Overview

Brief Summary

Pipilo erythrophthalmus

A medium-sized (7-8 ½ inches) songbird, the male Eastern Towhee is most easily identified by its black head and body, black breast, pale belly, and rufous flanks. Female Eastern Towhees are similar but paler, with the black replaced by brown. Most individuals have red eyes, although birds from Florida and Georgia tend to have white eyes instead. The Eastern Towhee breeds across much of the eastern United States and southern Canada west to Nebraska and Saskatchewan. In winter, many northern birds move south to the southern part of this species’ range as well as east Texas, Oklahoma, and southern Louisiana, where this species does not breed. Southern birds generally migrate short distances, if at all. Eastern Towhees breed in a variety of woodland habitats, particularly in heavily-vegetated undergrowth near forest edges or clearings. This species utilizes similar habitats in summer as in winter. Eastern Towhees mainly eat a variety of plant and animal foods, including fruits, berries, and insects. In appropriate habitat, Eastern Towhees may be seen foraging for food on the ground or, less frequently, in the canopy. Birdwatchers may also listen for this species’ song, a trilled “drink-your-teeeeee” or “towhee towheeeee,” from which this species derives its name. Eastern Towhees are primarily active during the day.

Threat Status: Least Concern

  • Eastern Towhee (Pipilo erythrophthalmus). The Internet Bird Collection. Lynx Edicions, n.d. Web. 20 July 2012. .
  • Greenlaw, Jon S. 1996. Eastern Towhee (Pipilo erythrophthalmus), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/262
  • Peterson, Roger Tory. Birds of Eastern and Central North America. Boston: Houghton Mifflin, 1980. Print.
  • Pipilo erythrophthalmus. Xeno-canto. Xeno-canto Foundation, n.d. Web. 20 July 2012. .
  • eBird Range Map - Eastern Towhee. eBird. Cornell Lab of Ornithology, N.d. Web. 20 July 2012. .
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Distribution

National Distribution

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Breeding

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) BREEDING: southern Manitoba and northeastern North Dakota eastward across the northernmost eastern states and southern Ontario and southwestern Quebec to coastal Maine, south to extreme northeastern Texas (at least formerly) and the Gulf coast states and southern Florida, west to western Iowa, southeastern Nebraska, eastern Colorado, eastern Oklahoma, eastern Arkansas, and eastern Louisiana (AOU 1983, Andrews and Righter 1992, Greenlaw 1996). NON-BREEDING: eastern Kansas, Iowa, southern Great Lakes region, southern New York, and southern New England (rarely farther north) south to southern Texas, Gulf coast, and southern Florida (AOU 1983, Greenlaw 1996).

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The eastern towhee occurs throughout the eastern United States. Occurrences from southern Saskatchewan and Quebec south to Florida, and west to eastern Texas are noted in a literature review. Populations north of southern New England through northern Indiana and Illinois to southern Iowa are primarily summer residents [42]. A general range map for the eastern towhee is available at Cornell's All About Birds, and a summer distribution map of the eastern towhee from Breeding Bird Survey data (1966-1996) can be found at Patuxent Wildlife Research Center.

Pipilo e. erythrophthalmus occurs in the most northerly part of the eastern towhee's distribution in the summer, and migrates to the southern and eastern portion of the species' range in the winter. The other subspecies are largely residents [69]. Pipilo e. canaster occurs from south-central Louisiana, north to northeastern Louisiana east through Mississippi, extreme southwestern Tennessee, northern Alabama and Georgia, central South Carolina to western North Carolina, and south to northwestern Florida and east along the Gulf Coast [1]. The range of P. e. rileyi extends from northern Florida through southern Georgia and coastal South Carolina to east-central North Carolina. Pipilo e. alleni occurs in peninsular Florida [69].

The following lists are based on eastern towhee distribution information and the habitat characteristics and plant species composition of vegetation communities eastern towhees are known to occupy. There is not conclusive evidence that eastern towhees occur in all the habitat types listed, and some community types, especially those in the western portion of the eastern towhee's range, may have been omitted. Abundance of eastern towhees in the community types listed is quite variable. Some plant communities support eastern towhees only in stands of specific ages or types, while others provide suitable habitat throughout. See Preferred Habitat/Cover for more detail.

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States or Provinces

(key to state/province abbreviations)
UNITED STATES
AL AR CT DE FL GA IL IN IA KS
KY LA ME MD MA MI MN MS MO NE
NH NJ NY NC ND OH OK PA RI SC
SD TN TX VT VA WV WI DC

CANADA
MB ON PQ SK

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Regional Distribution in the Western United States

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This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

BLM PHYSIOGRAPHIC REGIONS [10]:

14 Great Plains

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Physical Description

Size

Length: 22 cm

Weight: 42 grams

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Ecology

Habitat

Habitat and Ecology

Systems
  • Terrestrial
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Comments: Inhabits forest and swamp edges, regenerating clearcuts, open-canopied forests (including deciduous, pine [PINUS], pine-hardwood and spruce-fir [PICEA-ABIES]; particularly those with a well-developed understory), reclaimed strip mines, mid-late successional fields, riparian thickets, overgrown fencerows, shrub/small-tree thickets, and other brushy habitats. Nest is typically constructed on the ground, concealed among dense, woody vegetation. Later nests (renests, second broods) are more likely to be built above the ground (usually to 1.5 m, but up to 5.5 m) than earlier nests (Palmer-Ball 1996, Greenlaw 1996). However, regional differences in nest placement occur. Even early nests are more likely to be above ground in the Southeast than in the Northeast (J. Greenlaw, pers. comm.). Habitat selection does not differ appreciably between the breeding and non-breeding seasons (Greenlaw 1996). Significantly more abundant in open-canopied deciduous forest having relatively dense understory growth than in closed-canopied forest with relatively sparse understory vegetation (Annand and Thompson 1997, Chadwick et al. 1986, Childers et al. 1986, Crawford et al. 1981, DeGraaf et al. 1991, Robinson and Robinson 1999, Yahner 1993).

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Habitat: Cover Types

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This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

More info for the terms: cover, swamp

SAF COVER TYPES [31]:

12 Black spruce

13 Black spruce-tamarack

14 Northern pin oak

16 Aspen

17 Pin cherry

18 Paper birch

19 Gray birch-red maple

20 White pine-northern red oak-red maple

21 Eastern white pine

22 White pine-hemlock

23 Eastern hemlock

24 Hemlock-yellow birch

25 Sugar maple-beech-yellow birch

26 Sugar maple-basswood

27 Sugar maple

28 Black cherry-maple

30 Red spruce-yellow birch

31 Red spruce-sugar maple-beech

32 Red spruce

33 Red spruce-balsam fir

34 Red spruce-Fraser fir

38 Tamarack

39 Black ash-American elm-red maple

40 Post oak-blackjack oak

42 Bur oak

43 Bear oak

44 Chestnut oak

45 Pitch pine

46 Eastern redcedar

50 Black locust

51 White pine-chestnut oak

52 White oak-black oak-northern red oak

53 White oak

55 Northern red oak

57 Yellow-poplar

58 Yellow-poplar-eastern hemlock

59 Yellow-poplar-white oak-northern red oak

60 Beech-sugar maple

61 River birch-sycamore

62 Silver maple-American elm

63 Cottonwood

64 Sassafras-persimmon

65 Pin oak-sweetgum

69 Sand pine

70 Longleaf pine

71 Longleaf pine-scrub oak

72 Southern scrub oak

73 Southern redcedar

74 Cabbage palmetto

75 Shortleaf pine

76 Shortleaf pine-oak

78 Virginia pine-oak

79 Virginia pine

80 Loblolly pine-shortleaf pine

81 Loblolly pine

82 Loblolly pine-hardwood

83 Longleaf pine-slash pine

84 Slash pine

85 Slash pine-hardwood

87 Sweetgum-yellow-poplar

88 Willow oak-water oak-diamondleaf (laurel) oak

91 Swamp chestnut oak-cherrybark oak

92 Sweetgum-willow oak

93 Sugarberry-American elm-green ash

94 Sycamore-sweetgum-American elm

95 Black willow

96 Overcup oak-water hickory

98 Pond pine

100 Pondcypress

101 Baldcypress

102 Baldcypress-tupelo

103 Water tupelo-swamp tupelo

104 Sweetbay-swamp tupelo-redbay

105 Tropical hardwoods

106 Mangrove

108 Red maple

109 Hawthorn

110 Black oak

111 South Florida slash pine

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Habitat: Plant Associations

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This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the term: bog

KUCHLER [55] PLANT ASSOCIATIONS:

K074 Bluestem prairie

K076 Blackland prairie

K077 Bluestem-sacahuista prairie

K078 Southern cordgrass prairie

K079 Palmetto prairie

K080 Marl everglades

K081 Oak savanna

K082 Mosaic of K074 and K100

K083 Cedar glades

K084 Cross Timbers

K089 Black Belt

K091 Cypress savanna

K092 Everglades

K093 Great Lakes spruce-fir forest

K094 Conifer bog

K095 Great Lakes pine forest

K096 Northeastern spruce-fir forest

K097 Southeastern spruce-fir forest

K098 Northern floodplain forest

K099 Maple-basswood forest

K100 Oak-hickory forest

K101 Elm-ash forest

K102 Beech-maple forest

K103 Mixed mesophytic forest

K104 Appalachian oak forest

K105 Mangrove

K106 Northern hardwoods

K107 Northern hardwoods-fir forest

K108 Northern hardwoods-spruce forest

K109 Transition between K104 and K106

K110 Northeastern oak-pine forest

K111 Oak-hickory-pine

K112 Southern mixed forest

K113 Southern floodplain forest

K114 Pocosin

K115 Sand pine scrub

K116 Subtropical pine forest

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Habitat: Ecosystem

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This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

ECOSYSTEMS [36]:

FRES10 White-red-jack pine

FRES11 Spruce-fir

FRES12 Longleaf-slash pine

FRES13 Loblolly-shortleaf pine

FRES14 Oak-pine

FRES15 Oak-hickory

FRES16 Oak-gum-cypress

FRES17 Elm-ash-cottonwood

FRES18 Maple-beech-birch

FRES19 Aspen-birch

FRES39 Prairie

FRES41 Wet grasslands

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Migration

Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.

Locally Migrant: Yes. At least some populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: Yes. At least some populations of this species make annual migrations of over 200 km.

Most individuals breeding in the northern portion of the range migrate southward in the fall. However, some individuals may overwinter when sufficient natural food is available or supplemental food (e.g., feeding station) is provided, particularly during mild winters. Southeastern and Gulf state populations are generally sedentary or partially migratory. Migratory birds arrive in northern states from late April through early May, and depart from mid-September through mid-October. Migrations dates vary considerably, depending on weather. Birds that breed in northeastern states migrate between the Appalachian Mountains and the Atlantic Coast, whereas those that breed in the Midwest migrate west of the Appalachians. Migrates at night. Older males generally arrive on northern breeding grounds ahead of females and younger males (Greenlaw 1996).

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Trophic Strategy

Comments: Typically forages on the ground in dense, low vegetative cover. Sometimes forages in the open (usually near dense cover) or arboreally, especially in late spring and summer. Scratches among loose ground debris (e.g., leaf litter) to uncover seeds and invertebrates. Omnivorous; consumes a wide variety of seeds, fruits and invertebrates. Diet sometimes includes foliage buds, young grass stems, and flower buds (Greenlaw 1996). Also consumes vertebrates such as small salamanders, snakes, and lizards (Terres 1991). The relative proportion of animal/plant material in the diet varies seasonally. Based on the examination of 648 specimens collected throughout the year, plant material comprises 85% of the diet in winter, 53% in spring, 43% in summer and 79% in fall (Martin et al. 1951).

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Food Habits

More info for the terms: frequency, tree

Eastern towhees primarily eat on the ground, although they also glean from vegetation. In a southeastern Massachusetts pitch pine barren, 73.5% of male and 80.4% of female foraging observations were on the ground [66]. When foraging on the ground eastern towhees use a scratching technique where both feet kick back simultaneously [40,86]. In a laboratory study 4 eastern towhees used this method to successfully obtain seed buried almost 1 inch (2.25 cm) deep [98]. When foraging above ground the majority of time is spent gleaning foliage [66]. In Massachusetts, 22.5% of male and 16.3% of female foraging observations were of food being gleaned from foliage. Eastern towhees were also observed gleaning from twigs, branches, and trunks. When gleaning, eastern towhees occurred significantly (p<0.01) more often on the distal half of tree branches compared to using distal and proximal portions equally (see Foraging Habitat). In 0.5% of male and 0.3% of female foraging observations, eastern towhees hovered. Eastern towhees were never observed catching food out of the air [66].

Eastern towhees eat a variety of plant and animal matter. In literature reviews, eastern towhees are reported to eat seeds and fruits, several invertebrates, and occasionally small amphibians, snakes, and lizards [86]. Reviews report eastern towhees foraging at feeders [17,42]. Reviews show that animal matter makes up a larger proportion of the diet in the breeding season [42,63]. In fall and winter, plants make up 79% and 85% of the diet, respectively. This drops to 53% in spring and 43% in summer [63]. Insects such as beetles (Coleoptera), grasshoppers and crickets (Orthoptera), ants, wasps, and bees (Hymenoptera), and moths and caterpillars (Lepidoptera) are common prey items. Eastern towhees eat other invertebrates such as spiders (Araneae), millipedes (Diplopoda), centipedes (Chilopoda), and snails (Gastropoda) to a lesser extent [42,63]. Plants that comprise at least 5% of the eastern towhee diet include ragweed (Ambrosia spp.), oak, smartweed (Polygonum spp.), and corn (Zea mays) in the Northeast and blackberry, oak, panicgrass (Panicum spp.), ragweed, and wax-myrtle (Morella cerifera) in the Southeast [63]. The following table, adapted from Greenlaw's [42] literature review, shows the relative total volume (%) and frequency of occurrence (%, in parentheses) of different food items found in the stomach contents of adult eastern towhees.

Northeastern Region Midwestern Region Southeast Atlantic/Mid-Gulf Region
May-June Sept-Oct May-June Sept-Oct May-June Sept-Oct
Insects (adult) 29.6 (93) 0.6 (40) 49.2 (98) 35.1 (91) 32.8 (95) 2.5 (73)
Insects (immature) 5.4 (24) 0 4.7 (25) 2.4 (13) 13.0 (46) 0
Spiders 0.3 (6) 0.8 (20) 1.3 (17) 0.5 (13) 1.7 (18) 0
Millipedes 0.5 (9) 0.6 (20) 2.8 (15) 0.3 (9) 0.1 (2) 0
Centipedes 0 0.4 (20) 0.1 (2) 0 0 0
Snails 0.3 (3) 0 0.1 (6) 0 0.1 (2) 0
Plant 63.8 (91) 97.6 (100) 42.0 (67) 61.4 (83) 49.8 (79) 97.5 (100)

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Associations

Predators

Many animals prey on eastern towhees and their eggs, including reptiles, mammals, and birds. A literature review summarizes several reports demonstrating that predators are a major cause of nest failure [42]. The highest nest predation rate noted was 88% in a New York study. Mammals that are likely nest predators include northern raccoons (Procyon lotor), domestic cats (Felis catus), and eastern chipmunks (Tamias striatus). Snakes such as bullsnakes (Pituophis catenifer), rat snakes (Elaphe spp.) and garter snakes (Thamnophis spp.) have been reported eating eastern towhee eggs. Weasels (Mustela spp.) and blue jays (Cyanocitta cristata) are also likely nest predators [42]. Several birds are known to prey on adult eastern towhees, including short-tailed (Buteo brachyurus), sharp-shinned (Accipiter striatus), and Cooper's hawks (Accipiter cooperii) [70,87]. At least some mammals also feed on adult eastern towhees. In Maryland, an eastern towhee was found in the stomach contents of a red fox (Vulpes vulpes) [47].

Brown-headed cowbirds (Molothrus ater) parasitize eastern towhee nests. In a South Carolina old field 5 of 19 eastern towhee nests were parasitized [100]. Each parasitized nest contained 1 brown-headed cowbird egg. The desertion rate for parasitized nests was 20%, which was similar to nests that had not been parasitized (21%). Two of the five brown-headed cowbird eggs produced fledglings. The study did not determine if there was a difference in nest success between parasitized and nonparasitized nests [100]. In West Virginia, only 3 of 41 eastern towhee nests were parasitized by the brown-headed cowbird. Average number of fledged young in nonparasitized nests was 2.8, which was similar to the average of 2.7 fledglings per parasitized nest [9]. In a Pennsylvania study site, only 2 of 36 nests were parasitized and both produced eastern towhee fledglings [26]. In a study of nest parasitism on Sanibel Island, none of 5 eastern towhee nests found were parasitized [74].

BEHAVIORS:
Greenlaw's [42] literature review provides a detailed summary on eastern towhee behaviors, such as vocalizations, courtship, territoriality (see Timing of Major Life History Events), and migration (see General Distribution).

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General Ecology

DENSITY: In Massachusetts, density of breeding birds varied from an average of 1.78 individuals/ha in closed-canopy forest stands to an average of 7.41 individuals/ha in selectively-harvested stands (Chadwick et al. 1986). In Pennsylvania, the mean number of breeding individuals ranged from 0.17-0.99/ha in six forest types (Yahner 1986). In Arkansas, the number of territorial males ranged from 0.025-0.23/ha. Lowest densities occurred in a burned field and a dry forest, whereas the highest density occurred along a forest edge (James and Neal 1986). In the Pine barrens of southeastern Massachusetts, breeding bird density ranged from about 1.5-3/ha during a three-year period (Morimoto and Wasserman 1991). In New Jersey, population density varied from 0.5 males/ha in mesic oak forest to 0.8 males/ha in xeric pine-oak woodland (Greenlaw 1996). Territory size can vary inversely with experimentally altered food supplies. However, territories appear to contain more than the minimal food stocks necessary for maintenance and reproduction (Wasserman 1983).

INTRA- AND INTERSPECIFIC AGGREGATIONS: Forms loose flocks during the winter, ranging in size from 8-27 (mean = 17) individuals. These wintering groups often form mixed-species foraging flocks with Dark-eyed Juncos (JUNCO HYEMALIS), White-throated Sparrows (ZONOTRICHIA ALBICOLLIS), Field Sparrows (SPIZELLA PUSILLA), Song Sparrows (MELOSPIZA MELODIA), Northern Cardinals (CARDINALIS CARDINALIS), Tufted Titmice (BAEOLOPHUS BICOLOR), Winter Wrens (TROGLODYTES TROGLODYTES) and Carolina Chickadees (POECILE CAROLINENSIS; Barbour 1941).

SITE FIDELITY: Exhibits breeding site fidelity. In New Jersey, one male returned to the same 24-hectare section of oak forest in 5 of 6 years (Greenlaw 1996). In another New Jersey study, the return rate was 20% the first year after banding and 43% thereafter (Leck et al. 1988).

POPULATION PARAMETERS: Sex ratio data are limited; however, sex ratios determined to date are male-biased (at least temporarily) or no different from parity (Greenlaw 1996). Estimated annual probability of survival among adults is 0.58 (Savidge and Davis 1974 cited in Greenlaw 1996). Oldest known individual was at least 12 years, 3 months old (Terres 1991).

PARASITES: Host to at least 14 species of ectoparasites and six internal parasites (Greenlaw 1996).

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Habitat-related Fire Effects

More info for the terms: cover, density, fire exclusion, fire interval, fire severity, frequency, litter, low-severity fire, severity, shrubs, stand-replacing fire, succession, wildfire

Despite the abundance of articles addressing the effect of fire on eastern towhees, results should be interpreted with caution. As noted by a literature review summarizing songbird responses to fire in southwestern ponderosa pine (Pinus ponderosa) forests [32], there are several limitations to many studies addressing bird response to fire. Many studies focus on breeding communities and use composite statistics, such as species richness. Many are opportunistic, restricted in spatial or temporal scale, include confounding factors and/or lack sufficient replication. Very few compare demographic parameters of a given species between burned and unburned vegetation, which is necessary to determine if the site is meeting the needs of a species [32].

Given that eastern towhees are strongly associated with low woody cover (see Preferred Habitat/Cover), their response is likely influenced by the interaction of habitat type, fire interval, fire uniformity, and fire severity. In habitats such as loblolly-shortleaf pineland and slash pine flatwoods, fairly frequent low-severity fires appear to benefit eastern towhees [14,30,99]. However in habitats such as tallgrass prairie, frequent fires eliminate eastern towhee from the habitat [106], likely due to a decline in shrubby cover. Patchy fires may benefit eastern towhees, due to increased edge (see Effects of spatial arrangement/area) and decreased damage to existing understory vegetation. Due to their cover requirements (see Preferred Habitat/Cover), eastern towhees may be affected by fires severe enough to seriously impact understory vegetation [94]. In cover types where eastern towhees occur primarily in early successional habitat, large stand-replacing fires may create patches that would provide eastern towhee habitat several years after the burn. Eastern towhees have been observed on a sand pine (Pinus clausa) scrub site in Florida 3 to 7 years after it was burned in a severe wildfire and subsequently salvage logged [38].

Eastern towhee's response to fire depends in part on the vegetation community. In some habitat types eastern towhees are present soon after burning, while in other habitat types eastern towhees do not occur on recently burned areas. For example, the average number of eastern towhee detections per census was approximately 5 on an old field pineland in Florida that had been winter-burned annually for approximately 30 years [30]. In Arkansas, eastern towhees occurred at a density of 6 territorial males/40 ha the spring following a winter burn in a field that had been dominated by broomsedge bluestem (Andropogon virginicus) and sassafras (Sassafras albidum) saplings [81]. Eastern towhees were also observed on wetlands burned 6 months earlier [95]. However, in an Indiana oak forest that had been surface burned 1 to 2 years and 3 to 4 years before sampling, eastern towhees were absent despite their occurrence in unburned control plots [5]. Eastern towhees did not occur in annually burned tallgrass prairies, but were observed in unburned (>10 years) tallgrass prairie in some years [106]. Eastern towhees did not occur in frequently burned longleaf pine sandhill plots [75]. In areas of Minnesota where frequent (12-26 fires in 32 years) fires were being used to restore oak savanna once dominated by bur oak (Quercus macrocarpa) and northern pin oak (Q. ellipsoidalis), eastern towhees were associated with unburned vegetation [21].

In some habitats eastern towhees increase after prescribed surface fires. Shortly after fire was excluded from an old-field loblolly-shortleaf (P. echinata) pineland that had previously been burned annually, eastern towhee occurrence increased. Eastern towhee detections per census increased from approximately 5 in postfire year 1 to over 12 in the 4th postfire year. After 4 years eastern towhee detections gradually declined. After 11 years of fire exclusion, eastern towhee detections were less than in the 1st postfire year. Due to eastern towhee absence in a beech-magnolia (Fagus-Magnolia spp.) stand, the authors predicted that as succession continues and the proportion of deciduous canopy increases, eastern towhees will eventually leave the site [30]. In coastal scrub and slash pine flatwoods in east-central Florida, eastern towhee densities were significantly (p=0.007) higher (4.5/ha) on sites that had burned 4 years before the survey than those that had been burned 1 year before (2.3/ha), 2 years before (3.3/ha), or more than 10 years before (2.6/ha) the survey [14]. The habitat with the greatest number of eastern towhees detected on Eglin Air Force Base in Florida was a "mature flatwoods" that had been burned 1 to 5 years before the survey. Eastern towhee detections in this habitat were significantly (p<0.05) greater than in both the burned (1-3 years previously) and unburned (for several decades) sandhills dominated by mature (>50 yr) longleaf pine [92]. In a loblolly pine forest with a mid-story dominated by oaks (Quercus spp.) and hickories (Carya spp.), eastern towhee abundance was significantly (p<0.05) greater on sites that had been burned either 1 year, 2 years, or 3 years before sampling (detections/count/site=3.79) compared to sites that had not been burned for more than 20 years (detections/count/site=1.44). Eastern towhees were also observed at higher frequencies in burned areas (89%) compared to unburned areas (74%) [99]. Three areas in Pennsylvania, one unburned area, one that had been burned 1 year earlier, and one that had been burned 2 years earlier were comprised of various habitats including a bluestem (Andropogon spp.) community, a scrub oak (bear oak and dwarf oak (Q. prinoides) community, a scrub oak-mixed aspen community, a scrub oak-pitch pine community, and a scrub oak-mixed-oak community. Eastern towhees were the most common bird in all three areas, but occurred at the highest density, approximately 0.67 singing male/ha, in the 2-year-old burn. In the unburned it occurred at a density of about 0.43 singing male/ha, and in the 1-year-old burn singing male eastern towhees occurred at a density of about 0.38/ha [83].

Differences in vegetation structure after burning may explain at least some of the variation in eastern towhee's response. A literature review demonstrated that changes in bird communities after fire are associated with the degree of structural change in the vegetation. More structurally complex habitats and more severe fires result in longer periods before the prefire bird community returns. For instance, the prefire bird community is typically restored about 3 years after a grassland fire, while it can take over 30 years for the prefire bird community to return after a stand-replacing forest fire [48]. Data on vegetation in areas where eastern towhee did and did not occur after fire show the effects of much smaller changes in vegetation structure. Vegetation on burned and unburned areas differed in an investigation that found significantly (p<0.05) higher eastern towhee abundance in burned areas. The unburned stands of loblolly pine with an understory of oaks and hickories had significantly (p<0.05) more hardwoods and total trees (≥7.5 cm dbh/ha), more snags and logs (≥7.5 cm/ha), higher percentage canopy closure, higher percentage of leaf litter, and lower percentage of herb cover than burned plots [99]. In contrast, an Indiana study area comprised of oak (Quercus spp.) dominated forests, chestnut oak (Q. prinus) woodlands, oak-hickory forests, and American beech (Fagus grandifolia)-sugar maple forests that were burned twice in 4 years had lower density of small 1 inch (<2.5 cm) dbh live woody stems and less horizontal vegetation cover below 1.6 feet (0.5 m) than the unburned site. In this study, eastern towhees were observed in the unburned vegetation, but not in the burned area [4,5].

Severity of the fire is also likely have a major impact on eastern towhee's response. In a literature review, it is suggested that fires of any severity are likely to affect ground-nesting birds, since even low-severity fire is likely to have large impacts on the vegetation they use [61]. Rotenberry and others [78] hypothesize in a literature review that "cool" and "intermediate" burns could increase food availability and leave adequate vegetation for ground-nesting birds. Data comparing eastern towhee densities among burns of varying severity are lacking. Observations of a pitch pine barren site in New Jersey shortly after a fairly extensive wildfire occurred in late winter/early spring did provide some information. The number of eastern towhees declined in some burned areas and increased in areas adjacent to the burn area. However, in areas within the burn where the vegetation was only slightly damaged and new growth provided cover, eastern towhee abundance was not affected [94]. In South Carolina, average weekly adult eastern towhee survival rates were not significantly (p=0.15) different in young (2-5 years old) longleaf pine stands than in mature (32-97 years old) longleaf pine stands that were occasionally thinned and surface burned under prescription every 3 to 5 years [52]. Eastern towhees were observed on a sand pine scrub site in Florida subject to a stand-replacing fire and salvage logging between 5 and 7 years before sampling. The following table shows the average density (number/km²) of eastern towhees on the burned and salvaged site and on a mature (≥55 years since stand-replacing fire) forest site [38].

Burned and salvaged Mature
Spring 286.1 147.2
Winter 197.2 111.1

Seasonal effect of burning on eastern towhee is uncertain. Literature reviews suggest that burns during the breeding season are likely to have a larger impact on ground-nesting birds, both directly through nest mortality and indirectly through greater and/or more persistent loss of cover than would likely occur from dormant season burns [61,77]. In addition, once territories are established eastern towhees may be less likely to leave a burned area [29]. No studies to date (2006) have compared eastern towhee demographics on sites burned in varying seasons. However, comparisons of eastern towhee abundance on sites burned during dormant and growing seasons have not resulted in detectable effects. On sites burned every 2 to 4 years in pure and mixed stands of longleaf, loblolly, and shortleaf pine in Georgia, eastern towhee abundance on sites burned between April and August (average of 1.54 birds/plot) was not significantly (p=0.32) different from the sites burned between January and March (average of 2.31 birds/plot) [50]. The following table shows the frequency (%) and abundance (total/40-m radius) of eastern towhees on control plots, plots burned in January, and plots burned in June in a dry prairie/shrubland in southwestern Florida [33]:

  Control Winter burn Summer burn
Frequency 44 44 64
Abundance 21 20 25

Fire is likely to have an affect on the availability of eastern towhee food items. Several literature reviews summarize information regarding changes in plant and animal food availability after fire [23,60,78]. Several shrubby species may increase fruit production after fire, although the response is dependent on the species and postfire environmental factors [23]. Seed and fruit production typically increase after fire in southern forests, with peak production from 2 to 6 years after fire. The frequency and season of burning influence plant recovery and fruit production [60]. Lyon and others' [60] review discusses the effect of fire on invertebrates, some of which are eastern towhee prey. Many of the studies found an increase in arthropod availability after understory fires in southern sandhill and loblolly-shortleaf pine forests [60].

Fire ecology: Eastern towhees occur in a variety of habitats with a wide range of FIRE REGIMES. However, areas with the highest towhee densities typically undergo low-severity surface fires at an interval of about 4 to 10 years [14,30]. Fires in the southeastern United States frequently occur in summer due to an increase in the occurrence of lightning strikes [73].

Due to their preference for early successional stands (see Preferred Habitat/Cover), eastern towhees are likely to occur in early to intermediate successional stages after stand-replacing fires.

In herbaceous habitats eastern towhees are most likely to occur in areas where fire exclusion [106] has resulted in establishment of some woody shrubs.

The following table provides fire return intervals for plant communities and ecosystems where eastern towhee is likely to occur. For further information, see the FEIS review of the dominant species listed below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
maple-beech-birch Acer-Fagus-Betula spp. >1,000
silver maple-American elm Acer saccharinum-Ulmus americana <5 to 200
sugar maple Acer saccharum >1,000
sugar maple-basswood Acer saccharum-Tilia americana >1,000 [96]
bluestem prairie Andropogon gerardii var. gerardii-Schizachyrium scoparium 54,71]
bluestem-Sacahuista prairie Andropogon littoralis-Spartina spartinae 71]
mangrove Avicennia nitida-Rhizophora mangle 35-200 [68]
sugarberry-America elm-green ash Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica <35 to 200
beech-sugar maple Fagus spp.-Acer saccharum >1,000 [96]
cedar glades Juniperus virginiana 3-22 [43,71]
tamarack Larix laricina 35-200 [71]
yellow-poplar Liriodendron tulipifera <35 [96]
Everglades Mariscus jamaicensis <10 [68]
Great Lakes spruce-fir Picea-Abies spp. 35 to >200
northeastern spruce-fir Picea-Abies spp. 35-200 [27]
southeastern spruce-fir Picea-Abies spp. 35 to >200 [96]
black spruce Picea mariana 35-200
conifer bog* Picea mariana-Larix laricina 35-200
red spruce* Picea rubens 35-200 [27]
pine-cypress forest Pinus-Cupressus spp. <35 to 200 [6]
shortleaf pine Pinus echinata 2-15
shortleaf pine-oak Pinus echinata-Quercus spp. <10
slash pine Pinus elliottii 3-8
slash pine-hardwood Pinus elliottii-variable <35
sand pine Pinus elliottii var. elliottii 25-45 [96]
South Florida slash pine Pinus elliottii var. densa 1-5
longleaf-slash pine Pinus palustris-P. elliottii 1-4 [68,96]
longleaf pine-scrub oak Pinus palustris-Quercus spp. 6-10 [96]
red-white-jack pine* Pinus resinosa-P. strobus-P. banksiana 10-300 [27,45]
pitch pine Pinus rigida 6-25 [16,46]
pocosin Pinus serotina 3-8
pond pine Pinus serotina 3-8
eastern white pine Pinus strobus 35-200
eastern white pine-eastern hemlock Pinus strobus-Tsuga canadensis 35-200
eastern white pine-northern red oak-red maple Pinus strobus-Quercus rubra-Acer rubrum 35-200
loblolly pine Pinus taeda 3-8
loblolly-shortleaf pine Pinus taeda-P. echinata 10 to <35
Virginia pine Pinus virginiana 10 to <35
Virginia pine-oak Pinus virginiana-Quercus spp. 10 to <35
sycamore-sweetgum-American elm Platanus occidentalis-Liquidambar styraciflua-Ulmus americana <35 to 200 [96]
eastern cottonwood Populus deltoides <35 to 200 [71]
aspen-birch Populus tremuloides-Betula papyrifera 35-200 [27,96]
black cherry-sugar maple Prunus serotina-Acer saccharum >1,000
oak-hickory Quercus-Carya spp. <35
northeastern oak-pine Quercus-Pinus spp. 10 to <35 [96]
oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to >200 [68]
southeastern oak-pine Quercus-Pinus spp. <10
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra <35
northern pin oak Quercus ellipsoidalis <35
bear oak Quercus ilicifolia <35
bur oak Quercus macrocarpa <10 [96]
oak savanna Quercus macrocarpa/Andropogon gerardii-Schizachyrium scoparium 2-14 [71,96]
chestnut oak Quercus prinus 3-8
northern red oak Quercus rubra 10 to <35
post oak-blackjack oak Quercus stellata-Q. marilandica <10
black oak Quercus velutina <35 [96]
cabbage palmetto-slash pine Sabal palmetto-Pinus elliottii <10 [68,96]
blackland prairie Schizachyrium scoparium-Nassella leucotricha <10 [96]
southern cordgrass prairie Spartina alterniflora 1-3 [71]
baldcypress Taxodium distichum var. distichum 100 to >300
pondcypress Taxodium distichum var. nutans <35 [68]
eastern hemlock-yellow birch Tsuga canadensis-Betula alleghaniensis >200 [96]
elm-ash-cottonwood Ulmus-Fraxinus-Populus spp. <35 to 200 [27,96]
*fire return interval varies widely; trends in variation are noted in the species review

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Direct Effects of Fire

More info for the terms: fire interval, fire severity, severity

Adult eastern towhees are unlikely to suffer directly from fire. It is generally accepted that large, fast-moving fires can result in mortality, but birds typically have the mobility to avoid fire [19,23,76]. Spotted towhees (Pipilo maculatus), birds that are closely related and have similar habits as eastern towhees, were observed alive after a fire in California coastal sage scrub [12].

It is likely that eggs and young birds are much more vulnerable to fire. Although there were no data directly investigating eastern towhee nest mortality due to fire as of early 2006, literature reviews have used life history characteristics to speculate on possible effects of fire on nesting success and bird populations [62,77]. Since eastern towhees nest near the ground, low-severity surface fires during the breeding season could result in considerable nest mortality. However, the degree to which a population would be affected by fire would depend on several factors including occurrence of renesting, season of burn, fire interval, fire uniformity, and fire severity. Since eastern towhees have the tendency to renest, this may mitigate at least some of the impact a breeding-season fire would have [62,77]. Over 2 breeding seasons, 7 of 33 eastern towhee nests observed in a mature loblolly pine forest that had burned between 1 to 2 and 3 to 4 years earlier produced at least 1 fledgling [99]. The daily nest survival rate was 89%, and a mean of 0.68 fledgling was produced per active nest. These values could not be compared to the unburned site due to the lack of nests found there [99]. Higher [93] and lower [52] daily nest survival rates have been reported for eastern towhees in other areas (see Timing of Major Life History Events).

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Life History and Behavior

Life Expectancy

Lifespan, longevity, and ageing

Maximum longevity: 12.3 years (wild)
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Reproduction

Both sexes become sexually mature in second year of life (Greenlaw 1996).

PHENOLOGY: Breeding season varies with latitude. In the southeastern U.S. (Florida and Georgia) nest building and oviposition begin in late March-early April, whereas in New England these activities are delayed until mid-May (Greenlaw 1996, Stevenson and Anderson 1994). In the Midwest, breeding activities begin in mid- to late April (Mumford and Keller 1984).

OVIPOSITION/INCUBATION: Dates of oviposition can vary among habitats at the same latitude. In New Jersey, for example, oviposition began earlier in mesic oak forest than in xeric pine barrens. This difference was attributed to earlier foliage development and the concomitant earlier appearance of additional invertebrates in the oak forest (Greenlaw 1978). Egg-laying generally terminates by mid- to late July, but can extend into mid to late August. Clutch size is 2-6 (usually 3-4 eggs), and incubation begins with the laying of the penultimate or ultimate egg (Greenlaw 1996). Clutch size can vary at the same latitude, with larger clutches being produced in habitats with greater food supplies (Greenlaw 1978). Incubation, by the female only, lasts 12-13 days. Both sexes feed the nestlings and fledglings.

FLEDGING: Young leave the nest when 10-11 days old. Produce 1-2 broods per season; will also renest after nest failure (Greenlaw 1996).

NEST SUCCESS: At two sites in New Jersey, egg success (number of eggs that produced fledged young) varied from 28.3-35.6%. Nest predation rates ranged from 31-73% at these two sites (Greenlaw 1996). In West Virginia, nest success (nests that fledged at least one young) was 36.5% (Bell and Whitmore 1997).

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Pipilo erythrophthalmus

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 3 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

CCTGTACCTAATTTTCGGCGCATGAGCCGGAATAGTAGGTACCGCCCTAAGCCTCCTCATTCGGGCCGAACTAGGCCAACCTGGAGCTCTCCTAGGAGACGACCAAATTTATAACGTAGTCGTCACAGCCCACGCTTTCGTGATAATTTTCTTCATAGTTATACCAATTATGATCGGAGGCTTCGGAAACTGACTAGTCCCCCTAATAATTGGAGCCCCAGACATAGCATTCCCACGAATAAATAACATAAGTTTCTGACTACTACCCCCATCTTTCCTCCTCCTCCTAGCATCCTCCACCGTTGAAGCAGGTGCTGGCACAGGCTGAACAGTGTACCCACCACTAGCCGGCAACTTAGCCCACGCTGGAGCTTCAGTCGACCTCGCAATTTTCTCCCTACACCTAGCCGGTATCTCCTCAATCTTAGGAGCAATCAACTTCATCACAACAGCAATTAACATAAAACCCCCTGCCCTCTCACAATACCAGACCCCCCTATTTGTATGATCAGTCCTAATCACCGCAGTCTTACTACTCCTATCACTCCCAGTCCTCGCCGCAGGAATCACAATGCTCCTTACAGACCGCAACCTAAACACTACATTCTTTGACCCCGCTGGAGGAGGGGACCCTGTCCTATACCAACATCTCNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNN
-- end --

Download FASTA File

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© Barcode of Life Data Systems

Source: Barcode of Life Data Systems (BOLD)

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Statistics of barcoding coverage: Pipilo erythrophthalmus

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 3
Specimens with Barcodes: 5
Species With Barcodes: 1
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© Barcode of Life Data Systems

Source: Barcode of Life Data Systems (BOLD)

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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2012

Assessor/s
BirdLife International

Reviewer/s
Butchart, S. & Symes, A.

Contributor/s

Justification
This species has an extremely large range, and hence does not approach the thresholds for Vulnerable under the range size criterion (Extent of Occurrence <20,000 km2 combined with a declining or fluctuating range size, habitat extent/quality, or population size and a small number of locations or severe fragmentation). The population trend appears to be stable, and hence the species does not approach the thresholds for Vulnerable under the population trend criterion (>30% decline over ten years or three generations). The population size is extremely large, and hence does not approach the thresholds for Vulnerable under the population size criterion (<10,000 mature individuals with a continuing decline estimated to be >10% in ten years or three generations, or with a specified population structure). For these reasons the species is evaluated as Least Concern.

History
  • Least Concern (LC)
  • Least Concern (LC)
  • Least Concern (LC)
  • Lower Risk/least concern (LR/lc)
  • Not Recognized (NR)
  • Not Recognized (NR)
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© International Union for Conservation of Nature and Natural Resources

Source: IUCN

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