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Pipilo erythrophthalmuseastern towhee

By Charnele Johnson

Geographic Range

Eastern towhees (Pipilo erythrophthalmus) have a geographic range that includes much of North America. Year-round, eastern towhees range as far north as Delaware and as far south as Florida in the United States. Their range extends westward as far as Louisiana and Arkansas. Towhees also take up residence in the southernmost part of British Columbia, Canada and extend south into California and small portions of Texas. They are also found in central Mexico and Guatemala. A small isolated population is also found near the tip of Baja California Sur.

Eastern towhees overwinter in southern and central parts of the United States (e.g., parts of Louisiana, Texas, and southern Oklahoma) and parts of Mexico (e.g., Chihuahua, Coahuila, Monterrey, and Ciudad Victoria).

During their breeding months in summer, eastern towhees inhabit the Rocky Mountains and Great Plains areas (e.g., Idaho, Montana, Wyoming, and British Columbia). Their summer range also includes the Appalachian Mountains (e.g., New York, Pennsylvania, and southern parts of Maine) and southern Canada (e.g., the provinces of Quebec, Ontario, Saskatchewan, Alberta, and British Columbia). (BirdLife International, 2012; Greenlaw, 1996; Root, 1988)

Habitat

Eastern towhees spend much of their time on or near the ground. They commonly nest and forage in areas of open woods, undergrowth, bushy edges, and areas with leaf litter. Eastern towhees do not favor any particular shrub species in their habitat, though they generally live in areas with small shrubs and developed undergrowth.

During winter, eastern towhees are typically found in areas with dense shrubs and leaf litter. Secondary succession and mountainous areas are ideal habitats for eastern towhees because they typically have small canopy trees that allow access to sunlight. Mountain ranges in southern and southwestern North America are examples of areas in which eastern towhees are commonly found.

During their summer breeding months, towhees are typically found in areas similar to the habitats they use in winter that are rich in undergrowth and ground litter. Outside of their breeding months, towhees can be found in wet, isolated areas, such as wetland thickets. However, they may still be found in areas similar to their breeding habitat. Indeed, eastern towhees outside of breeding season have been observed using areas that were confirmed to be used as breeding areas. (Greenlaw, 1996; Peterson, 1980)

Physical Description

Eastern towhees are typically 18 to 21 cm tall with an average wingspan of 20 cm. Adults weigh an average of 40 g, although males are slightly larger than females.

Eastern towhees are sexually dimorphic, meaning different sexes and stages of maturity can be distinguished by their size and the color of their feathers. Adult males have black on the tops of their heads and their backs. They have white underbellies and thick rufous stripes along their sides and underneath their wings. Adult females are nearly identical to adult males, although they have brown coloration where males have black coloration. Eastern towhees commonly have red eyes, but white-eyed variants exist along the southern Atlantic coast and in Florida. Juvenile eastern towhees are altricial when they hatch, with very few feathers, pink skin, and closed eyes. Juvenile males are brownish-black in color and juvenile females are light brown. Juveniles of both sexes also have lighter streaks along their bodies, whereas adults are solid black or brown, depending on their sex. (Barbour, 1950; Greenlaw, 1996; Peterson, 1980; Simpson and Simpson, 2013)

  • Sexual Dimorphism
  • male larger
  • sexes colored or patterned differently
  • male more colorful
  • Average mass
    40 g
    1.41 oz
  • Range length
    18 to 21 cm
    7.09 to 8.27 in
  • Average length
    20 cm
    7.87 in
  • Average wingspan
    20 cm
    7.87 in

Reproduction

Eastern towhees are monogamous and breed seasonally from early March to late July, depending on how far they migrate. They typically begin breeding earlier in warmer, southern states and later in northern states. For example, they begin breeding in early March in Georgia, Florida, and Alabama, whereas they do not begin nesting and breeding in New England states until late May.

Males typically start the mating process with aggressive behaviors towards potential mates. These aggressive behaviors include producing muted primary songs, raising their tail feathers, and flapping their wings. Males typically select one female and pursue them throughout the surrounding area for an unspecified amount of time. While following females, males exhibit other mating behaviors such as complex song displays, debris carrying, and growling. Over time, these behaviors become infrequent and eventually stop. Females engage in song as well, but are passive in their overall behavior. Following successful courtship, paired males and females become inseparable and females typically begin producing whinny calls frequently. The exact amount of time mating behaviors are exhibited is unknown.

After they select a mate, female eastern towhees begin searching for a site to build their nests. Females spend 5 days building their nests alone. During this time, males are most commonly seen closely following their mate. Copulation does not occur until nest-building begins and is initiated by males. In response, females display copulatory postures consisting of head and tail elevation, back arching, and wing quivering. During copulation, males mount females for 2 to 3 seconds before either jumping off or flying away. Copulation occurs on the ground, on logs, or on perches as high as 10 m above the ground.

Once egg-laying has begun, paired males and females do not engage in further copulation. Any initiations of copulation by males are commonly rejected by female after they have laid their eggs. (BirdLife International, 2012; Greenlaw, 1996; Root, 1988)

Eastern towhees begin their breeding season in early March in the southern United States (e.g., Georgia and Florida), mid to late April in the Midwest, and early May in the northern parts of their range (e.g., southern Canada and in New England states). The delay in mating behavior farther north occurs because it remains colder for longer and individuals must migrate farther to reach northern parts of their range. The latitude and local climate also impacts the duration of the breeding season and the number of broods. In the southern United States, eastern towhees have a longer breeding season and typically have more than one brood per year. Towhees build nests about 1 m above the ground. Nests are usually composed of leaf litter, tree branches, and bark. Those that are low to the ground are hidden by leaf litter. Males do not contribute to nest building.

Egg-laying typically occurs early in the morning, with females laying one egg per day until their clutch is complete. Females usually have a single clutch of 3 to 5 eggs. Eggs incubate in the nest for 12 to 13 days before hatching. Hatchlings weigh about 3 g at birth and typically take 7 to 10 days to gain weight, grow their feathers, and become fledglings. Fledglings quickly learn to fly and, once they are volant, fledglings leave the nest. However, parents often continue to provide food for their offspring for 3 to 4 weeks after they leave the nest. (BirdLife International, 2012; Greenlaw, 1996; Robbins and Blom, 1996; Root, 1988)

  • Breeding interval
    Eastern Towhees normally breed once per year in the spring.
  • Breeding season
    Early March to early May
  • Range eggs per season
    3-4 to 2-6
  • Average eggs per season
    3-4
  • Range time to hatching
    12 to 13 days
  • Average time to hatching
    12 days
  • Average fledging age
    10 days
  • Average time to independence
    3-4 weeks
  • Average age at sexual or reproductive maturity (female)
    1 years
  • Average age at sexual or reproductive maturity (male)
    1 years

Both male and female eastern towhees take care of their nestlings after hatching and also for 3 to 4 weeks after they leave the nest. Females build nests, lay their eggs, and incubate them. Males protect their mate and developing eggs by sitting on the rim of the nest or perching nearby while females are incubating eggs. Males also leave food in or around the nest before eggs hatch and, on rare occasions, they provide food directly to their mates. Before nestlings hatch, if females must leave the nest to forage for themselves, their mate will follow and guard them from predators.

After eggs hatch, both parents feed nestlings and work to keep their nests clean. To prevent clutter, females typically eats the shells from which nestlings hatched. Both parents continue to feed and care for their offspring after they have left the nest. Juvenile eastern towhees will return to the nest to be fed by their parents, and become fully independent after about 1 month.

Depending on the length of the breeding season and the abundance of food in their area, eastern towhees may produce a second brood of eggs. Generally, by the time females have a second brood the first brood has already fledged. However, if fledglings are still in the nest when the second brood hatches, parents engage in parental division, wherein males care for the fledglings from the first brood while female care for the second brood. Parental division is also common when a brood parasite is present. While females care for their offspring, males care for any brood parasite offspring. (BirdLife International, 2012; Greenlaw, 1996)

  • Parental Investment
  • altricial
  • pre-fertilization
    • protecting
      • male
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • male
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • male
  • pre-independence
    • provisioning
      • female
    • protecting
      • male

Lifespan/Longevity

Eastern towhees are reported to live an average of 6 years in the wild. The longest recorded lifespan of a wild eastern towhee was a male that lived to be about 12 years old. Not much information is known about the longevity of eastern towhees in captivity. (BirdLife International, 2012)

  • Range lifespan
    Status: wild
    12 (high) years
  • Average lifespan
    Status: wild
    6 years
  • Typical lifespan
    Status: wild
    1 to 6 years

Behavior

During breeding season, males spend their time establishing their territory, chasing or fighting intruding males, and courting females. Males are reported to adjust their territory sizes depending on population density and food abundance in a particular area. If a male is successful in mating with a female, it spends the rest of the breeding season guarding the nest and providing for its offspring until they are independent. Mated females spent most of their time incubating eggs and taking care of hatchlings.

Eastern towhees are solitary outside of mating season, spending most of their time foraging or resting. They are diurnal and are typically found hopping or walking along the ground foraging for insects or, during the breeding season, looking for materials to build nests. Eastern towhees often roost during the morning, before sunrise, in dense foliage. They are typically solitary when roosting. Eastern towhees perching in trees usually hop from perch to perch, using their wings for balance. If preparing for flight, they engage in pre-flight behaviors, such as wing-flicking. The amount and intensity of pre-flight behaviors depend on the length they are preparing to travel. If they are flying long distance, they perform undisturbed wing flaps evenly on both sides. If their destination is a short distance away, they perform periodic and uneven wing flaps. During the breeding season, eastern towhees also use uneven wing flaps to alert other eastern towhees of predators near nesting sites.

Eastern towhees frequently engage in self-maintenance behaviors, including preening, stretching, scratching, bathing, sunbathing, and anting. Preening typically occurs between bouts of song or during periods of rest. Stretching and scratching occur after long periods of rest. Bathing occurs in shallow waters, or on wet foliage if water is scarce. Eastern towhees typically wash the top half of their bodies first and their hind ends afterwards. Sunbathing typically occurs around midday. If perched in a tree, eastern towhees seek sunny areas and fluff out their feathers while facing the direction of the sun. If on the ground, they lie on their side with their wings exposed and their mouth open. Eastern towhees also engage in a behavior called "anting", in which they rub insects, often ants, on their feathers or skin. This most commonly occurs in juveniles between July and October. (Greenlaw, 1996)

  • Range territory size
    8000 to 27000 m^2

Home Range

Home ranges are rarely reported for eastern towhees. A study of two populations in 1941 reported that eastern towhees had winter home range sizes of 3.8 and 12.6 ha. Territory sizes are much smaller, with measurements of 0.8 to 1.1 ha in a California population, and 0.8 to 2.7 ha in a Kansas population. Territory size depends partly on population density and food availability in a given area. ("California wildlife and their habitats: western Sierra Nevada", 1980; Greenlaw, 1996)

Communication and Perception

Eastern towhees have a variety of songs and calls that they use for communication with their mates, offspring, and other conspecifics. Some calls are typically heard from only one sex. For instance, males produce growling calls and females make whinny calls. Nestlings and juveniles also have specific songs, such as screams and trill calls, that are associated with begging behaviors to get food from their parent. Young hatchlings with eyes that are not fully developed give trill calls for their parents. As nestlings develop and fledge, they learn more complex and situation-appropriate calls.

Eastern towhees most commonly make a “chewink” call, which serves as an alarm call to warn others of approaching predators. All eastern towhees use this call and respond to it similarly, regardless of their sex, stage in development, or the season. This call differs slightly in its sound between eastern towhee populations and its sound is interpreted differently by humans, depending on the geographic region. In the north, their call has been translated from “chewink” to “chewee”. Near the Gulf Coast the call is described as “shree” whereas in Florida it is described as “swee”. For towhees on the east coast, the sound of their trill is compared to the phrase “drink-your-tea”. (Greenlaw, 1996; Peterson, 1980; Robbins and Blom, 1996)

Food Habits

Eastern towhees are omnivores with a diet consisting of about 70% vegetation and 30% insects. They commonly eat plants such as ragweed (Ambrosia psilostachya), foxtail (genus Alopercurus), and wild fruits. They also eat various insect species, including moths and butterflies (order Lepidoptera), spiders (order Araneae), snails (class Gastropoda), grasshoppers (order Orthoptera), and beetles (order Coleoptera). The diet of eastern towhees shifts seasonally. For example, beetles compose the majority of their diet in winter, while moth and butterfly caterpillars are the most common food items during the breeding season.

Eastern towhees typically forage near the ground in densely covered areas of forests. They may also forage in open areas, but this is uncommon. Eastern towhees move along the ground and scratch at leaf litter to uncover insects. Scratching at leaves and dirt causes insects to leave areas where they normally hide from birds that search for food in the air. ("California wildlife and their habitats: western Sierra Nevada", 1980; Greenlaw, 1996; Robbins and Blom, 1996; Root, 1988)

  • Animal Foods
  • insects
  • terrestrial non-insect arthropods
  • Plant Foods
  • leaves
  • seeds, grains, and nuts
  • fruit

Predation

Eastern towhees have many predators, some of which prey on eggs and hatchlings and others which prey on adults. Known nest predators include bull snakes (Pituophis melanoleucus), western rat snakes (Pantherophis obsoleta), eastern garter snakes (Thamnophis sirtalis), and eastern chipmunks (Tamias striatus). Known predators of adult eastern towhees include short-eared owls (Asio flammeus), Cooper’s hawks (Accipiter cooperii), and short-tailed hawks (Buteo brachyurus).

To evade predators, eastern towhees initiate alarm calls to warn others of danger. They also engage in mobbing, where multiple eastern towhees attack singular predators. If eastern towhees sense a predator while foraging, they remain motionless for several minutes and wait for the predator to move on before silently escaping. If females detect a predator near their nests, they walk on the ground and drag their tails as if they are injured. By appearing injured, females can distract predators and potentially lure them away from nests. (BirdLife International, 2012)

Ecosystem Roles

Eastern towhees have important roles in their ecosystems. They serve as prey for other vertebrates, including snakes, small mammals, and other birds. They also prey on several invertebrate species and likely play a role in controlling invertebrate populations. Furthermore, eastern towhees serve as seed dispersers for some plant species. They eat fruits and seeds that drop to the ground and carry them to new areas in their digestive tracts before defecating. This aids in the distribution of seedlings, especially for those plant species that require the process of digestion to germinate.

Many parasites affect eastern towhees. Ectoparasites include chewing lice (suborder Mallophaga), ticks (subclass Acarina), and louse flies (family Hippoboscidae). Endoparasites, such as protozoans (genus Leucocytozoon) and trematodes (genus Lyperosomum), have been found in their blood and their livers, respectively. These parasites have also been found on adult eastern towhees rather than inside them.

Brood parasites also impact eastern towhees. Brown-headed cowbirds (Molothrus ater) lay their eggs in the nests of other birds, including eastern towhees. Eastern towhees cannot tell the difference between their eggs and brood parasite eggs, so they often raise cowbird offspring alongside their young. In cases where brood parasites are present, parental care is split: females feed and care for their own offspring while males feed and care for brood parasite hatchlings. (BirdLife International, 2012; Denton and Krissinger, 2011; Greenlaw, 1996)

  • Ecosystem Impact
  • disperses seeds
Commensal/Parasitic Species
  • Ticks (subclass Acarina)
  • Louse flies (family Hippoboscidae)
  • Trematodes (genus Lyperosomum)
  • Brown-headed cowbirds (Molothrus ater)
  • Chewing lice (suborder Mallophaga)
  • Protozoan (genus Leucocytozoon)

Economic Importance for Humans: Positive

Eastern towhees are common across large parts of North America, both during and after their breeding season. Because they are widespread and males have bright coloration, eastern towhees are a source of ecotourism and recreational enjoyment through bird watching. (BirdLife International, 2012; Ewert, 1979; Greenlaw, 1996)

Economic Importance for Humans: Negative

Eastern towhees are not known to negatively impact human populations. (Greenlaw, 1996)

Conservation Status

Eastern towhees are considered a species of "least concern" on the IUCN Red List. They are listed as protected under the Migratory Bird Act, which prohibits their capture and distribution in the pet trade or for other purposes. Eastern towhees have no special status in the CITES appendices or on other international or national conservation lists.

Long-term breeding surveys of eastern towhees suggest their populations have experienced significant declines in the last 50 years. It is estimated that breeding birds decline at a rate of about 1% per year and the amount of breeding individuals has decreased by 44% since the 1960s. Population declines are attributed to the destruction of suitable habitats for commercial and residential purposes. (BirdLife International, 2012; Greenlaw, 1996)

Contributors

Charnele Johnson (author), Radford University, Karen Powers (editor), Radford University, April Tingle (editor), Radford University, Emily Clark (editor), Radford University, Cari Mcgregor (editor), Radford University, Jacob Vaught (editor), Radford University, Galen Burrell (editor), Special Projects.

Glossary

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.

World Map

acoustic

uses sound to communicate

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

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.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

choruses

to jointly display, usually with sounds, at the same time as two or more other individuals of the same or different species

crepuscular

active at dawn and dusk

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.

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.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

granivore

an animal that mainly eats seeds

herbivore

An animal that eats mainly plants or parts of plants.

insectivore

An animal that eats mainly insects or spiders.

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).

marsh

marshes are wetland areas often dominated by grasses and reeds.

migratory

makes seasonal movements between breeding and wintering grounds

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

polymorphic

"many forms." A species is polymorphic if its individuals can be divided into two or more easily recognized groups, based on structure, color, or other similar characteristics. The term only applies when the distinct groups can be found in the same area; graded or clinal variation throughout the range of a species (e.g. a north-to-south decrease in size) is not polymorphism. Polymorphic characteristics may be inherited because the differences have a genetic basis, or they may be the result of environmental influences. We do not consider sexual differences (i.e. sexual dimorphism), seasonal changes (e.g. change in fur color), or age-related changes to be polymorphic. Polymorphism in a local population can be an adaptation to prevent density-dependent predation, where predators preferentially prey on the most common morph.

scrub forest

scrub forests develop in areas that experience dry seasons.

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

solitary

lives alone

suburban

living in residential areas on the outskirts of large cities or towns.

tactile

uses touch to communicate

terrestrial

Living on the ground.

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

visual

uses sight to communicate

References

USDA Forest Service. California wildlife and their habitats: western Sierra Nevada. Gen. Tech. Rep. PSW-GTR-37. Berkeley, California: Pacific Southwest Forest and Range Exp. Stn., Forest Service, U.S. Department of Agriculture. 1980. Accessed April 22, 2015 at http://www.fs.fed.us/psw/publications/documents/psw_gtr037/.

Baicich, P., C. Harrison. 1997. A Guide to the Nests, Eggs, and Nestlings of North American Birds: Second Edition. San Diego, CA: AP Natural World.

Barbour, R. 1950. Growth and feather development of towhee nestlings. American Midland Naturalist, 44/3: 742-749.

Barbour, R. 1941. Winter habits of the red-eyed towhee in eastern Kentucky. American Midland Naturalist, 26/3: 593-595.

Bell, J., R. Whitmore. 2000. Bird nesting ecology in a forest defoliated by gypsy moths. The Wilson Bulletin, 112/4: 524-531.

Bell, J., R. Whitmore. 1997. Eastern towhee numbers increase following defoliation by gypsy moths. The Auk, 114/4: 708-716.

BirdLife International, 2012. "The IUCN Red List of Threatened Species. Version 2014.3" (On-line). Pipilo erythrophthalmus. Accessed February 05, 2015 at http://www.iucnredlist.org/details/22721310/0.

Denton, F., W. Krissinger. 2011. Lyperosomum byrdi sp. n. (Dignea: Dicrocoeliidae) from the rufous-sided towhee, Pipilo erythrophthalmus (L.), with a revised synopsis of the genus. Proceedings of the Helminthological Society of Washington, 42/1: 38-42.

Ewert, D. 1979. Development of song of a rufous-sided towhee raised in acoustic isolation. Condor, 81/3: 313.

Greenlaw, J. 1996. "Eastern Towhee" (On-line). Birds of North America (on-line). Accessed February 13, 2015 at http://bna.birds.cornell.edu/bna/species/262/articles/introduction.

Greenlaw, J. 1976. Use of bilateral scratching behavior by emberizines and icterids. The Condor, 78/1: 94-97.

Krementz, D., L. Powell. 2000. Breeding season demography and movements of eastern towhees at the Savannah River Site, South Carolina. Wilson Journal of Ornithology, 112/2: 243-248.

Peterson, R. 1980. A Field Guide to the Birds. Boston, MA: Houghton Mifflin Company.

Pyle, P. 1997. Identification Guide to North American Birds: Part One. Bolinas, CA: Slate Creek Press.

Richards, D. 1981. Alerting and messaging components in songs of rufous-sided towhees. Behavior, 76/3/4: 223-249.

Robbins, C., E. Blom. 1996. Atlas of the Breeding Birds of Maryland and the District of Columbia. Pittsburgh, PA: University of Pittsburgh Press.

Root, T. 1988. Atlas of Wintering North American Birds: An Analysis of Christmas Bird Count Data. Chicago: The University of Chicago Press.

Sibley, C., D. West. 1959. Hybridization in the rufous-sided towhees of the Great Plains. The Auk, 76/3: 326-338.

Simpson, A., R. Simpson. 2013. Nature Guide to Shenandoah National Park. CT: Morris Book Publishing.

Verner, J., M. Willson. 1969. Mating system, sexual dimorphism, and the role of male North American passerine birds in the nesting cycle. Ornithological Monographs, 9: 1-76.

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