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Lifespan, longevity, and ageing

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Maximum longevity: 12.3 years (wild)
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Pipilo erythrophthalmus

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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

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Reid Rumelt

Pipilo erythrophthalmus

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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.

References

  • 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
  • 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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Rumelt, Reid B. Pipilo erythrophthalmus. June-July 2012. Brief natural history summary of Pipilo erythrophthalmus. Smithsonian's National Museum of Natural History, Washington, D.C.
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Common Names

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COMMON NAME: [3]




eastern towhee
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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Direct Effects of Fire

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More info for the terms: fire interval, fire severity, forest, 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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Distribution

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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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Food Habits

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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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Habitat-related Fire Effects

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More info for the terms: cover, density, fire exclusion, fire interval, fire regime, fire severity, forest, frequency, fruit, grassland, habitat type, herb, herbaceous, litter, low-severity fire, seed, severity, shrubs, species richness, 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.

FIRE REGIMES: The following table provides fire return intervals for plant communities and ecosystems where eastern towhee is likely to occur. Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find FIRE REGIMES".

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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Habitat: Cover Types

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More info on this topic.

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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

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 terms: bog, forest

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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Management Considerations

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MANAGEMENT CONSIDERATIONS:
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 [44]. A literature review provides some information suggesting the decline has slowed in some areas in recent years [42].

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 [42] 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.
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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Predators

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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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

States or Provinces

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(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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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Taxonomy

provided by Fire Effects Information System Animals
Pipilo erythrophthalmus (Linnaeus) is the scientific name for eastern
towhee, a member of the Emberizidae family [3]. In 1995, the American Ornithologists'
Union split the rufous-sided towhee (P. erythrophthalmus)
into the eastern and spotted towhee (P. maculatus) [2].
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 [1] are:



P. e. ssp. erythrophthalmus (Linnaeus)

P. e. ssp. alleni Coues

P. e. ssp. canaster Howell

P. e. ssp. rileyi Koelz
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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Use of Fire in Population Management

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More info for the terms: cover, fire exclusion, fire interval, forest, stand-replacing fire

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 [30].

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 [38].

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 [78].
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Meyer, Rachelle. 2006. Pipilo erythrophthalmus. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: www.fs.fed.us/database/feis/animals/bird/pier/all.html

Comprehensive Description

provided by Smithsonian Contributions to Zoology
Pipilo erythrophthalmus (Linnaeus)

In the last general survey of the brown-headed cowbird's hosts (Friedmann, 1963:150–151), it was noted that few instances of parasitism on this towhee had been reported from California, but that the recent spread and increase of the parasite in the southern parts of that state would probably result in further records. Three such instances have come to light: 1, collected in San Luis Obispo County, 2 June 1934, is now in the Santa Barbara Museum of Natural History, and 2 more—1 from near Mendota, Fresno County, 1 June 1935, and 1 from near Stockton, 21 June 1942—are in the collections of the Western Foundation. The San Luis Obispo record involves the host subspecies P. erythrophthalmus megalonyx; the other 2 have to do with the race P. erythrophthalmus falcinellus. The last mentioned subspecies is new to the host catalog of the parasite.

The Western Foundation's collections contain 393 sets of eggs of this towhee (all races), 11 of which (2.8 percent) are parasitized. A list of only those races of the host known to be victimized by the cowbird, yields the following data: P. erythrophthalmus erythrophthalmus, 55 sets, 9 (16.3 percent) parasitized; P. erythrophthalmus falcinellus, 8 sets, 2 (25.0 percent) parasitized; P. erythrophthalmus canaster, 3 sets, none parasitized; P. erythrophthalmus arcticus, 7 sets, none parasitized; P. erythrophthalmus montanus, 14 sets, none parasitized; P. erythrophthalmus curtatus, 1 set, not parasitized; P. erythrophthalmus oreganus, 37 sets, none parasitized; P. erythrophthalmus falcifer, 57 sets, none parasitized; and P. erythrophthalmus megalonyx, 173 sets, none parasitized.

The Ontario nest records files at Toronto have data on 70 nests, 14 of which (20.0 percent) were parasitized.

BROWN TOWHEE
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bibliographic citation
Friedmann, Herbert, Kiff, Lloyd F., and Rothstein, Stephen I. 1977. "A further contribution of knowledge of the host relations of the parasitic cowbirds." Smithsonian Contributions to Zoology. 1-75. https://doi.org/10.5479/si.00810282.235

Eastern towhee

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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.[2]

Taxonomy

The eastern towhee was described by the Swedish naturalist Carl Linnaeus in 1758 in the tenth edition of his Systema Naturae and given the binomial name Fringilla erythrophthalma.[3] This species is now placed in the genus Pipilo that was introduced by the French ornithologist Louis Jean Pierre Vieillot in 1816.[4] The specific name erythrophthalmus/erythrophthalma combines the Ancient Greek words ερυθρος eruthros "red" and οφθαλμος ophthalmos "eye".[5]

Four subspecies are recognised:[6]

  • P. e. erythrophalmus (Linnaeus, 1758), found south-central and southeast Canada to the eastern United States
  • P. e. canaster (Howell, AH, 1913), found in the interior southeast United States
  • P. e. rileyi (Koelz, 1939), found in the coastal southeast United States except for central and southern Florida
  • P. e. alleni (Coues, 1871) found in south and central Florida

Description

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).[7][8] 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).[9] 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.

Distribution

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.[2]

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.[10] 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.[11] 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.[10]

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%.[12] Shrubby vegetation along power lines is commonly used by eastern towhees.[13] 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).[13]

From 1966 to 2015 the eastern towhee experienced a greater than 1.5% annual population decrease throughout the Atlantic seaboard and the northern part of its breeding range.[14]

Timing of major life events

Female

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.[15] 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.[16] The Pipilo e. erythrophthalmus subspecies is the most migratory of the subspecies.[10]

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.[2] Literature reviews also report nest construction by the female, which takes about three to five days.[2][15] Egg laying typically occurs until August. For example, a review of eastern towhees in Indiana notes nesting from 15 April to 20 August.[17] However, a literature review of eastern towhees in Florida included a report of a nest observed on 2 September 1983 that contained two eggs.[18] 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.[2][15][18]

In a literature review, Greenlaw [2] reports mean breeding territory size of 4 acres (1.6 hectares) (range 1.6–6 acres (0.65–2.43 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 hectares) (range 1.8–4 acres (0.73–1.62 ha), n=20).[2] In Massachusetts, mean male eastern towhee territory size was about 1.3 acres (0.53 hectares), and female eastern towhee territory size was 1.1 acres (0.45 hectares). 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.[2] 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.[19]

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.[20] In a Pennsylvania woodlot observed between 1962 and 1967, an eastern towhee returned to the site for 4 consecutive years.[21]

Several reviews report eastern towhee clutch sizes from 2 to 6 eggs, with means ranging from 2.45 to 3.6 eggs per nest.[2][15][17][18] All 5 eastern towhee nests on Sanibel Island, Florida, contained 3 eggs.[22] 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.[2][15][17][18]

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%.[23] 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.[19]

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.[19] Between 1962 and 1967 in Pennsylvania, annual survival of breeding eastern towhees calculated from mistnetting recaptures was 58%.[21] According to a literature review, both males and females become reproductively mature in their second year.[2] Eastern towhees of over 12 years old have been reported in the wild.[24]

Preferred habitat

Eastern towhee singing in Jamaica Bay Wildlife Refuge in New York

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.[16] A literature review reports eastern towhees up to 3,000 ft (910 m) in New Hampshire.[15]

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.[25] 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%.[26]

Eastern towhees occur in many habitats, from tallgrass prairies and marshes to mature forests.[19] 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.[25][27][28][29] 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.[27] 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.[28]

Although eastern towhees generally prefer young successional sites, variation between habitat types and years has been observed. Krementz and Powell [19] 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.[30] 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.[31]

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.[12][30] 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.[30] 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.[32]

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; 0.0–3.0 m) pine volume was significantly (p<0.001) positively correlated with the average number of eastern towhee territories per experimental unit.[32] Yahner [30] 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.[33]

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.[34] 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).[31] 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.[35]

Nest with hatchlings in North Carolina

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).[2][15][17][18] In a study of cowbird parasitism on Sanibel Island, all 5 eastern towhee nests located were within 6 feet (1.8 m) of the ground.[22] Nests as high as 18 feet (5.5 m) have been reported in literature reviews.[2][15][17] 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 (0.30 m) above the ground.[23]

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.[36] Nesting success was not significantly (p>0.05) affected by stand age or distance to edge in even-aged mixed-aspen stands in Pennsylvania.[23]

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.).[19] 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.).[36] 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.[2][17]

Foraging habitat: Selection of foraging 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.[37] On 2 New Jersey sites, eastern towhee foraging preference switched over the course of the breeding season.[38] 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.[38]

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.[39] 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.[40]

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 (0.20–23.88 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 (1.0 hectare) 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 48 to 62 acres (7.7 to 19.4 to 25.1 ha).[29] Eastern towhees only bred in riparian vegetation patches in Iowa that were at least 650 feet (200 m) wide.[34] 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.[41]

Food habits

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.[37] When foraging on the ground eastern towhees use a scratching technique where both feet kick back simultaneously.[18] In a laboratory study 4 eastern towhees used this method to successfully obtain seed buried almost 1 inch (2.5 cm) deep. When foraging above ground the majority of time is spent gleaning foliage.[37] 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.[37]

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.[18] Reviews report eastern towhees foraging at feeders.[2] Reviews show that animal matter makes up a larger proportion of the diet in the breeding season.[2][42] In fall and winter, plants make up 79% and 85% of the diet, respectively. This drops to 53% in spring and 43% in summer.[42] 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.[2][42] 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.[42]

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.[2] 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.[2] 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.[43][44][45] 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).[46]

Brown-headed cowbirds (Molothrus ater) parasitize eastern towhee nests. In a South Carolina old field, 5 of 19 eastern towhee nests were parasitized.[47] 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.[47] 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.[36] In a Pennsylvania study site, only 2 of 36 nests were parasitized and both produced eastern towhee fledglings.[41] In a study of nest parasitism on Sanibel Island, none of 5 eastern towhee nests found were parasitized.[22]

References

Public Domain This article incorporates public domain material from Pipilo erythrophthalmus. United States Department of Agriculture.

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Eastern towhee: Brief Summary

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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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