (
A. a. cameloides
), which are scarce, and a population in Nova Scotia (
A. a. americana
), which Canada has deemed endangered.
Other Comments
Alces
, which has been traditionally classified as a monotypic genus, has been recently
divided into two species by some researchers: the European species,
Alces alces
, and the North American species,
Alces americanus
. Such a division has caused confusion and does not represent the latest genetic research.
The division of moose into European and North American species was believed to be
supported by morphological differences and evidence that European moose have 68 chromosomes
and North American moose have 70. It is now known that moose in Central Asia (Yakutia)
also have 70 chromosomes and share mitochondrial DNA halotypes with European and North
American moose. Furthermore, moose exhibit low variability in mitochondrial DNA worldwide
and have relatively low overall genetic diversity compared to other mammals. It is
more likely that all extant lineages of moose originated from Central Asia within
the last 60,000 years, supporting a single species hypothesis rather than a two or
three species hypothesis.
Additional Links
Contributors
Tanya Dewey (editor), Animal Diversity Web.
Daniel De Bord (author), University of Alaska Fairbanks, Link E. Olson (editor, instructor), University of Alaska Fairbanks.
- 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.
- 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.
- 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.
- 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).
- terrestrial
-
Living on the ground.
- 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.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- 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.
- marsh
-
marshes are wetland areas often dominated by grasses and reeds.
- swamp
-
a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.
- bog
-
a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- 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.
- polygynous
-
having more than one female as a mate at one time
- 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
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- 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
- diurnal
-
- active during the day, 2. lasting for one day.
- crepuscular
-
active at dawn and dusk
- motile
-
having the capacity to move from one place to another.
- migratory
-
makes seasonal movements between breeding and wintering grounds
- sedentary
-
remains in the same area
- solitary
-
lives alone
- 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
- pheromones
-
chemicals released into air or water that are detected by and responded to by other animals of the same species
- 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
- chemical
-
uses smells or other chemicals to communicate
- 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.
- keystone species
-
a species whose presence or absence strongly affects populations of other species in that area such that the extirpation of the keystone species in an area will result in the ultimate extirpation of many more species in that area (Example: sea otter).
- 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.
- herbivore
-
An animal that eats mainly plants or parts of plants.
- folivore
-
an animal that mainly eats leaves.
References
Ballard, W., V. Van Ballenberghe. 2007. Predator-Prey Relationships. Pp. 247-273 in Ecology and Management of the North American Moose . Boulder, Colorado: University Press of Colorado.
Bowyer, R., V. Van Ballenberghe, J. Kie. 2003. Moose: Alces alces . Pp. 931-964 in Wild mammals of North America: Biology, management and conservation . Baltimore, MD: John Hopkins University Press.
Boyer, R. 2004. Sexual segregation in Ruminants: definitions, hypotheses, and implications for conservation and management.. Journal of Mammalogy , 85: 1039-1052.
Bubenik, A. 2007. Behavior. Pp. 173-221 in Ecology and Management of the North American Moose . Boulder, Colorado: University Press of Colorado.
Child, K. 2007. Incidental mortality. Pp. 275-301 in Ecology and Management of the North American Moose . Boulder, Colorado: University Press of Colorado.
Engan, J. 2001. Changes in the relationship between palmate and cervine antlers in moose in southeastern Norway. Alces , 37: 79-88.
Franzmann, A. 1981. Alces alces. Mammalian Species , 154: 1-7.
Franzmann, A. 2000. Moose. Pp. 578-600 in Ecology and management of large mammals in North America . Upper Saddle River, NJ: Prentice-Hall.
Gaillard, J., M. Festa-Bianchet, N. Yoccoz. 1998. Population Dynamics of Large Herbivores: Variable Recruitment with Constant Adult Survival. Trends in Ecology & Evolution , 13: 58-63.
Geist, V., M. Ferguson, J. Rachlow. 2008. "Alces americanus" (On-line). IUCN Red List of Threatened Species. Accessed November 16, 2008 at http://www.iucnredlist.org/details/818 .
Grubb, P. 2005. Artiodactyla: Cervidae: Capreolinae. Pp. 652-653 in Mammal species of the world: a taxonomic and geographic reference . Baltimore, MD: The Johns Hopkins University Press.
Henttonen, H., M. Stubbe, T. Maran, A. Tikhonov. 2008. "Alces alces" (On-line). 2008 IUCN Red List of Threatened Species. Accessed November 16, 2008 at http://www.iucnredlist.org/details/41782 .
Hundertmark, K., R. Bowyer. 2004. Genetics, evolution, and phylogeography of moose. Alces , 40: 103-122.
Hundertmark, K., G. Shields, I. Udina, R. Bowyer, A. Danilkin, C. Schwartz. 2002. Mitochondrial phylogeography of moose (Alces alces): late Pleistocene divergence and population expansion. Molecular Phylogenetics and Evolution , 22: 375-387.
Hundertmark, K. 2007. Home range, dispersal and migration. Pp. 303-335 in Ecology and management of the North American moose . Boulder, CO: University Press of Colorado.
Karns, P. 2007. Population distribution, density, and trends. Pp. 125-139 in Ecology and Management of the North American Moose . Boulder, CO: University Press of Colorado.
Kelsall, J., E. Tefler. 1974. Biogeography of moose with particular reference to western North America. Naturaliste Canadien , 101: 117-130.
Kie, J., R. Bowyer, K. Stewart. 2003. Ungulates in western forests: Habitat requirements, population dynamics, and ecosystem processes. Pp. 296-340 in Mammal community dynamics in western coniferous forests: Management and conservation . Baltimore, MD: Johns Hopkins University Press.
Kielland, K., J. Bryant. 1998. Moose herbivory in taiga: effects on biogeochemistry and vegetation dynamics in primary succession. Oikos , 82: 377-383.
Kuznetsov, G. 2002. Moose and forest problems in Russia. Alces supplement , 2: 65-70.
LeResche, R. 1974. Moose migrations in North America. Naturaliste Canadien , 101: 393-415.
Miquelle, D., E. Smirnov, H. Quigley, M. Hornocker, E. Nikolaev, E. Matyushkin. 1996. Food habits of Amur tigers in Sikhote-Alin Zapovednik and the Russian Far East, and implications for conservation. Journal of Wildlife Research , 1: 138-147.
Molvar, E., R. Bowyer, V. Van Ballenberghe. 1993. Moose herbivory, browse quality, and nutrient cycling in an Alaskan treeline community. Oecologia , 94: 472-479.
Novak, R. 1999. Walker's Mammals of the World . Baltimore, MD: The John Hopkins University Press.
Peek, J. 2007. Habitat relationships. Pp. 351-375 in Ecology and management of North American moose . Boulder, CO: University Press of Colorado.
Persson, I., K. Danell, R. Bergstrom. 2000. Disturbance by large herbivores in Boreal forests with special reference to moose. Annales Zoologici Fennici , 37: 251-263.
Pulliainen, E. 1974. Seasonal movements of moose in Europe. Naturaliste Canadien , 101: 379-392.
Regelin, W., A. Franzmann. 1998. Past, present and future moose management and research in Alaska. Alces , 34: 279-286.
Renecker, L., C. Schwartz. 2007. Food habits and feeding behavior. Pp. 403-439 in Ecology and Management of the North American Moose . Boulder, CO: University Press of Colorado.
Schwartz, C., K. Hundertmark. 1993. Reproductive characteristics of Alaskan moose. Journal of Wildlife Management , 57: 454-468.
Schwartz, C. 2007. Reproduction, natality, and growth. Pp. 141-171 in Ecology and Management of the North American Moose . Boulder, CO: University Press of Colorado.
Schwartz, C. 1992. Reproductive biology of North American moose. Alces , 28: 165-173.
Seiler, A. 2005. Predicting locations of moose-vehicle collisions in Sweden. Journal of Applied Ecology , 42: 371-382.
Timmermann, H. 2003. The status and management of moose in North America circa 2000-01. Alces , 39: 131-151.
Wilson, D., S. Ruff. 1999. The Smithsonian Book of North American Mammals . Washington, D.C.: Smithsonian Institution Press.