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.
Regularity: Regularly occurring
Type of Residency: Breeding
Regularity: Regularly occurring
Type of Residency: Year-round
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).
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 . 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 . 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 . 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 .
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.
States or Provinces
Regional Distribution in the Western United States
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 :
14 Great Plains
Length: 22 cm
Weight: 42 grams
Habitat and Ecology
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).
Habitat: Cover Types
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 :
12 Black spruce
13 Black spruce-tamarack
14 Northern pin oak
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
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
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
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
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
103 Water tupelo-swamp tupelo
104 Sweetbay-swamp tupelo-redbay
105 Tropical hardwoods
108 Red maple
110 Black oak
111 South Florida slash pine
Habitat: Plant Associations
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  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
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
K106 Northern hardwoods
K107 Northern hardwoods-fir forest
K108 Northern hardwoods-spruce forest
K109 Transition between K104 and K106
K110 Northeastern oak-pine forest
K112 Southern mixed forest
K113 Southern floodplain forest
K115 Sand pine scrub
K116 Subtropical pine forest
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):
FRES10 White-red-jack pine
FRES12 Longleaf-slash pine
FRES13 Loblolly-shortleaf pine
FRES41 Wet grasslands
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).
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).
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 . 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 . When foraging above ground the majority of time is spent gleaning foliage . 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 .
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 . 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 . 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 . The following table, adapted from Greenlaw's  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|
|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)|
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 . 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 . 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) .
Brown-headed cowbirds (Molothrus ater) parasitize eastern towhee nests. In a South Carolina old field 5 of 19 eastern towhee nests were parasitized . 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 . 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 . In a Pennsylvania study site, only 2 of 36 nests were parasitized and both produced eastern towhee fledglings . In a study of nest parasitism on Sanibel Island, none of 5 eastern towhee nests found were parasitized .BEHAVIORS:
Greenlaw's  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).
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).
Habitat-related Fire Effects
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 , 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 .
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 , 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 . 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 .
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 . 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 . Eastern towhees were also observed on wetlands burned 6 months earlier . 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 . Eastern towhees did not occur in annually burned tallgrass prairies, but were observed in unburned (>10 years) tallgrass prairie in some years . Eastern towhees did not occur in frequently burned longleaf pine sandhill plots . 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 .
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 . 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 . 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 . 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%) . 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 .
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 . 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 . 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 . Rotenberry and others  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 . 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 . 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/kmÂ²) of eastern towhees on the burned and salvaged site and on a mature (≥55 years since stand-replacing fire) forest site .
|Burned and salvaged||Mature|
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 . 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) . 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 :
|Control||Winter burn||Summer burn|
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 . 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 . Lyon and others'  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 .
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 .
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  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)|
|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 |
|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 |
|sugarberry-America elm-green ash||Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica||<35 to 200|
|beech-sugar maple||Fagus spp.-Acer saccharum||>1,000 |
|cedar glades||Juniperus virginiana||3-22 [43,71]|
|tamarack||Larix laricina||35-200 |
|yellow-poplar||Liriodendron tulipifera||<35 |
|Everglades||Mariscus jamaicensis||<10 |
|Great Lakes spruce-fir||Picea-Abies spp.||35 to >200|
|northeastern spruce-fir||Picea-Abies spp.||35-200 |
|southeastern spruce-fir||Picea-Abies spp.||35 to >200 |
|black spruce||Picea mariana||35-200|
|conifer bog*||Picea mariana-Larix laricina||35-200|
|red spruce*||Picea rubens||35-200 |
|pine-cypress forest||Pinus-Cupressus spp.||<35 to 200 |
|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 |
|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 |
|red-white-jack pine*||Pinus resinosa-P. strobus-P. banksiana||10-300 [27,45]|
|pitch pine||Pinus rigida||6-25 [16,46]|
|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 |
|eastern cottonwood||Populus deltoides||<35 to 200 |
|aspen-birch||Populus tremuloides-Betula papyrifera||35-200 [27,96]|
|black cherry-sugar maple||Prunus serotina-Acer saccharum||>1,000|
|northeastern oak-pine||Quercus-Pinus spp.||10 to <35 |
|oak-gum-cypress||Quercus-Nyssa-spp.-Taxodium distichum||35 to >200 |
|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 |
|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 |
|cabbage palmetto-slash pine||Sabal palmetto-Pinus elliottii||<10 [68,96]|
|blackland prairie||Schizachyrium scoparium-Nassella leucotricha||<10 |
|southern cordgrass prairie||Spartina alterniflora||1-3 |
|baldcypress||Taxodium distichum var. distichum||100 to >300|
|pondcypress||Taxodium distichum var. nutans||<35 |
|eastern hemlock-yellow birch||Tsuga canadensis-Betula alleghaniensis||>200 |
|elm-ash-cottonwood||Ulmus-Fraxinus-Populus spp.||<35 to 200 [27,96]|
Direct Effects of Fire
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 .
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 . 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 . Higher  and lower  daily nest survival rates have been reported for eastern towhees in other areas (see Timing of Major Life History Events).
Life History and Behavior
Lifespan, longevity, and ageing
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).
Molecular Biology and Genetics
Barcode data: Pipilo erythrophthalmus
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.
-- end --
Download FASTA File
Statistics of barcoding coverage: Pipilo erythrophthalmus
Public Records: 3
Specimens with Barcodes: 5
Species With Barcodes: 1
IUCN Red List Assessment
Red List Category
Red List Criteria
National NatureServe Conservation Status
Rounded National Status Rank: N4B - Apparently Secure
Rounded National Status Rank: N5 - Secure
NatureServe Conservation Status
Rounded Global Status Rank: G5 - Secure
Global Short Term Trend: Decline of 10-30%
Comments: BBS data indicate a significant population decline of 2%/year from 1966-1998. The rate of statistically significant declines (for states with >25 survey routes) ranges from 1.1-9.1%/year (Alabama and Massachusetts, respectively). Population declines are greater in the eastern BBS region (2.0%/year) than the central region (0.5%/year; Sauer et al. 1999). This downward population trend as indicated by BBS data is corroborated by CBC data. Between 1959 and 1988, CBC data reveal a significant decline of 1.1%/year (Sauer, et al. 1996). Autumn banding data, collected at Manomet Bird Observatory, Massachusetts from 1970-1988, corroborate the BBS and CBC data sets (Hagan 1993). The steady decline since the 1960s is thought to be the result of forest succession (reduction in early successional habitat; Hagan 1993). In a study of Chestnut Oak (QUERCUS PRINUS) forests of the Smoky Mountains, towhee numbers declined between 1947 and the early 1980s. The decline followed canopy closure and a reduction of understory vegetation (Wilcove 1988). Although not statistically significant, counts of breeding birds have increased in Kansas, Louisiana, Mississippi and Nebraska (Sauer et al. 1999).
Comments: Significant declines over last 30 years thought to be primarily the result of forest succession (reduction in early successional habitat). HABITAT CHANGE: Preferred habitat can be lost through vegetation succession or overgrazing. Nearly eliminated from deciduous forest overgrazed by White-tailed Deer (ODOCOILEUS VIRGINIANUS), Elk (CERVUS ELAPHUS), and Mouflon Sheep (OVIS MUSIMON) due to the loss of understory vegetation (Casey and Hein 1983). Abundance was also much lower in unthinned oak woodlands inhabited by numerous White-tailed Deer (13-23/km2) than similar stands inhabited by fewer deer (1-3/km2). Woody understory vegetation was sparse in stands having high deer densities. In thinned stands, however, deer density had no impact on abundance (DeGraaf et al. 1991). Population densities are lower in urbanized areas relative to forested areas due to reduction in suitable habitat (Beissinger and Osborne 1982). Maturation of successional habitats also results in lower population densities (Hagan 1993, Wilcove 1988). NEST PARASITISM: Although forest-dividing corridors can improve habitat conditions for towhees, they also attract Brown-headed Cowbirds (MOLOTHRUS ATER; Rich et al. 1994), which commonly parasitize nests (Friedmann 1963). Frequency of cowbird nest parasitism varies from 3.4% in New York and New Jersey (Greenlaw 1996) to 54.2% in Pennsylvania (Norris 1947). PREDATION: Known nest predators include Pine Snake (PITUOPHIS MELANOLEUCUS), Rat Snake (ELAPHE OBSOLETA), Common Garter Snake (THAMNOPHIS SIRTALIS), and Eastern Chipmunk (TAMIAS STRIATUS). Suspected nest predators include Blue Jay (CYANOCITTA CRISTATA) and weasels (MUSTELA spp.). Known predators of adults include Short-eared Owl (ASIO FLAMMEUS), Sharp-shinned Hawk (ACCIPITER STRIATUS), Cooper's Hawk (ACCIPITER COOPERII), Short-tailed Hawk (BUTEO BRACHYURUS), and Loggerhead Shrike (LANIUS LUDOVICIANUS; Barbour 1951, Greenlaw 1996, Ogden 1974, Storer 1966). OTHER: Migrants are sometimes killed in collisions with towers (Taylor and Anderson 1973).
Restoration Potential: Restoration potential is good; previously populations apparently increased in the eastern U.S. in response to post-Columbian forest clearing and fragmentation of the deciduous forest (Hagan 1993).
Preserve Selection and Design Considerations: Classified as an forest interior and forest edge species (Blake and Karr 1987). Selection of forest fragments in fragmented landscapes is generally area independent. Has been observed in forest fragments as small as 0.01 hectares in New Jersey, 0.5 hectares in South Carolina, and 4.7 hectares in Illinois (Blake and Karr 1987, Forman et al. 1976, Galli et al. 1976, Kilgo et al. 1997). Equally as likely to inhabit isolated hardwood stands surrounded by fields as those surrounded by pine plantations in South Carolina (Kilgo et al. 1997).
Management Requirements: Because open-canopied, shrubby habitats are preferred, management should include activities that promote early- to mid-seral successional habitats. In forested habitat, opening the canopy benefits this species by favoring growth of the understory. Open-canopied, shrubby secondary forests can be created by clearcutting, intensive thinning, or by making large group selection cuts (1-2 ha; Annand and Thompson 1997, Crawford et al. 1981). Forest-dividing corridors can also promote habitat suitability (Anderson et al. 1977, Chasko and Gates 1982, Rich et al. 1994). Fire exclusion in Southeastern pinelands allows previously fire-suppressed hardwoods to flourish, which initially benefits this species. Habitat changes associated with long-term fire exclusion (>15 years), however, do not (Engstrom et al. 1984). A fire regime that favors shrubby understory vegetation while maintaining an open canopy will benefit towhees.
Management Research Needs: Need research on breeding success in different seral stages to determine optimal versus marginal habitat (J. Greenlaw, pers. comm.), as abundance alone is not always a reliable indicator of relative habitat quality (Vickery et al. 1992).
Biological Research Needs: Need information on dispersal of juveniles from natal territories, juvenile survival and recruitment, age- and sex- specific survivorship, and breeding and wintering ecology (Greenlaw 1996).
Use of Fire in Population Management
When managing for eastern towhees, fire interval should be carefully considered before implementing a burning plan [14,42]. Surface burns in open woodlands, grasslands, and wetlands at intervals that would eventually reduce low woody cover would likely result in eastern towhee declines. Eastern towhees have been observed to peak 4 years after fire in Florida on an old-field loblolly-shortleaf pineland, in slash pine flatwoods with saw palmetto (Serenoa repens), myrtle oak, and sand live oak in the understory, and in a scrub community comprised of scattered slash pine and cabbage palm [14,30]. The most favorable fire interval for eastern towhee depends on the habitat and other management objectives, but fire intervals of ≤3 yrs will likely result in eastern towhee absence or smaller population sizes. Fire exclusion in these areas is also likely to cause eastern towhee declines. As the canopy closes the low shrubby species that provide cover and food decline. Four years after fire in a Florida old-field loblolly-shortleaf pineland, eastern towhee detections gradually declined. Eleven years after fire eastern towhee detections were less than detections 1 year after fire .
Less is known about eastern towhees response to severe fires. In dense-canopied forests, stand-replacing fires may provide patches of early successional vegetation necessary for eastern towhee occurrence. Although there are no data on eastern towhee occurrence in severely burned forest, eastern towhees have increased on sites subjected to other canopy-opening disturbances that may have less effect on the understory, such as wind throw or logging [28,39,91]. On a sand pine scrub site in Florida that was burned in a stand-replacing fire and salvage logged 5 to 7 years earlier, eastern towhees were more abundant than on a mature, control site. The extent to which eastern towhees were responding to the effects salvage logging is unknown .Although it is likely that reasonable inferences regarding eastern towhees response can be made from the impact fire has on understory vegetation, other factors such as a fire's affect on food availability and abundance of predators and competitors should also be considered when using fire as a management tool for eastern towhee .
Data from the Breeding Bird Survey and Christmas Bird Counts show moderate to large declines in eastern towhee populations, especially in the northern part of its range [11,18,44,72]. For example, trends from 3 surveys showed that 1989 population levels were 13% of 1966 levels in the Northeast . A literature review provides some information suggesting the decline has slowed in some areas in recent years .
Managing habitat for the eastern towhee and species with similar habitat requirements has been the subject of many articles including [20,22,65,90,104]. Greenlaw's  literature review notes the impact of urbanization and agriculture on eastern towhee habitat and asserts the importance of preserving pitch pine barren habitats in the Northeast, where eastern towhees still occur in relatively high densities.
Relevance to Humans and Ecosystems
Stewardship Overview: Significant population declines have occurred in the last 30 years, particularly in the northeastern portion of the range. This is thought to be the result of forest succession (reduction in early successional habitat). Considered a forest interior/forest edge, area-independent species. During the breeding season, has been observed in forest fragments as small as 0.01 ha. Because open-canopied, shrubby habitats are preferred, management should include activities that promote early- to mid-seral successional habitats. Preferred habitat can be created/maintained by forest management techniques such as clearcutting, intensive thinning, or group selection cutting. In southern pinelands, a fire regime that favors understory growth while maintaining an open canopy benefits this species.
The eastern towhee (Pipilo erythrophthalmus) is a large New World sparrow. The taxonomy of the towhees has been under debate in recent decades, and formerly this bird and the spotted towhee were considered a single species, the rufous-sided towhee.
Their breeding habitat is brushy areas across eastern North America. They nest either low in bushes or on the ground under shrubs. Northern birds migrate to the southern United States. There has been one record of this species as a vagrant to western Europe; a single bird in Great Britain in 1966.
The song is a short drink your teeeeea lasting around one second, starting with a sharp call ("drink!") and ending with a short trill "teeeeea". The name "towhee" is onomatopoeic description of one of the towhee's most common calls, a short two-part call rising in pitch and sometimes also called a "chewink" call.
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The eastern towhee is a large and striking species of sparrow. The total length ranges from 17.3 to 23 cm (6.8 to 9.1 in) and the wingspan is 20–30 cm (7.9–11.8 in). The body of mass of this species ranges from 32 to 53 g (1.1 to 1.9 oz), with an average of 40 g (1.4 oz). Adults have rufous sides, a white belly, and a long dark tail with white edges. The eyes are red, white for birds in the southeast. Males have a black head, upper body, and tail; these parts are brown in the female. Juveniles are brown overall. Eastern towhees of all ages and both sexes generally are unmistakable and are not known to co-exist with the similar western spotted towhee.
The eastern towhee occurs throughout the eastern United States and southeast Canada. Occurrences from southern Saskatchewan, southwest Ontario 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 primarily are summer residents.
Pipilo e. erythrophthalmus occurs in the most northerly part of the eastern towhee's distribution in the summer, and in winter migrates to the southern and eastern portion of the range of the species. The other subspecies are largely residents. 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. 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.
The eastern towhee occurs in vegetation of disturbed areas, such as old-field successional vegetation and shrubby areas of power line right-of-ways. In northwestern Arkansas, eastern towhees occurred in old-field vegetation where dwarf sumac (Rhus copallina) occurred at a frequency of 28.6%, winged elm (Ulmus alata) at a frequency of 21%, and black cherry (Prunus serotina) at a frequency of 19.2%. Shrubby vegetation along power lines commonly is used by eastern towhees. For example in Maryland, eastern towhee territories along a power line right-of-way corresponded with shrubby areas containing species such as Allegheny blackberry (Rubus allegheniensis) and blueberry (Vaccinium spp.). Other species included hawthorn (Crataegus spp.), red maple (Acer rubrum), black cherry, and black locust (Robinia pseudoacacia).
Timing of major life events
Arrival and departure of eastern towhees into summer breeding grounds varies with location. According to a literature review, eastern towhees typically arrive in New York in early April and leave by the middle of November. A review of eastern towhees in New Hampshire describes arrival in late April to May with the majority departing in September. Further south, on the border of Tennessee and North Carolina, migration to high-elevation areas within the Great Smoky Mountains begins as early as March. Eastern towhees typically leave these sites in October. The Pipilo e. erythrophthalmus subspecies is the most migratory of the subspecies.
Breeding begins in spring and continues to late summer. Reports of eastern towhees nesting as early as late March in Florida and Georgia, in mid- to late April in some midwestern states, and as late as mid-May in northern New England were summarized in a literature review. Literature reviews also report nest construction by the female, which takes about three to five days. Egg laying typically occurs until August. For example, a review of eastern towhees in Indiana notes nesting from 15 April to 20 August. However, a literature review of eastern towhees in Florida included a report of a nest observed on 2 September 1983 that contained two eggs. According to several literature reviews, eastern towhees may renest after failed nesting attempts and can raise two, and in the south sometimes three, broods per season.
In a literature review, Greenlaw  reports mean breeding territory size of 4 acres (1.6 ha) (range 1.6–6 acres (0.64–2.44 ha), n=24) in a mesic oak (Quercus species) forest where eastern towhees occurred at a density of 21 males/40 ha. In a xeric pine (Pinus species)-oak woodland where the eastern towhee density was 32 males/40 ha, mean eastern towhee territory size was 3 acres (1.2 ha) (range 1.8–4 acres (0.71–1.65 ha), n=20). In Massachusetts, mean male eastern towhee territory size was about 1.3 acres (0.52 ha), and female eastern towhee territory size was 1.1 acres (0.45 ha). Territory size changed over the course of the breeding season and was not significantly (p>0.05) affected by reductions in food availability of 30% or less. During the winter eastern towhees are not as territorial and may be seen in mixed species flocks. Daily movement of eastern towhees in loblolly (P. taeda) and longleaf pine (P. palustris) forests and clearcuts in South Carolina averaged 325 feet (99 m) per day. Only 2 females, out of 11 females and 9 males, stayed within the stand where they were captured for the duration of a 10-week study.
Eastern towhees have fairly strong fidelity to breeding territories. In an oak forest in New Jersey, adult eastern towhee return rates were 20% the 1st year after banding and 43% in subsequent years. Between 1960 and 1967, the maximum number of eastern towhee returns to the site was 5. In a Pennsylvania woodlot observed between 1962 and 1967, an eastern towhee returned to the site for 4 consecutive years.
Several reviews report eastern towhee clutch sizes from 2 to 6 eggs, with means ranging from 2.45 to 3.6 eggs per nest. All 5 eastern towhee nests on Sanibel Island, Florida, contained 3 eggs. Eastern towhees in 2 pitch pine (P. rigida) barrens sites in New Jersey and New York had a later median egg laying date (mid-June) and significantly (p<0.05) smaller average early nest clutch sizes (NJ=2.67, NY=3.25) than those in an oak-hickory (Carya spp.) site, which had a median egg-laying date in early June and an average early nest clutch size of 3.88. Food availability likely explains at least some of the differences between the 2 habitat types. Eggs are incubated by the female for 12 or 13 days. After hatching both parents feed the young, which fledge 10 to 12 days later and are dependent on parental care for about another month.
A wide range of eastern towhee nest success values have been reported. On Sanibel Island, 1 of the 5 eastern towhee nests observed was successful. In Louisiana, average daily nest success rate was 95.3% on a bottomland hardwood forest site. The same study found a 92.6% average daily nest success rate in a 6-year-old managed cottonwood (Populus spp.) plantation in Alabama. Average eastern towhee nest success across mixed bigtooth (P. grandidentata) and quaking aspen (P. tremuloides) stands of varying ages in Pennsylvania was 48.1%. In South Carolina, only 1 of 10 nests was successful, and the mean daily nest survival rate was 62.9%. This low value was explained by high levels of predation. Due to lower nest success rates of Bachman's sparrow (Aimophila aestivalis) than the previous year, it is suggested that eastern towhee nest success may have been measured during a comparatively poor year.
Compared to nests, adult towhee survival rates are high. Average weekly adult survival rate of eastern towhees in a South Carolina study area was 99.3%. This rate was obtained from radio-marked eastern towhees and represented the pooled survival of both sexes and from 2 South Carolina sites, young and mature stands of loblolly and longleaf pine. Between 1962 and 1967 in Pennsylvania, annual survival of breeding eastern towhees calculated from mistnetting recaptures was 58%. According to a literature review, both males and females become reproductively mature in their second year. Eastern towhees of over 12 years old have been reported in the wild.
Eastern towhees range from near sea level to as high as 6,500 ft (2,000 m) along the border of Tennessee and North Carolina during the summer. A literature review reports eastern towhees up to 3,000 ft (910 m) in New Hampshire.
Eastern towhees spend the majority of their time near the ground. For instance, in Pennsylvania in spring, observations of eastern towhees below 3 ft (0.91 m) from the ground occurred significantly (p<0.05) more than expected based on random spatial distribution, and observations above 3 ft (0.91 m) occurred significantly (p<0.05) less than would be expected. In a Louisiana bottomland forest 62% of eastern towhee observations were within 2 ft (0.61 m) of the ground, and only 4% were observed above 25 ft (7.6 m). In the spring this changed, with detections of eastern towhees below 25 ft (7.6 m) declining from 70% to 65% and detections in the canopy (>25 ft (>7.6 m)) increasing from 4% to 7%.
Eastern towhees occur in many habitats, from tallgrass prairies and marshes to mature forests. However, eastern towhees are most common in early successional stands, habitat edges, and areas with similar vegetation structure throughout eastern forests.
In most communities eastern towhees are more abundant in young successional stands. Several studies found increased eastern towhee abundance on early successional sites compared to later-successional sites. Mean number of breeding eastern towhees (0.70 bird/50-m radius) and nest success rate (58%) were higher in a 15-year-old clearcut in West Virginia, than in other treatments, including a stand of yellow-poplar (Liriodendron tulipifera), black cherry, red maple, sugar maple (Acer saccharum), and white ash (Fraxinus americana) that was not harvested. In a southern Missouri oak-hickory forest, eastern towhees were not present before clearcutting or in the nearby uncut forest after cutting, but occurred at a mean density of 9.3 birds/10 ha in a 3-year-old clearcut. A study of stands of varying ages in central New York found that eastern towhee density peaked in early successional stands.
Although eastern towhees generally prefer young successional sites, variation between habitat types and years has been observed. Krementz and Powell  found higher relative abundance of eastern towhee in young (2–6 years old) stands of loblolly and longleaf pine than mature (32–98 years old) stands when investigated in 1995. However, in stands compared in 1996, the 95% confidence intervals of eastern towhee relative abundance on the 2 sites had a substantial degree of overlap. The degree to which eastern towhee responds to succession is influenced by habitat. For example, in Pennsylvania there was a significant (p<0.05) difference between eastern towhee densities (number/10 ha) between mature mixed-oak forest and stands that had been clearcut about 5 years previously. However, eastern towhee densities did not differ significantly between a 1-year old mixed aspen (Populus tremuloides, P. grandidentata) clearcut, a 5-year old aspen clearcut, and a mature aspen stand. Eastern towhee abundance has been shown to peak at different times in different habitats. For instance, although in central hardwood forests eastern towhees were most abundant in regenerating stands, in loblolly and shortleaf pine forest they were most common in pole timber and mature stands. In addition, Bell and Whitmore concluded that early successional is likely too broad of a term for describing optimal towhee habitat, since high density of small trees was negatively associated with eastern towhee density in the eastern panhandle of West Virginia.
Eastern towhees seem to prefer sites with characteristics generally associated with early successional vegetation, such as low canopy cover and dense understory. Negative correlations between eastern towhee abundance and various measurements of overstory density have been found in several studies. Average density of eastern towhees across 6 habitat types in Pennsylvania was significantly (p<0.05) negatively correlated with density of overstory trees and basal area of overstory trees. Number of eastern towhees in a western Virginia hardwood forest was also significantly (p<0.05) inversely correlated with total percent canopy cover. In a loblolly pine forest in South Carolina, the average number of eastern towhee breeding territories per experimental unit was significantly (p≤0.008) negatively correlated with mid-story (10–46 ft (3.0–14.0 m)) pine (Pinus spp.) and deciduous volume.
Many studies have demonstrated a positive correlation between eastern towhee abundance and understory density. In a loblolly pine forest in South Carolina, understory (0–10 ft (0–3 m)) pine volume was significantly (p<0.001) positively correlated with the average number of eastern towhee territories per experimental unit. Yahner  found the average density of eastern towhees over 6 habitat types was significantly (p<0.05) positively correlated with density of short (2–5 foot (0.5–1.5 m)) shrubs. In east-central Florida slash pine (P. elliottii) flatwoods with understories dominated by myrtle oak (Q. myrtifolia) and sand live oak (Q. geminata) and in scrub sites with scattered slash pine and cabbage palmetto (Sabal palmetto), eastern towhee densities were significantly (p=0.01) negatively correlated with mean shrub height.
Eastern towhees may associate with and avoid certain plants. In Nevada, Towhees seem to prefer a habitat in which they are camouflaged such as flowering quince which matches their rufous coloration. In riparian vegetation in Iowa, eastern towhee density was significantly (p≤0.01) positively associated with total plant and vine species richness and negatively correlated with forb and deciduous tree species richness. In West Virginia, eastern towhees were associated with plant species that occurred on drier ridgetops, such as blackberry (p<0.02), black cherry (p<0.002), and black locust (p<0.04). These sites tended to have open canopies and low tree density. Eastern towhee density was negatively associated with plants of the moister parts of this study area, such as black tupelo (Nyssa sylvatica, p<0.006), red maple (p<0.001), and witch hazel (p<0.03). In central New Jersey eastern towhees were significantly (p=0.03) more abundant in gray dogwood (Cornus racemosa) shrubland than either eastern redcedar (Juniperus virginiana) or multiflora rose (Rosa multiflora) shrublands.
Nesting habitat: Eastern towhees typically nest on or near the ground. Several literature reviews note the predominance of eastern towhee nests below 5 feet (1.5 m). In a study of cowbird parasitism on Sanibel Island, all 5 eastern towhee nests located were within 6 feet (2 m) of the ground. Nests as high as 18 feet (5.5 m) have been reported in literature reviews. Nests higher off the ground in mixed aspen stands of varying ages in Pennsylvania had significantly (p<0.001) lower nest success. Of 13 unsuccessful eastern towhee nests, 11 were greater than 1 foot (0.5 m) above the ground.
In West Virginia, there were no significant (p>0.05) differences in habitat surrounding successful and unsuccessful nests. Large snags (≥9 inches diameter at breast height (≥22.9 cm)) did not have an effect on nesting success. Nesting success was not significantly (p>0.05) affected by stand age or distance to edge in even-aged mixed-aspen stands in Pennsylvania.
Eastern towhees nest in a variety of species including grape and blueberry (Vaccinium spp.). The majority of nests observed in South Carolina loblolly and longleaf pine forests and clearcuts were located in grape, tree sparkleberry (V. arboreum), and oak (Quercus spp.). On an oak-hickory site in West Virginia, 27% of 41 eastern towhee nests were found in grape, 17% in blackberry (Rubus spp.), 12% in greenbrier (Smilax spp.), and 12% in mountain-laurel (Kalmia latifolia). Nests also occurred in Virginia creeper (Parthenocissus quinquefolia), spice bush (Lindera benzoin), and azalea (Rhododendron spp.). In a power line right-of-way in Pennsylvania, the 6 eastern towhee nests observed occurred in Allegheny blackberry, witch-hazel (Hamamelis virginiana), blueberry (Vaccinium spp.), white oak (Q. alba), eastern hayscented fern (Dennstaedtia punctilobula) and sweetfern (Comptonia peregrina) combined, and on ground level. According to literature reviews, eastern towhee nests located on the ground are embedded in litter in dry areas and typically occur at the base of grasses, forbs, low shrubs, or small trees.
Foraging habitat: Selection of forging habitat by eastern towhees has been investigated in Massachusetts and New Jersey. When gleaning in a southeastern Massachusetts pitch pine barren, eastern towhees preferred species such as pitch pine, bear oak (Q. ilicifolia), and other deciduous trees, mainly oaks. Ericaceous species were avoided. Use differed significantly (p<0.001) from availability. On 2 New Jersey sites, eastern towhee foraging preference switched over the course of the breeding season. On a site dominated by oaks, primarily black oak (Q. velutina), eastern towhees used oaks in May, as would be expected due to their density. However, in June and July, as relative arthropod biomass declined in oaks, use of oaks was less than would be expected. On a pitch pine-dominated site, use of oaks (primarily bear oak and blackjack oak (Q. marilandica)) was greater than would be expected in May, but was proportionate to availability in June and July. These negative correlations between date and oak use were significant (p<0.025) for both sites. Use of the oak-dominated site also decreased significantly (p<0.05) through the summer.
Effects of spatial area: Eastern towhees appear to prefer edge habitats in many areas. For instance, the mean abundance of eastern towhees in a baldcypress (Taxodium distichum) in northern Florida was 18, while eastern towhees did not occur in either the baldcypress forest or the clearcut. At the interface of the baldcypress stand and a 13-year-old planted slash pine stand, mean abundance of eastern towhees was 22 breeding birds, while in the planted slash pine stand the average abundance was 15 breeding birds. Density of eastern towhees was found to decline with distance from the edge of a power line right-of-way and an oak-hickory forest in eastern Tennessee. At the edge, eastern towhees occurred at a density of just over 10 pairs/40 ha, while 197 feet (60 m) from the edge eastern towhee density had dropped to 1 pair/40 ha. In addition, in experimentally clearcut Pennsylvanian forests composed of white oak, northern red oak (Q. rubra), chestnut oak (Q. prinus), scarlet oak (Q. coccinea), red maple, quaking aspen, bigtooth aspen, and pitch pine, male towhees were detected significantly (p<0.05) more often than expected in the areas where the spatial arrangement of clearcuts was most patchy.
Several studies have addressed the effect of the size of habitat patches on eastern towhees. In mixed-oak forest in New Jersey, eastern towhee frequency generally increased with patch size, although eastern towhees were detected in all plot sizes (0.5–59 acres (0.2–24 ha)) except 0.02–acre (0.01 ha) plots. On a site in South Carolina, eastern towhee frequency of occurrence increased as clearcut size increased from <2.5 acres (<1 ha) to clearcut sizes from 21 to about 32 acres (8.5–12.8 ha). On another site eastern towhee frequency declined as clearcut size increased from 19 to 33 acres (7.6–13.2 ha) to 48 to 62 acres (19.5–25.2 ha). Eastern towhees only bred in riparian vegetation patches in Iowa that were at least 650 feet (200 m) wide. In southern and eastern Pennsylvania eastern towhee nest success was not significantly (p≥0.10) different on sites with gradual edges and those with more distinct edges between "wildlife habitat openings" and oak-hickory forest.
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. When foraging on the ground eastern towhees use a scratching technique where both feet kick back simultaneously. In a laboratory study 4 eastern towhees used this method to successfully obtain seed buried almost 1 inch (2.25 cm) deep. When foraging above ground the majority of time is spent gleaning foliage. 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. 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.
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. Reviews report eastern towhees foraging at feeders. Reviews show that animal matter makes up a larger proportion of the diet in the breeding season. In fall and winter, plants make up 79% and 85% of the diet, respectively. This drops to 53% in spring and 43% in summer. 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. 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.
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. 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. Several birds are known to prey on both young and adult eastern towhees, including northern goshawk (Accipiter gentilis), Broad-winged (Buteo platypterus), short-tailed (Buteo brachyurus), sharp-shinned (Accipiter striatus) and Cooper's hawks (Accipiter cooperii). Other predators include barred (Strix varia), short-eared (Asio flammeus) and eastern screech-owls (Megascops asio) and even the loggerhead shrike (Lanius ludovicianus), which is scarcely larger than a towhee. 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).
Brown-headed cowbirds (Molothrus ater) parasitize eastern towhee nests. In a South Carolina old field, 5 of 19 eastern towhee nests were parasitized. 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. 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. In a Pennsylvania study site, only 2 of 36 nests were parasitized and both produced eastern towhee fledglings. In a study of nest parasitism on Sanibel Island, none of 5 eastern towhee nests found were parasitized.
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|Wikimedia Commons has media related to Eastern towhee.|
|Wikispecies has information related to: Pipilo erythrophthalmus|
- Eastern towhee species account – Cornell Lab of Ornithology
- Eastern towhee – Pipilo erythropthalmus – USGS Patuxent Bird Identification InfoCenter
- Eastern towhee videos, photos, and sounds at the Internet Bird Collection
- Stamps (for United States) with range map at bird-stamps.org
- Eastern towhee photo gallery at VIREO (Drexel University)
- Eastern towhee bird sound at Florida Museum of Natural History
- Interactive range map of Pipilo erythrophthalmus at IUCN Red List maps
Names and Taxonomy
Comments: Formerly regarded as conspecific with P. MACULATUS (AOU 1998). Citing morphological, behavioral, and mtDNA differences (e.g., Ball and Avise 1992), AOU (1995) split the rufous-sided towhee into two species, P. ERYTHROPTHALMUS (eastern towhee, central and eastern North America) and P. MACULATUS (spotted towhee, mainly west of the Great Plains). Limited hybridization occurs in a narrow zone in the central Great Plains. See Banks and Browning (1995) for a discussion of nomenclatural issues involving PIPILO.
towhee, a member of the Emberizidae family . In 1995, the American Ornithologists'
Union split the rufous-sided towhee (P. erythrophthalmus)
into the eastern and spotted towhee (P. maculatus) .
Hybridization between the 2 species occurs along riparian corridors in the Great
Plains, especially on the Platte River [42,82]. The 4 eastern towhee
subspecies recognized by the American Ornithological Union  are:
P. e. ssp. erythrophthalmus (Linnaeus)
P. e. ssp. alleni Coues
P. e. ssp. canaster Howell
P. e. ssp. rileyi Koelz