Limnothlypis swainsonii



INTRODUCTORY


  2005 Đ Linda K. Williams, Martha Lafite Thompson Nature Sanctuary

AUTHORSHIP AND CITATION:
Meyer, Rachelle. 2006. Limnothlypis swainsonii. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

FEIS ABBREVIATION:
LISW

COMMON NAMES:
Swainson's warbler

TAXONOMY:
Limnothlypis swainsonii (Audubon) is the scientific name of the Swainson's warbler, a member of the Parulidae family [1].

SYNONYMS:
None

ORDER:
Passeriformes

CLASS:
Bird

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
Information on state-level protected status of animals in the United States is available at NatureServe, although recent changes in status may not be included.

ANIMAL DISTRIBUTION AND OCCURRENCE

SPECIES: Limnothlypis swainsonii

GENERAL DISTRIBUTION:
According to reviews [7,43,63], Swainson's warbler breeds in the southeastern United States and winters on Caribbean Islands and portions of the Yucatan Peninsula. Swainson's warbler's breeding range extends along the Gulf Coast from northwestern Florida to eastern Texas, north through eastern Oklahoma, and east through southern Missouri, southern Illinois, and parts of Tennessee, Kentucky, Virginia, and West Virginia to the coast of North Carolina. Although rare, Swainson's warbler also breeds in Maryland and Delaware. Swainson's warblers migrate through southern Florida or along the Gulf Coast to wintering grounds on the Yucatan Peninsula including northern Guatemala, Belize, and the Mexican states of Quintana Roo and Campeche. Swainson's warblers also overwinter in Cuba, Jamaica, the Bahamas and other Caribbean islands [7,43,63]. A map of Swainson's warbler's range can be found at NatureServe.

The following lists are speculative and are based on Swainson's warbler's distribution information and the habitat characteristics and species composition of communities Swainson's warblers are known to occupy during migration and breeding. There is not conclusive evidence that Swainson's warblers occur in all the habitat types listed, and some community types may have been omitted. Swainson's warblers are observed rarely in some of the communities listed and more often in others. See the Preferred Habitat section for more detail.

ECOSYSTEMS [22]:
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

STATES/PROVINCES: (key to state/province abbreviations)
UNITED STATES
AL AR DE FL GA IL KY LA MD MS
MO OK NC SC TN TX VA WV PR

MEXICO
Camp. Q.R. Tab. Tamps. Ver. Yuc.

BLM PHYSIOGRAPHIC REGIONS [5]:
14 Great Plains

KUCHLER [35] PLANT ASSOCIATIONS:
K100 Oak-hickory forest
K103 Mixed mesophytic forest
K104 Appalachian oak forest
K106 Northern hardwoods
K111 Oak-hickory-pine
K112 Southern mixed forest
K113 Southern floodplain forest
K114 Pocosin

SAF COVER TYPES [20]:
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
27 Sugar maple
28 Black cherry-maple
43 Bear oak
44 Chestnut oak
45 Pitch pine
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
65 Pin oak-sweetgum
70 Longleaf pine
73 Southern redcedar
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
89 Live 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
97 Atlantic white-cedar
98 Pond pine
100 Pondcypress
101 Baldcypress
102 Baldcypress-tupelo
103 Water tupelo-swamp tupelo
104 Sweetbay-swamp tupelo-redbay
108 Red maple
109 Hawthorn
110 Black oak

SRM (RANGELAND) COVER TYPES [59]:
805 Riparian
809 Mixed hardwood and pine
812 North Florida flatwoods
815 Upland hardwood hammocks

PLANT COMMUNITIES:
Several studies of Swainson's warbler have been performed in "bottomland hardwood forests". These are forests near rivers and include a mosaic of swamps, floodplain forests, and riverfront vegetation [43]. Oaks (Quercus spp.), elms (Ulmus spp.), and sweetgum (Liquidambar styraciflua) are usually major components of the canopy in these habitats [3,43,61], but several other species may be present including bald cypress (Taxodium distichum) [61], sugarberry (Celtis laevigata), ashes (Fraxinus spp.), maples (Acer spp.), and yellow-poplar (Liriodendron tulipifera) [43]. The understory is variable but typically includes saplings of canopy and subcanopy trees, cane (Arundinaria gigantea), dwarf palmetto (Sabal minor), holly (Ilex spp.), and/or vines such as greenbrier (Smilax spp.) and peppervine (Ampelopsis arborea) [3,43,52,61].

Swainson's warblers often occur in areas with cane-dominated understories [4,45,52,61]. These habitats occur in several coniferous and deciduous forests of the southeast. Cane-dominated stands are sometimes referred to as canebrakes [43,69]. In southern Missouri, 75% of canopy trees in canebrakes occupied by Swainson's warbler were either boxelder (Acer negundo, 34%), American elm (Ulmus americana, 13%), hackberry (Celtis occidentalis, 12%), honey-locust (Gleditsia triacanthos, 10%), black walnut (Juglans nigra, 4%), or American hornbeam (Carpinus caroliniana, 3%) [69]. More information is available in the Distribution and Occurrence section of the FEIS review of cane.

In addition to the plant communities listed above, Swainson's warbler is associated with the rhododendron-mountain laurel-eastern hemlock-American holly (Rhododendron spp.-Kalmia latifolia-Tsuga canadensis-Ilex opaca) community type of the Appalachian Mountains [40].


BIOLOGICAL DATA AND HABITAT REQUIREMENTS

SPECIES: Limnothlypis swainsonii
TIMING OF MAJOR LIFE HISTORY EVENTS:
Migration: According to literature reviews, Swainson's warblers migrate into the southeastern United States from late March to late April. Males typically arrive several days before females. During fall migration birds generally leave sometime from August to October, although Swainson's warblers have been recorded in southern Florida in November. More detailed information regarding migration routes and timing is available [7,43,63].

Breeding: Nesting begins in late April and early May [7,41,43,46]. Females build bulky nests in 2 to 9 days using materials near the nest site [41,42,66]. Swainson's warbler clutches typically contain 3 or 4 eggs [7,43,63]. A literature review found a mean clutch size of 3.3 eggs (n=90), with 68% of nests containing 3 eggs [7]. In canebrakes of southern Missouri, the average clutch size was 3.65 eggs [69]. Eggs are incubated for 13 to 15 days [42,43,66] and the young fledge in 10 to 12 days [7,43]. According to a review, Swainson's warblers may begin breeding at 1 year, although data supporting this are lacking [7]. A male Swainson's warbler returned to a territory in the Great Dismal Swamp in Virginia 3 years in a row [43], and a review notes site fidelity to wintering areas in Jamaica [7]. For information on breeding behaviors such as courtship, brooding, and parental care, see [7,42].

Swainson's warblers will renest, if initial nesting attempts fail. A review summarizes accounts of renesting, including a female that had 3 failed nests in 1 breeding season [7]. Although it appears fairly rare, double brooding has also been reported [43,46]. In Franklin L. Gravely Wildlife Management Area in South Carolina, the second nest of a pair of Swainson's warblers with young that fledged in early June was found on June 12, and nestlings from this nest were banded on July 8 [46].

Values of nest success for Swainson's warblers range from low to average. For example, Meanley [43] observed only 3 successful nests out of 16, and a literature review states that nest success "seems remarkably low" [7]. However, in canebrakes of Missouri the daily clutch survival (x = 0.9675) and survival during the 15-day incubation period (x = 0.6095) were within the range of other forest song birds. On average each Swainson's warbler nest produced 2.12 fledglings [69]. In a managed hardwood forest in South Carolina, 8 of 15 nests with known outcomes were successful. The estimated average daily nest survival rate was 0.961 which resulted in a 46% probability of nest success [52].

Lifespan: According to a review, the oldest documented Swainson's warbler was recaptured in Jamaica when it was 7 years and 9 months old [7].

Behavior: Descriptions of behaviors such as vocalizations, territory defense, courtship, and preening are reviewed by [7,41,43].

PREFERRED HABITAT:
Swainson's warbler breeding habitat is generally similar to migratory and wintering habitats. Although Swainson's warblers are most commonly associated with floodplain forests and mountain ravines during the breeding season [7,18,36,63], they also occur in heavily disturbed habitats such as old fields and loblolly pine (Pinus taeda) plantations [11,18,68]. Swainson's warbler wintering habitat in Jamaica and Cuba includes scrub and lowland and montane forests, including "dry limestone" and second growth forests [33,36,65].

Swainson's warblers are found at low and mid-elevation sites. Reported elevations of breeding areas range from 220 feet (67.3 m) [4] to 2,800 feet (850 m) [43]. A review notes occurrences to 4,000 feet (1,220 m) in the Appalachians [63]. Swainson's warblers were reported from sea level [36] to 4,350 feet (1,325 m) on wintering sites in Jamaica [24] and on sites up to 4,900 (1,500 m) during migration [56].

Swainson's warbler habitats have large amounts of cover especially in the understory. For more information on the vegetative characteristics of Swainson's warbler habitat see Cover Requirements.

Due to its use as a foraging substrate [3,25,36] (see Food Habits), litter is an important component of Swainson's warbler habitat. In Arkansas, sites occupied by Swainson's warbler had significantly (P<0.001) greater litter cover (x = 59.5%) than random sites (x = 41.1%) in a study area comprised of oak-hickory (Quercus-Carya) forest and box canyons. There was also significantly (P=0.003) more litter cover on occupied sites (70%) compared to unoccupied sites (50.4%) in a bottomland hardwood forest study area. Data pooled from these and 2 other Arkansas study areas showed significantly (P=0.005) greater litter cover (x = 69.5%) at sites with Swainson's warblers than random sites (x = 51.0%). In addition, sites where Swainson's warblers were detected had insignificantly (0.298P>0.05) deeper litter than random sites [4]. Litter was the third most important variable in models generated from data collected at Bond Swamp National Wildlife Refuge in Georgia to predict occupancy of habitat patches by Swainson's warblers. On average, litter depth on plots in Swainson's warbler territories was 50% greater than on unoccupied plots [45]. The average litter cover at foraging locations in bottomland hardwood forest in Louisiana (98%) was significantly (P0.05) greater than at random sites (~91%) [3]. However, leaf litter cover was significantly (P=0.011) lower in areas of Missouri canebrakes where Swainson's warblers were present (38.5%) than in areas where Swainson's warblers were not detected (55.6%) in one year of a two-year study [69]. Importance of litter in wintering habitat in Jamaica is addressed by [65].

Although Swainson's warblers are typically located near water, standing water within their territories is rare. In lowlands of southern Illinois, all observations of male Swainson's warblers were within 660 feet (200 m) of open water [18]. In a model for predicting their presence on sites in South Carolina, Swainson's warblers were more likely to occur in areas close to water [47]. However, territories usually contain little or no open water. In the Great Dismal Swamp of Virginia and North Carolina, water was not recorded at any of the 600 reference points within Swainson's warbler territories, while standing water was observed at 43% of 880 reference points in unoccupied habitat [26]. A model for predicting occupancy of habitat patches by Swainson's warbler based on data collected in Bond Swamp, Georgia included a negative relationship with standing water [45]. In Tensas River National Wildlife Refuge, Louisiana, Swainson's warblers occurred most often in the second terrace floodplain forest, which does not seasonally flood [3]. On average only 5.1% of 1-acre (0.04 ha) plots within Swainson's warbler territories in southern Illinois lowlands were completely covered by water [18]. The proximity of territories to water and the primarily terrestrial habits of the Swainson's warbler leave the birds vulnerable to flooding [26,40,43].

Swainson's warbler may avoid areas where the soil is comprised of gravel [18,27]. In southern Illinois, only 1 of 42 males occurred on sites with gravel dominated soils despite thorough searches in such areas [18].

Density/Territory: Reported territory sizes vary widely across habitats and throughout Swainson's warbler's range. In a review, Meanley [43] summarizes territory sizes observed over several years of investigation. The smallest territory was 0.3 acres (0.1 ha), while the largest territories were over 4 acres (1.6 ha). This included a 6-acre (2.4 ha) territory in the Great Dismal Swamp in Virginia and another that expanded from 4.1 acres (1.7 ha) earlier in the breeding season to 8 acres (3.2 ha) in July. Size and arrangement of habitat were suggested as possible factors influencing Swainson's warbler territory size and distribution [43]. In a second-growth forest comprised of yellow-poplar, red maple (Acer rubrum), and eastern hemlock (Tsuga canadensis) a male Swainson's warbler had a territory with a diameter of approximately 820 feet (250 m) [46]. In a managed hardwood forest in South Carolina territories averaged 4.5 acres (1.84 ha) on a sheared site (saplings and stumps removed) and 5.7 acres (2.32 ha) on an unsheared site [52]. A Swainson's warbler territory in a lowland hardwood forest of northeastern Alabama was 1.7 acres (0.69 ha) [66]. According to a review, territories in southern Illinois ranged from 0.3 to 8 acres (0.12-3.2 ha) with a mean of 2.2 acres (0.9 ha) [7].

Population densities are similarly variable. High density estimates include 25 male Swainson's warblers/kmē in oak-gum (Quercus-Nyssa) forest in Louisiana [16] and 20.5 males/kmē on a mixed-aged second-growth floodplain forest site in Louisiana. One of the lowest reported densities of Swainson's warbler was 3.8 males/kmē in second growth floodplain forest in Arkansas [27]. A fairly comprehensive summary of population densities is included in [7].

Landscape factors: It is generally accepted that Swainson's warblers prefer large areas of habitat [18,26,27]. In southern Illinois, the smallest lowland forest stand with a Swainson's warbler territory was about 865 acres (350 ha) [18]. Using typical densities of Swainson's warblers, Graves [27] suggested an area of 15,440 to 24,710 acres (6,250-10,000 ha) would be needed to support 500 pairs of Swainson's warblers.

Scale of investigation was important for detecting effects of various habitat characteristics on presence of Swainson's warbler in South Carolina [47]. A model for predicting Swainson's warbler presence included opposite relationships to the same variables at differing scales. For instance, at fine scales Swainson's warblers were less likely to occur in older stands (5-acre (2 ha) scale) of relatively uniform age (49-acre (20 ha) scale), while at large scales Swainson's warbler was positively associated with mean stand age (6,990-acre (2,827 ha) scale ) and "continuity of age" (1,210-acre (491 ha) scale) [47].

At a small scale, patchy habitat may be important to Swainson's warblers. In addition to the finding noted above regarding "continuity of age" at the 49-acre scale [47], it is possible that in some areas Swainson's warbler's require territories that have closed canopies and litter-covered, relatively open areas at ground level as well as dense understory thickets typically associated with canopy gaps [27] (see Cover Requirements). More research is needed on the relative importance of each of these features for nesting, perching, and/or foraging and the impact of their size and arrangement in several landscape contexts [4,52,61].

COVER REQUIREMENTS:
Swainson's warblers require a well-developed understory. Four study areas in Arkansas comprised mainly of bottomland hardwood forest had significantly (P<0.001) greater understory density in occupied (10.9 stems/mē) than random (5.1 stems/mē) sites. Occupied sites also had greater (P=0.022) shrub cover (13.3%) than unoccupied sites (10%) [4]. There was significantly (P<0.01) greater cover of low vegetation (0-5 feet (0-1.5 m)) at nests sites (57%) than at random sites (40%) within Swainson's warbler territories in a managed hardwood forest in South Carolina [52]. The 2 most important variables in models predicting Swainson's warbler territories in Bond Swamp National Wildlife Refuge in Georgia were cane density and the combined density of shrubs, vines, and saplings [45]. On 5 sites in Arkansas, Louisiana, Mississippi, and Florida, Swainson's warblers were observed in the greatest abundance (10-20 territorial males/kmē) on sites with many (620-820 stems/ha) small (<25 cm DBH) trees and thickets of understory vegetation. Median stem densities on these sites ranged from 31,592 stems/ha to 48,281 stems/ha. Researchers concluded that stem densities from 30,000 to 50,000 stems/ha provide the cover necessary for high-quality Swainson's warbler habitat [27]. In southern Illinois, the average stem density in Swainson's warbler territories was 26,390 stems/ha. No Swainson's warblers were detected in areas with <5,000 stems/ha, and in 40 of 53 plots stem densities were greater than 10,000 stems/ha [18].

J. Michael Meyers, USGS Patuxent Wildlife Research Center


Although cane is an important habitat component, Swainson's warblers do occur in areas where it is absent. Cane density was significantly (P=0.019) higher on sites in 4 Arkansas study areas occupied by Swainson's warblers (x = 6.1/mē) than on unoccupied sites (x = 4.6/mē) [4]. The presence of cane was an important variable in models for predicting Swainson's warbler occupancy in a South Carolina managed bottomland forest, and cane ground cover was significantly (P<0.001) higher in territory plots (sheared site = 2.66%, unsheared site = 6.00%) than unused plots (sheared site = 0.48%, unsheared site = 0.54%) [52]. Cane density was the most important variable in models predicting Swainson's warbler presence in Bond Swamp, Georgia [45]. In Big Hammock Wildlife Management Area in Georgia, cane density was insignificantly (P>0.05) greater in areas occupied by Swainson's warblers (x = 1,748; range 0 to 7,747 stems/ha) than in areas adjacent to territories (x = 601; range 0 to 2,819 stems/ha) and unoccupied control areas (x = 163; range = 0 to 1,443 stems/ha) [61]. Cane is not a required component of Swainson's warbler habitat [3,26,27,66]. In the Great Dismal Swamp, for example, cane was present on only 17 of 30 sampling plots in Swainson's warbler territories, and cane density was actually lower on sites occupied by Swainson's warblers (median = 0 culms/ha; range = 0 to 50,400 culms/ha) than unoccupied sites (median = 23,300 culms/ha; range 0 to 88,000 culms/ha) [26].

Other understory vegetation has been associated with Swainson's warblers. Vines may be an important component of Swainson's warbler habitat. In a managed bottomland hardwood forest in South Carolina, success rate of Swainson's warbler nests located in vines was higher (63%, n=8) than success rate of nests in cane (20%, n=5). In addition, categorical abundance of vines was the variable that was most consistently included in models for predicting Swainson's warbler occurrence. When cane and vines were abundant there was a 59% probability of detecting a Swainson's warbler on a site. Vine species on these sites included greenbrier, grape (Vitis spp.), trumpet creeper (Campsis radicans), and peppervine [52]. In a microhabitat model for predicting Swainson's warbler occupancy of sites in South Carolina, number of vines was positively associated with Swainson's warbler presence [47]. Swainson's warbler territories in the Great Dismal Swamp had greenbrier (Smilax laurifolia and S. rotundifolia) tangles significantly (P<0.001) more often (x = 3,300 stems/ha; range = 0 to 41,000 stems/ha) than unoccupied areas (median=0; range = 0-6,000 stems/ha). Redbay (Persea borbonia) was also significantly (P<0.001) more dense in Swainson's warbler territories. Coastal sweetpepperbush (Clethra alnifolia) occurred at significantly (P<0.001) lower densities in Swainson's warbler territories [26]. Foraging sites in northeastern Louisiana had significantly (P0.05) greater dwarf palmetto density than random sites (4.0) [3].

Although several sources suggest that Swainson's warbler habitat has little herbaceous vegetation [18,40,43], the relationship between Swainson's warblers and herbaceous ground cover is uncertain. For example, southern Missouri canebrakes occupied by Swainson's warblers had significantly (P0.05) less herbaceous ground cover (24.4%) than unoccupied areas (34.6%) in 1992, but significantly (P0.05) more (35.6%) than unoccupied areas (20.8%) in 1993 [69]. Despite no significant differences in forb cover at 3 other Arkansas study areas, Swainson's warblers were detected on sites with significantly (P<0.001) less forb cover (19.6%) than random sites (49.7%) in bottomland hardwood forest in White River National Wildlife Refuge [4]. In contrast, herbaceous ground cover in Swainson's warbler territories and adjacent unoccupied areas was greater (55-90%) than in unoccupied control plots in the bottomland forest of Big Hammock Wildlife Management Area [61]. In bottomland hardwood forest in Louisiana, the density of ground foliage in plots centered on Swainson's warbler feeding locations were in accordance with availability [3].

Swainson's warbler habitat generally has a closed canopy. Sites in 4 Arkansas study areas where Swainson's warbler were detected had significantly (P=0.024) greater canopy cover (89.3%) than unoccupied sites [4]. In bottomland forest of southern Illinois, sites where male Swainson's warblers were observed singing had >55% canopy cover and typically canopy cover exceeded 75% [18]. Despite occupied and adjacent plots in Big Hammock Wildlife Management Area in Georgia having fewer trees than control plots, canopy closure was still high, ranging from 70% to 100% [61]. Although similar to canopy closure in unoccupied (x = 82.5%) canebrakes in Missouri, sites where Swainson's warblers were detected had high canopy coverage (x = 84.1%) [69]. However, Swainson's warblers can occur in or near areas with open canopies [11,27].

Species composition of the canopy does not have a major influence on Swainson's warblers. Floristics were relatively unimportant on 5 sites in Arkansas, Mississippi, Louisiana, and Florida [27], in lowlands of southern Illinois [18], and in the Great Dismal Swamp. Although black tupelo (Nyssa sylvatica) had a negative relationship with Swainson's warblers in several accurate habitat occupancy models and occurred at significantly (P<0.001) lower basal area in Swainson's warbler territories, it is possible this relationship is due to a significant (P0.002) positive correlation (rs= 0.47) between basal area of black gum and pooled water and not avoidance of the species itself [26].

Swainson's warbler's association with other canopy characteristics is more ambiguous. On 5 sites in Arkansas, Mississippi, Louisiana, and Florida canopy height was a relatively unimportant Swainson's warbler habitat feature [27], and in Missouri canebrakes significant (P<0.05) associations with canopy height and diameter at breast height were only observed in 1 of 2 years. The minimum canopy height of occupied canebrakes was 12 feet (3.6 m) [69]. In southern Illinois, Swainson's warblers were not observed in forests with tree heights less than 25 feet (7.6 m) [18]. In the Great Dismal Swamp Swainson's warbler territories had significantly (P=0.001) fewer medium-sized (10-16 inches (25-39.9 cm)) trees, while the number of larger trees (>16 inches (40 cm)) was similar on unoccupied and occupied sites [26].

Swainson's warblers occur in habitats at various stages of succession. In bottomland hardwood forests in North Carolina, Swainson's warblers were captured at a rate of 0.24 birds/100 mist net hours in young stands (6-8 years), 0.38 birds/100 mist net hours in intermediate-aged stands (17-21 years), and 0.71 birds/100 mist net hours in mature stands (>60 years) [15]. In South Carolina, Swainson's warblers occurred at a frequency of 0.04 in mature (≥40 years) upland hardwood forest and 0.11 in a young (17 years) white pine stand [37]. In the lowlands of southern Illinois, Swainson's warblers were found in late successional forests and in old fields surrounded by mature forest [18]. On 5 sites in Arkansas, Mississippi, Louisiana, and Florida, Swainson's warbler territories in mature forest were generally near disturbance gaps [27].

Nesting: Nests are typically found at or below about 6 feet (2 m) in various vegetation. In the Great Dismal Swamp in Virginia, the average height of Swainson's warbler nests was 4 feet (1.3 m) and ranged from 18 inches (0.5 m) to 6 feet 3 inches (2 m). Nests were built in abundant understory species, including greenbrier, Japanese honeysuckle (Lonicera japonica), coastal sweetpepperbush, and cane [43]. In Franklin L. Gravely Wildlife Management Area, 2 nests were found in eastern hemlock at 6 feet (1.9 m) and just under 7 feet (2.1 m) [46]. A review notes occurrence of nests from just over 1 to 10 feet (0.4-3.1 m) in saplings, shrubs, and vines [7].

Nests are often placed near patches of dense vegetation, typically associated with canopy gaps. In a managed hardwood forest in South Carolina nests were significantly (P<0.05) closer to vine masses (11 feet (3.3 m)) than were random territory plots. They ranged from 10 to 16.4 feet (3-5 m) from these vine masses, which occurred in canopy gaps that averaged 22,600 feetē (2,100 mē) [52]. Meanley [43] found nests typically occurred along the edge of canebrakes. Similarly, a summary of a report from a hardwood forest in West Virginia states that Swainson's warblers' nests were typically placed within 50 feet (15 m) of, but not within, the densest vegetation [43]. In addition, Eddleman [18] notes a report of 2 nests occurring on the edge of canebrakes in southern Illinois.

Other vegetation characteristics at nest sites were reported by [52,69]. In a managed hardwood forest in South Carolina, cover of low vegetation (0-5 feet (0-1.5 m)) was significantly (P<0.01) greater at nest sites (57%) than at random sites within Swainson's warbler territories (40%). In contrast, vegetation cover at the highest stratum (5-8 feet (1.5-2.5 m)) was significantly (P<0.01) lower at nest sites (3%) compared to unused sites (13%). Overall, nest sites typically occurred in areas with 20% to 30% ground cover, 60% to 65% cover at 3 to 5 feet (1-1.5 m) above ground, and 65% to 75% cover at 5 to 8 feet (1.5-2.5 m) above ground [52]. In 1993, nest sites (n=17) were in areas with significantly (P=0.009) taller cane (10 feet (3.1 m)), lower (P=0.047) grass cover (6.9%), and higher (P=0.051) leaf litter coverage (58.4%) than random sites (cane height = 9 feet (2.7 m), grass cover = 15.4%, leaf litter cover = 49.2%) in canebrakes of southern Missouri [69].

Foraging: Swainson's warbler foraging habitat may be less dense than nesting habitat at and just above ground level [3,25,27,43]. Swainson's warblers in 70 locations throughout the southeast tended to forage in vine tents and shady glades less of than 430 feetē (40 mē) [25]. In addition, a review notes use of small <50 feetē (5 mē) areas with few obstructions near ground level by foraging male Swainson's warblers [43]. However, the ground foliage densities on foraging sites in northeastern Louisiana were in accordance with availability [3].

It is possible that foraging occurs in more interior locations within a stand. Swainson's warblers were not observed foraging on the forest edge in Jamaica [36].

FOOD HABITS:
Swainson's warblers forage on and near the ground. In northeastern Louisiana, Swainson's warblers had a mean foraging height of 1.3 feet (0.4 m). This was the lowest relative foraging height observed and included the most occurrences of a species foraging on fallen debris. Of 17 foraging observations, 71% were on the ground and the remainder were in shrubs [3]. In Jamaica, Swainson's warblers captured 95% of insects on the ground [36].

Swainson's warbler's diet is comprised almost entirely of arthropods. In the southeast, crickets (Gryllidae), ground beetles (Carabidae), ants (Formicidae), and spiders (Araneae) are the primary prey items of Swainson's warblers. Caterpillers (Lepidoptera), stinkbugs (Pentatomidae), and grasshoppers (Acrididae) have also been reported as major food items [43]. Less often Swainson's warblers eat millipedes (Diplopoda), beetle larvae, and moths (Lepidoptera) [25,43]. In Jamaica, Swainson's warblers ate insects in all (n=32) foraging observations [36]. Regurgiation samples from Swainson's warblers in Jamaica were comprised primarily of beetles (Coleoptera), spiders, and ants. In addition to the typical prey items, seeds and a species of gecko (Sphaerodactylus goniorhynchus) were also eaten. See the table below for occurrence and frequency of prey items.

Prey in regurgitation samples of 13 Swainson's warblers in Jamaica [64]
Prey Type

Number
(% of total)

Frequency
(% of birds)

Formicidae 51 (19) 77
Seeds 7 (3) 31
Coleoptera 103 (39) 100
Larvae 10 (4) 54
Snails 1 (<1) 8
Spiders 58 (22) 100
Hymenoptera 1 (<1) 8
Orthoptera 13 (5) 62
Dermaptera 5 (2) 31
Mites 2(1) 15
Hemiptera 3 (1) 15
Pseudoscorpion 3 (1) 23
Gecko 9 (3) 69
Unknown 1 (<1) 8


 

Steve Nanz, http://www.stevenanz.com/

Swainson's warblers forage primarily by using their bills to search through and capture prey in loose litter. For example, in northeast Louisiana sweeping motions of the bill through ground leaf litter was used in 63% of foraging observations (n=41 observations, n=17 Swainson's warblers). Gleaning insects off of the ground and low shrubs occurred in 32% of observations. Only 5% of observations were of flying Swainson's warblers capturing prey off a substrate [3]. Of 399 individuals observed in over 70 locations only 2 used foraging methods other than the "leaf-lifting" technique. The author suggests observations of arboreal gleaning by Swainson's warblers are of agitated individuals [25]. A review of the foraging habits of the Swainson's warbler is included in [7].

PREDATORS:
There are few confirmed Swainson's warbler predators. No predation of adults has been reported. Nests are likely predated by several snakes, birds, and mammals [7,43,63,66]. Given their occurrence near the ground, nests would be susceptible to both mammal and snake predation [43,63]. Predation of a nest near Macon, Georgia was attributed to a mouse [43]. A review lists nest predation by the common milksnake (Lampropeltis triangulum) and possible predation by common grackles (Quiscalus quiscula) and blue jays (Cyanocitta cristata) [7].

According to a review, Swainson's warbler nests are parasitized by brown-headed cowbirds (Molothrus ater) throughout their breeding range [7]. The extent to which brown-headed cowbird parasitism affects Swainson's warbler is uncertain, with both rather high [43] and low rates [52] of parasitism reported. It has been suggested that Swainson's warbler's habit of nesting comparatively late in the spring prevents much brown-headed cowbird parasitism [44].

MANAGEMENT CONSIDERATIONS:
The Swainson's warbler is considered vulnerable in many portions of its range. Partners in Flight consider Swainson's warbler a priority species [21,49,71]. Their range has apparently receded in Oklahoma [28] and Delaware [7]. In addition, Swainson's warbler populations in the Blue Ridge mountains may be declining. Populations may be increasing in some areas, such as the South Atlantic Coastal Plain [32,57]. However these estimates are limited by small sample size, low power to detect population trends, and/or low abundance [7,57].

Playback of Swainson's warbler songs is an effective censusing technique [23,24].

Methods of establishing and/or maintaining cane are addressed in [18,53].

Although demographic analysis in varying habitats and landscapes has not been performed [4], Swainson's warblers may benefit from some timber harvesting. Habitat characteristics of foraging sites in Tensas River National Wildlife Refuge in Louisiana led to the conclusion that selective cutting with rotations ≥100 years would have a positive effect on Swainson's warblers [3]. Small (0.6 acre (0.25 ha)), patchy clearcuts (25/kmē) were recommended as a way to promote Swainson's warbler establishment in marginal habitat in Virginia [27]. In South Carolina, a viable Swainson's warbler population occurred in large (741-1,730 acres (300-700 ha)), approximately 15-year-old clearcuts where some lowland areas were left intact [52]. The degree of fragmentation and size of clearcuts at which impacts to Swainson's warbler become detrimental are uncertain [4]. Restricting harvesting to the nonbreeding season (1 October to 1 April) is likely to minimize negative impacts to Swainson's warblers [4,18]. A more detailed review of the impact of various management techniques on Swainson's warblers is provided by [7] and water management issues are discussed in [4,27].


FIRE EFFECTS AND USE

SPECIES: Limnothlypis swainsonii
DIRECT FIRE EFFECTS ON ANIMALS:
Although there are no data on the direct impact of fire on Swainson's warblers, adults are likely much less vulnerable than eggs and young birds. It is generally accepted that adult birds typically have the mobility to avoid fire, although large, fast-moving fires can result in mortality [12,17,39,54]. In contrast, the relatively low placement of Swainson's warblers nests (see Nesting cover) suggests that even low-severity surface fires during the breeding season could result in considerable nest mortality [17,39,55]. A severe reduction in nest success due to flooding to about 6 feet (2 m) was observed in a floodplain forest of Georgia [43]. The degree to which a population would be affected by fire would depend on several factors including occurrence of renesting, timing of the fire, fire interval, fire uniformity, and fire severity [39,55]. Although Swainson's warblers have a tendency to renest after failed attempts (see Timing of Major Life History Events), the extent to which this could mitigate the impact of a breeding-season fire is unknown.

HABITAT-RELATED FIRE EFFECTS:
Very little information is available on the effects of fire on Swainson's warblers. The available information is comprised mainly of Swainson's warbler abundances on sites with varying fire histories and does not include information at varying temporal and spatial scales. There is also a lack of information regarding the effects of various site and fire characteristics on Swainson's warblers' response to burning, and as of early 2007 there were no comparisons of Swainson's warbler demographic data from burned and unburned areas or experiments with replicates and/or controls. The majority of the information that follows is speculation based on Swainson's warblers' habitat and cover requirements.

Swainson's warblers have been observed in recently burned areas. Swainson's warblers were transients in fire-suppressed longleaf pine (Pinus palustris) stands, but did not occur in fire-maintained stands, which had significantly (P<0.05) lower percent canopy cover (62.78% vs. 77.29% on fire-suppressed sites), mean number of small trees (4.35 vs. 14.28 on fire-suppressed sites), and mean number of large saplings (2.56 vs. 4.71 on fire-suppressed sites) [34]. Swainson's warblers were observed in 4.3% of surveys performed on a grass-forb regeneration site in South Carolina that had been cut and burned the previous year, and they occurred in 15% of censuses in shrub-scrub regeneration sites that were cut and burned 2 to 5 years previously [37].

Several factors are likely to influence Swainson's warblers response to fire, including fire season, frequency, uniformity, severity and site characteristics.

Burning during the nonbreeding season will probably reduce impacts of fire on Swainson's warbler survival and reproduction. Recommendations regarding timing of silvicultural treatments include scheduling disturbances from October 1 to April 1 [4,18].

Frequent fires are likely to have a negative impact on Swainson's warbler. Swainson's warblers were observed on sites in North Carolina that were not burned in the growing season and had relatively few dormant season burns, but were not seen on sites maintained with at least 1 prescribed fire in the growing season and 5-year dormant season fire intervals [34]. A similar trend occurred in southwestern Mississippi [10]. In contrast, regular fires in canebrakes at intervals of about 4 to 10 years [30,38] may help maintain Swainson's warbler habitat.

Given the possibility of a preference for patchy habitats (see Landscape factors), Swainson's warbler may have a better response to small and/or patchy fires compared to large and/or uniform burns. Small (0.6 acre (0.25 ha)), patchy clearcuts (25/kmē) were recommended as a way to promote establishment in marginal habitat in Virginia [27] and small (<1.2 acres (<0.5 ha)) group selection cuts were recommended on sites in Arkansas [4].

The influence of fire severity would depend in part on the time required for necessary habitat components to recover. Given their occurrence on sites with thick cover, Swainson's warbler would not be expected to occur in areas after severe fires. However, in some cases it is possible that stand-replacing fire could benefit Swainson's warbler by opening the canopy and allowing the development of a dense understory. Mixed-severity fires could potentially create a patchy mosaic of open and closed habitats, which may benefit Swainson's warblers (see Landscape factors), as well as thin out dense canopy cover. Less severe fires could improve understory cover by simulating sprouting of rhizomatous understory species, such as cane [4,31,58]. However, even low-severity surface fires could consume leaf litter [48] and nesting habitat. The impact this would have on Swainson's warbler is unknown, but it would likely be influenced by the availability of these features in neighboring areas and the time necessary for vegetation regeneration, leaf litter accumulation, and prey species recovery. Depending on fire characteristics, understory vegetation could recover quickly. For instance, cane can reach preburn heights within 2 growing seasons of burning [31,58]. Several other species Swainson's warblers use for nesting and/or cover sprout following fire, including mountain laurel, coastal sweetpepperbush and common and laurelleaf greenbrier. In some circumstances, prey species may also be able to recover in a relatively short time. In an oak-pine (Quercus-Pinus) forest in Kentucky, arthropod communities began recovering 2 years after prescribed burning [14], and some beetles were not substantially affected by prescribed burning in mixed-oak forests of Ohio [60,62]. However, fires can cause large [6,14], potentially long-term [9] changes in the forest floor arthropod community.

Fire Ecology: Swainson's warblers occur in habitats with a wide range of fire regimes. Canebrakes can be maintained with low severity fires about every 4 to 10 years [30,38]. See the FEIS review of cane for more details regarding fire regimes of forests where cane occurs and the effects of fire on cane. Although little is known about presettlement fire regimes in many Swainson's warbler habitats including oak-hickory and bottomland forests, a literature review [70] summarizes the information available and suggests that fire-return intervals in bottomland forests were from <35 to 200 years and that fires were typically of low severity due to rapid decomposition on these moist sites resulting in low fuel loadings. Stand replacing fires at intervals exceeding 1,000 years occur in most maple-beech-birch (Acer-Fagus-Betula spp.) communities [70].

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

Community or Ecosystem Dominant Species Fire-Return Interval Range (years)
maple-beech Acer-Fagus spp. 684-1,385 [13,70]
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
sugarberry-America elm-green ash Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica <35 to 200
Atlantic white-cedar Chamaecyparis thyoides 35 to >200
beech-sugar maple Fagus spp.-Acer saccharum >1,000 [70]
green ash Fraxinus pennsylvanica <35 to >300 [19,70]
yellow-poplar Liriodendron tulipifera <35
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 [70]
longleaf-slash pine Pinus palustris-P. elliottii 1-4 [50,70]
pitch pine Pinus rigida 6-25 [8,29]
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 [70]
eastern cottonwood Populus deltoides <35 to 200 [51]
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 [70]
oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to >200 [50]
southeastern oak-pine Quercus-Pinus spp. <10
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra <35
bear oak Quercus ilicifolia <35
chestnut oak Quercus prinus 3-8
northern red oak Quercus rubra 10 to <35
black oak Quercus velutina <35
live oak Quercus virginiana 10 to<100 [70]
interior live oak Quercus wislizenii <35 [2]
cabbage palmetto-slash pine Sabal palmetto-Pinus elliottii <10 [50,70]
baldcypress Taxodium distichum var. distichum 100 to >300
pondcypress Taxodium distichum var. nutans <35 [50]
eastern hemlock-yellow birch Tsuga canadensis-Betula alleghaniensis 100-240 [67,70]
eastern hemlock-white pine Tsuga canadensis-Pinus strobus x=47 [13]

FIRE USE:
There is a large amount of uncertainty regarding Swainson's warblers' response to fire, and much more data are needed to make recommendations. The data currently available suggest small and/or low-severity fires during the nonbreeding season would probably have the least detrimental impact on Swainson's warblers. Certain communities important to Swainson's warblers, such as canebrakes, are fire dependent [31,58], and low-severity burns in these areas may create or improve this type of Swainson's warbler habitat [4]. Burning intervals of 2 to 3 years appear incompatible with Swainson's warbler habitats [10,30,38], likely due to a lack of understory cover and/or litter in these areas. Appropriate fire frequencies depend in a large part on the response of vegetation and litter arthropod community to burning.

Limnothlypis swainsonii: REFERENCES


1. American Ornithologists' Union. 2007. The A.O.U. check-list of North American birds, 7th edition, [Online]. American Ornithologists' Union (Producer). Available: http://www.aou.org/checklist/index.php3. [50863]
2. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. [36984]
3. Barrow, Wylie Cark. 1990. Ecology of small insectivorous birds in a bottomland hardwood forest. Baton Rouge, LA: Louisiana State University. 212 p. Dissertation. [63884]
4. Bednarz, James C.; Stiller-Krehel, Petra; Cannon, Brian. 2005. Distribution and habitat use of Swainson's warblers in eastern and northern Arkansas. In: Ralph, C. John; Rich, Terrell D., eds. Bird conservation implementation and integration in the Americas: proceedings of the 3rd international Partners in Flight conference. Vol. 1; 2002 March 20-24; Asilomar, CA. Gen. Tech. Rep. PSW-GTR-191. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 576-588. [63743]
5. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]
6. Bess, Emilie C.; Parmenter, Robert R.; McCoy, Stacey; Molles, Manuel C., Jr. 2002. Responses of a riparian forest-floor arthropod community to wildfire in the middle Rio Grande valley, New Mexico. Environmental Entomology. 31(5): 774-784. [46990]
7. Brown, R. E.; Dickson, J. G. 1994. Swainson's warbler--Limnothlypis swainsonii. In: Poole, A.; Gill, F., eds. The birds of North America. No. 126. Philadelphia, PA: The Academy of Natural Sciences; Washington, DC: The American Ornithologists' Union: 1-20. [65356]
8. Buchholz, Kenneth; Good, Ralph E. 1982. Density, age structure, biomass and net annual aboveground productivity of dwarfed Pinus rigida Moll. from the New Jersey Pine Barren Plains. Bulletin of the Torrey Botanical Club. 109(1): 24-34. [8639]
9. Buddle, Christopher M.; Langor, David W.; Pohl, Greg R.; Spence, John R. 2006. Arthropod responses to harvesting and wildfire: implications for emulation of natural disturbance in forest management. Biological Conservation. 128(3): 346-357. [60653]
10. Burger, Loren W., Jr.; Hardy, Carol; Bein, Jeff. 1998. Effects of prescribed fire and midstory removal on breeding bird communities in mixed pine-hardwood ecosystems. In: Pruden, Teresa L.; Brennan, Leonard A., eds. Fire in ecosystem management: shifting the paradigm from suppression to prescription: Proceedings, Tall Timbers fire ecology conference; 1996 May 7-10; Boise, ID. No. 20. Tallahassee, FL: Tall Timbers Research Station: 107-113. [35612]
11. Carrie, N. Ross. 1996. Swainson's warblers nesting in early seral pine forests in east Texas. The Wilson Bulletin. 108(4): 802-804. [62501]
12. Chandler, Craig; Cheney, Phillip; Thomas, Philip; Trabaud, Louis; Williams, Dave. 1983. Fire in forestry: Vol. I. Forest fire behavior and effects. New York: John Wiley & Sons. 450 p. [12241]
13. Cleland, David T.; Crow, Thomas R.; Saunders, Sari C.; Dickmann, Donald I.; Maclean, Ann L.; Jordan, James K.; Watson, Richard L.; Sloan, Alyssa M.; Brosofske, Kimberley D. 2004. Characterizing historical and modern fire regimes in Michigan (USA): a landscape ecosystem approach. Landscape Ecology. 19: 311-325. [54326]
14. Coleman, T. W.; Rieske, L. K. 2006. Arthropod response to prescription burning at the soil-litter interface in oak-pine forests. Forest Ecology and Management. 233(1): 52-60. [64057]
15. de Cruz, Justine B. 1996. Bird-habitat relations in regenerating forests of eastern North Carolina and Izabal, Guatemala. Durham, NC: Duke University. 345 p. Dissertation. [65645]
16. Dickson, James G. 1978. Forest bird communities of the bottomland hardwoods. In: DeGraaf, Richard M, technical coordinator. Proceedings of the Workshop Management of Southern Forests for Nongame Birds; 1978 January 24 - January 26; Atlanta, GA. Gen. Tech. Rep. SE-14. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 66-73. [17949]
17. Dickson, James G. 2002. Fire and bird communities in the South. In: Ford, W. Mark; Russell, Kevin R.; Moorman, Christopher E., eds. The role of fire in nongame wildlife management and community restoration: traditional uses and new directions: Proceedings of a special workshop; 2000 December 15; Nashville, TN. Gen. Tech. Rep. NE-288. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 52-57. [41557]
18. Eddleman, William R.; Evans, Keith E.; Elder, William H. 1980. Habitat characteristics and management of Swainson's warbler in southern Illinois. Wildlife Society Bulletin. 8(3): 228-233. [62504]
19. Eggler, Willis A. 1980. Live oak. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 63-64. [49984]
20. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
21. Fitzgerald, Jane A.; True, C. Diane; Diamond, David D.; Ettel, Troy; Moore, Laurel; Nigh, Timothy A.; Vorisek, Shawchyi; Wathen, Greg. 2005. Delineating focus areas for bird conservation in the central hardwoods bird conservation region. In: Ralph, C. John; Rich, Terrell D., eds. Bird conservation implementation and integration in the Americas: proceedings of the 3rd international Partners in Flight conference. Vol. 1; 2002 March 20-24; Asilomar, CA. Gen. Tech. Rep. PSW-GTR-191. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 192-202. [63710]
22. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]
23. George, William G. 1972. Breeding status of the purple gallinule, brown creeper, and Swainson's warbler in Illinois. The Wilson Bulletin. 84(2): 208-210. [62505]
24. Graves, Gary R. 1996. Censusing wintering populations of Swainson's warblers: surveys in the Blue Mountains of Jamaica. The Wilson Bulletin. 108(1): 94-103. [62507]
25. Graves, Gary R. 1998. Stereotyped foraging behavior of the Swainson's warbler. Journal of Field Ornithology. 69(1): 121-127. [62509]
26. Graves, Gary R. 2001. Factors governing the distribution of Swainson's Warbler along a hydrological gradient in Great Dismal Swamp. The Auk. 118(3): 650-664. [62510]
27. Graves, Gary R. 2002. Habitat characteristics in the core breeding range of the Swainson's warbler. The Wilson Bulletin. 114(2): 210-220. [62511]
28. Heck, Berlin A. 2001. History and status of the Swainson's warbler in Oklahoma. Bulletin of the Oklahoma Ornithological Society. 34(2): 5-11. [62512]
29. Hendrickson, William H. 1972. Perspective on fire and ecosystems in the United States. In: Fire in the environment: Symposium proceedings; 1972 May 1-5; Denver, CO. FS-276. [Washington, DC]: U.S. Department of Agriculture, Forest Service: 29-33. In cooperation with: Fire Services of Canada, Mexico, and the United States; Members of the Fire Management Study Group; North American Forestry Commission; FAO. [17276]
30. Hughes, Ralph H. 1957. Response of cane to burning in the North Carolina coastal plain. Bulletin No. 402. Raleigh, NC: North Carolina State College, Agricultural Experiment Station. 24 p. [34716]
31. Hughes, Ralph H. 1966. Fire ecology of canebrakes. In: Proceedings, 5th annual Tall Timbers fire ecology conference; 1966 March 24-25; Tallahassee, FL. No. 5. Tallahassee, FL: Tall Timbers Research Station: 148-158. [16236]
32. Hunter, William C.; Pashley, David N.; Escano, Ronald E. F. 1993. Neotropical migratory landbird species and their habitats of special concern within the Southeast Region. In: Finch, Deborah M.; Stangel, Peter W., eds. Status and management of neotropical migratory birds: Proceedings; 1992 September 21-25; Estes Park, CO. Gen. Tech. Rep. RM-229. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 159-171. [65648]
33. Kirkconnell, Arturo; Wallace, George E.; Garrido, Orlando H. 1996. Notes on the status and behavior of the Swainson's warbler in Cuba. The Wilson Bulletin. 108(1): 175-178. [62514]
34. Krieger, Sharlene Marie. 1997. Abundance, habitat associations, and nest success of passerines within a fire maintained longleaf pine (Pinus palustris) ecosystem. Raleigh, NC: North Carolina State University. 78 p. Thesis. [62074]
35. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. [3455]
36. Lack, David; Lack, Peter. 1972. Wintering warblers in Jamaica. The Living bird. 11: 129-153. [63887]
37. Lanham, Joseph Drew. 1997. Attributes of avian communities in early-successional, clearcut habitats in the mountains and Upper Piedmont of South Carolina. Clemson, SC: Clemson University. 126 p. Dissertation. [60756]
38. Leithead, Horace L.; Yarlett, Lewis L.; Shiflet, Thomas N. 1971. 100 native forage grasses in 11 southern states. Agric. Handb. 389. Washington, DC: U.S. Department of Agriculture, Forest Service. 216 p. [17551]
39. Lyon, L. Jack; Telfer, Edmund S.; Schreiner, David Scott. 2000. Direct effects of fire and animal responses. In: Smith, Jane Kapler, ed. Wildland fire in ecosystems: Effects of fire on fauna. Gen. Tech. Rep. RMRS-GTR-42-vol. 1. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 17-23. [44435]
40. Meanley, Brooke. 1966. Some observations on habitats of the Swainson's warbler. The Living Bird. 5: 151-165. [62523]
41. Meanley, Brooke. 1969. Pre-nesting and nesting behavior of the Swainson's warbler. The Wilson Bulletin. 81(3): 246-257. [62522]
42. Meanley, Brooke. 1971. Additional notes on pre-nesting and nesting behavior of the Swainson's warbler. The Wilson Bulletin. 83(2): 194. [62518]
43. Meanley, Brooke. 1971. Natural history of Swainson's warbler. North American Fauna No. 69. Washington, DC: U.S. Department of the Interior, Bureau of Sport Fish and Wildlife. 90 p. [62520]
44. Meanley, Brooke. 1982. Swainson's warbler and the cowbird in the Dismal Swamp. The Raven. 53(3): 47-49. [62524]
45. Meyers, J. Michael; Wright, Elizabeth A. 2003. Models for managing habitat of a Swainson's warbler breeding population, Bond Swamp National Wildlife Regufe, Georgia, [Online]. In: Products: research showcase. Laurel, MD: U.S. Department of the Interior, U.S. Geological Survey, Patuxent Wildlife Research Center (Producer). Available: http://www.pwrc.usgs.gov/resshow/meyers/meyers1.htm [2007, January 26]. [65351]
46. Miller, Stanlee M. 2003. First report of a double-brooded Swainson's warbler. The Wilson Bulletin. 115(1): 94-95. [62525]
47. Mitchell, Michael S.; Lancia, Richard A.; Gerwin, John A. 2001. Using landscape-level data to predict the distribution of birds on a managed forest: effects of scale. Ecological Applications. 11(6): 1692-1708. [62496]
48. Moseley, Kurtis R.; Castleberry, Steven B.; Schweitzer, Sara H. 2003. Effects of prescribed fire on herpetofauna in bottomland hardwood forests. Southeastern Naturalist. 2(4): 475-486. [62076]
49. Mueller, Allan J.; Dikeman, Hayley M.; Edwards, Thomas L.; Holbrook, Robert S.; Rowe, Karen L. 2005. Integrated migratory bird planning in a corps of engineers irrigation project, Bayou Meto, Arkansas. In: Ralph, C. John; Rich, Terrell D., eds. Bird conservation implementation and integration in the Americas: proceedings of the 3rd international Partners in Flight conference. Vol. 1; 2002 March 20-24; Asilomar, CA. Gen. Tech. Rep. PSW-GTR-191. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 86-92. [63519]
50. Myers, Ronald L. 2000. Fire in tropical and subtropical ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 161-173. [36985]
51. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; Gottfried, Gerald J.; Haase, Sally M.; Harrington, Michael G.; Narog, Marcia G.; Sackett, Stephen S.; Wilson, Ruth C. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. [36978]
52. Peters, Kimberly A.; Lancia, Richard A.; Gerwin, John A. 2005. Swainson's warbler habitat selection in a managed bottomland hardwood forest. Journal of Wildlife Management. 69(1): 409-417. [62526]
53. Platt, Steven G.; Brantley, Christopher G. 1993. Switchcane: Propagation and establishment in the southeastern United States. Restoration & Management Notes. 11(2): 134-137. [22802]
54. Quinn, Ronald D. 1994. Animals, fire and vertebrate herbivory in Californian chaparral and other Mediterranean-type ecosystems. In: Moreno, Jose M.; Oechel, Walter C., eds. The role of fire in Mediterranean-type ecosystems. New York: Springer Verlag: 46-78. [26804]
55. Robbins, Louise E.; Myers, Ronald L. 1992. Seasonal effects of prescribed burning in Florida: a review. Misc. Publ. No. 8. Tallahassee, FL: Tall Timbers Research, Inc. 96 p. [21094]
56. Ruelas Inzunza, Ernesto; Hoffman, Stephen W.; Goodrich, Laurie J. 2005. Stopover ecology of neotropical migrants in central Veracruz, Mexico. In: Ralph, C. John; Rich, Terrell D., eds. Bird conservation implementation and integration in the Americas: proceedings of the 3rd international Partners in Flight conference: Vol. 2; 2002 March 20-24; Asilomar, CA. Gen. Tech. Rep. PSW-GTR-191. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 657-672. [63459]
57. Sauer, J. R.; Hines, J. E.; Fallon, J. 2005. Red-headed woodpecker Melanerpes erythrocephalus: North American breeding bird survey trend results. In: The North American breeding bird survey, results and analysis 1966-2005. Version 6.2.2006, [Online]. U.S. Department of the Interior, Geological Survey, Patuxent Wildlife Research Center, Migratory Bird Research (Producer). Available: http://www.mbr-pwrc.usgs.gov/bbs/bbs.html [2007, January 29]. [65354]
58. Shepherd, W. O.; Dillard, E. U.; Lucas, H. L. 1951. Grazing and fire influences in pond pine forests. Tech. Bull. No. 97. Raleigh, NC: North Carolina State College, Agricultural Experiment Station. 56 p. In cooperation with: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. [14546]
59. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. [23362]
60. Smith, A. E.; Horn, D. J. 2000. Monitoring scarab populations exposed to prescribed fire in mixed-oak forests of southeast Ohio. In: Yaussy, Daniel A., compiler. Proceedings: workshop on fire, people, and the central hardwoods landscape; 2000 March 12-14; Richmond, KY. Gen. Tech. Rep. NE-274. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 129. Abstract. [40445]
61. Somershoe, Scott G.; Hudman, Stephen P.; Chandler, C. Ray. 2003. Habitat use by Swainson's warblers in a managed bottomland forest. The Wilson Bulletin. 115(2): 148-154. [62527]
62. Stanton, Chris; Purrington, Foster F.; Horn, David J. 2000. Ground beetle responses to prescribed burning in southern Ohio's hardwood forests. In: Yaussy, Daniel A., compiler. Proceedings: workshop on fire, people, and the central hardwoods landscape; 2000 March 12-14; Richmond, KY. Gen. Tech. Rep. NE-274. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 129. Abstract. [40446]
63. Stevenson, Henry M.; Anderson, Bruce H. 1994. The birdlife of Florida. Gainesville, FL: University of Florida Press. 892 p. [60776]
64. Strong, Allan M. 2000. Divergent foraging strategies of two neotropical migrant warblers: implications for winter habitat use. The Auk. 117(2): 381-392. [62528]
65. Strong, Allen M.; Sherry, Thomas W. 2001. Body condition of Swainson's warblers wintering in Jamaica and the conservation value of Caribbean dry forests. The Wilson Bulletin. 113(4): 410-418. [62529]
66. Summerour, C. W. 1979. Nesting of the Swainson's warbler in Alabama. Alabama Birdlife. 27(1/2): 6-10. [62531]
67. Swain, Albert M. 1978. Environmental changes during the past 2000 years in north-central Wisconsin: analysis of pollen, charcoal, and seeds from varved lake sediments. Quaternary Research. 10: 55-68. [6968]
68. Tappe, Philip A.; Thill, Ronald E.; Melchiors, M. Anthony; Wigley, T. Bently. 2004. Breeding bird communities on four watersheds under different forest management scenarios in the Ouachita Mountains of Arkansas. In: Guldin, James M., tech. comp. Ouachita and Ozark Mountains symposium: ecosystem management research; [Meeting date unknown]; [Location unknown]. Gen. Tech. Rep. SRS-74. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station: 154-163. [55739]
69. Thomas, Brian G.; Wiggers, Ernie P.; Clawson, Richard L. 1996. Habitat selection and breeding status of Swainson's warblers in southern Missouri. Journal of Wildlife Management. 60(3): 611-616. [62532]
70. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; Grace, James B.; Hoch, Greg A.; Patterson, William A., III. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. [36983]
71. Watson, Craig; Hayes, Chuck; McCauley, Joseph; Milliken, Andrew. 2005. The South Atlantic Migratory Bird Initiative - an integrated approach to conservation of "all birds across all habitats". In: Ralph, C. John; Rich, Terrell D., eds. Bird conservation implementation and integration in the Americas: proceedings of the 3rd international Partners in Flight conference. Vol. 1; 2002 March 20-24; Asilomar, CA. Gen. Tech. Rep. PSW-GTR-191. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 266-276. [63712]

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