Brown-headed Nuthatch Sitta pusilla
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
Breeding Bird Survey
AL
MS
LA
TX
OK
AR
State
(NatureServe)
VA TN
NC MD
SC DE
GA
FL
Physiographic
Province
Coastal Plain
Piedmont
Highlands
LANDFIRE
MapZone
37 44 45 46 48 54 55 56
57 58 59 60 98 99
Southeast Region Version of map better
The brown-headed nuthatch (BHNU) is an endemic, resident (non-migratory) breeding, insect and seed eating bird of mature, open-canopied, southeastern pine forest (Engstrom et al 1984, Connor and Dickson 1997, The Nature
Conservancy 2000, Dornak et al 2004, Barnhill and Imm 2006, Cox and Slater 2007, Lower MS Joint Venture 2009, Tall
Timbers Research Station 2009). Preferred BHNU habitat consists of mature pine forests with open understory and snags
( i.e
. standing dead trees) suitable for the excavation of cavity nests (Wilson and Watts 1999, Lloyd and Slater 2006, Haas
2007). The BHNU forages for insects almost exclusively (>98% of observations) in live and dead overstory pines
(Withgott and Smith 1998, Barnhill and Imm 2006) of pine and pine-hardwood forest and woodland. Deciduous forest apparently does not support nuthatches (Engstrom et al 1984, The Nature Conservancy 2000, Connor et al 2002). A small
BHNU breeding population on the urbanized campus of Stephen F. Austin University in eastern TX used pines 82% (1.75
X availability) of the time while using hardwoods just 18% (0.34 X availability) of the time (Herb and Burt 2000). Time spent in trees was dominated by foraging on bark (50.4% for pines, 54.2% for hardwoods), or pine cone (16.9%) substrates in the branches of the upper 2/3’s of these trees. No BHNU were detected in 9-10 year-old closed-canopy, commercial loblolly pine plantations studied in eastern NC (Wilson and Watts 1999). BHNU across GA piedmont forest sites strongly favored mature (60+ year old) open pineland (mean = 11.3 BHNU/50 ha; SE = ±2.28), used dense commercial (20-30 years old) loblolly pine plantations only rarely (mean = 0.007 BHNU/50 ha; SE = ±0.007), and were entirely absent from upland hardwood forest sites (White et al 1996) in winter.
Increased habitat openness promoted winter BHNU abundance in longleaf pine sand hills habitats in the FL panhandle
(Provencher et al 2002a). BHNU was identified as one of four strong and consistent avian species contributing to proportional similarity and endpoint difference between hardwood reduction and [desired “open” condition] reference plots (Provencher et al 2002b). BHNU were most abundant at upland, open (fire treated) locations in longleaf pine sand hills on Fort Bragg, NC, where BHNU presence was: positively associated with grass cover and the number of longleaf pine trees; and negatively with leaf litter at the microhabitat (50m radius), territory scale (Allen 2001, and Allen et al
2006). BHNU preferred open pine stands with a mean pine basal area of 5.6 m 2 /ha (O'Halloran and Conner 1987, Dornak
2004) with few hardwoods (≤17.4 stems/ha and basal area ≤5 m 2 /ha) and an open midstory (Wilson and Watts 1999).
BHNU point counts in commercial loblolly pine plantations in eastern NC (Wilson and Watts 1999) found abundance was negatively correlated with canopy cover, hardwood density, and basal area of hardwoods and positively correlated with groundcover density. No BHNU were detected in forest patches prior to a first commercial thinning while detections were greatest in the year immediately following thinning, declining with time after thinning. BHNU density was not significantly correlated with canopy height, pine density or pine basal area. Wilson et al (1995) compared untreated with restoration treated (overstory thinning, hardwood reduction and prescribed burning, favoring herb-forb groundcover) shortleaf pine-bluestem stands in western AR. BHNU were present in all restoration treatments but entirely absent from untreated plots. Optimal canopy closure is variable (15-85 percent), but closed canopies are avoided (O’Halloran and
Conner 1987, Wilson and Watts 1999, Dornak 2004, Engstrom et al 1984).
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
4/10/2020
Brown-headed Nuthatch Sitta pusilla
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
Specific pine species composition was not as critical as d.b.h., with an average d.b.h. of 25.6 cm optimal (O’Halloran and
Conner 1987). At Stephen F. Austin University in east TX, 47% of available trees campus-wide were pine and 53% hardwoods (Herb and Burt 2000). Within territories, mean total tree density was 39.15 trees/ha (range = 23.8-75.7 trees/ha), with 57% (22.3 trees/ha; ranges = 13.5-46.9 trees/ha and 45-60%) pine and 43% (16.8 trees/ha; ranges =10.2-
28.8 trees/ha and 36-55%) hardwood stems. Territories with relatively dense pine composition provide more foraging substrate and may allow increased foraging efficiency and reproductive success. Greater abundances of BHNU within 20 breeding cluster sites managed for red-cockaded woodpeckers (RCW’s) were detected during both breeding and wintering seasons than in untreated control sites in pine cover types in east TX (Connor et al 2002). RCW management restored a more open park-like condition, and t he greater BHNU abundance within managed sites was attributed to the species apparent avoidance of hardwood vegetation. Hardwood basal area was 97% less (mean = 0.1 m 2 /ha, SD = 0.3) in woodpecker cluster areas (mean = 3.6-4.9, m 2 /ha SD = 2.4-3.7) and hardwood midstory stem-density, also tended to be less (88-100%, mean = 0.1-0.2 stems/ha, SD = 0.0-0.3) in RCW cluster areas than in untreated pine forest sites (mean =
4.3-5.9 stems/ha, SD = 1.3-4.8). Woody shrub layer vegetation was 20-140% more abundant in cluster areas (mean =
10.3-22.1%, SD = 0.2-5.6) than in control sites (mean = 9.3-13.1%, SD = 0.8-5.2), associated with a 200-300% increase in the grass component (mean = 19.8-27.5 % cover, SD = 22.2-22.6 for RCW clusters vs. mean = 7.0-8.5% cover, SD =
12.7-18.0 for controls), and a concurrent 57-63% decrease in the fern and dicotyledonous component of the ground cover
(mean = 18.0-30.8 % cover, SD = 10.1-18.8 for RCW clusters vs. mean = 48.0-72.5% cover, SD = 23.3-24.2 for controls).
Breeding BHNU abundance did not however differ between native (undisturbed) herbaceous ground cover and more ruderal “old field” herbaceous ground cover with prior soil disturbance within open, mature longleaf pine stands studied in southwest GA, suggesting density or % cover of herbaceous ground cover is more important than ground cover species composition (Rutledge and Conner 2002).
BHNU were absent from 10-year old slash pine plantations during all seasons in northern FL (Repenning and Labisky
1985). During the breeding season, BHNU were also absent from 24-year old slash pine plantations and absent from 1year old slash pine plantations during the winter. While most abundant in both breeding and winter seasons in open, mature longleaf pine forest (45 birds/Km 2 breeding and 19/Km 2 winter) and 40-year old slash pine plantations (33/Km 2 breeding and 17/Km 2 winter), BHNU were also present but with low abundance (7/Km 2 ) in 1-year old, grass-forb dominated slash pine plantations during the breeding season, and in 24-year old slash pine plantations in winter (3/Km 2 ).
In older stands, increases in mid-story hardwoods lead to decreases in BHNU, possibly because the vegetation may obscure potential nest cavity locations (Wilson and Watts 1999, Engstrom et al 1984, Allen et al 2006). Undergrowth is typically sparse (~35 percent) in occupied BHNU habitat (Dornak et al 2004). The LMJV (2009) BHNU habitat suitability index model used an inverse logistic function for small stem density to account for the preference of BHNU for open understories. Frequent fire maintains a sparse midstory and diverse groundcover composition, increasing arthropod biomass for forage (Engstrom et al 1984, Taylor 2003, Barnhill and Imm 2006). The LMJV (2009) BHNU habitat suitability index model incorporated an inverse logistic function for hardwood basal area based upon Wilson and Watts
(1999) data to represent the inverse relationship between hardwood basal area and BHNU abundance.
BHNU characteristically nest in moderately (Harrison 1975, Headstrom 1965) to well-decayed pine or hardwood snags
(Miller and Jones 1999, O'Halloran and Conner 1987, McNair 1984) and show a strong preference for snags blackened by fire (Harrison 1975, Withgott and Smith 1998), lacking crowns, and missing bark (O'Halloran and Conner 1987). BHNU may enlarge an existing cavity, reuse an old one, or use artificial nest boxes or other man-made cavities such as streetlights (The Nature Conservancy 2000, Herb and Burt 2000, LMJV 2009, McNair 1984). BHNU nests occurred in light poles and electrical box in addition to natural cavities in dead portions of live trees on an urbanized east TX college campus (Herb and Burt 2000) where natural cavity sitesavailability was likely at a premium due to snag and dead limb removal and intensive landscaping practices.
BHNU tend to select standing dead, well decayed trees or stumps that are <2.5 m in height (Weiss 2003). Weiss determined that BHNU in a “declining” breeding population in coastal VA were selecting nest sites (n=29 nests) that were higher (mean = 10.1 m; range = 2.2-29.3 m), and 34% (10/29 nests) were in dead portions (branch wounds) of live trees rather than in dead snags/stumps when compared to published historic nest records. BHNU’s were positively correlated with standing snags ( τ = 0.15, n = 144, P < 0.009), and were > 3X more likely to be detected in eastern NC commercial loblolly pine plantation plotss (Wilson and Watts 1999) containing standing snags (12 of 32 plots, 37.5%) compared to plots that did not (13 of 122 plots, 11.6%; X 2 = 7.35, df = 1, P < 0.007). BHNU primarily nest in large d.b.h. snags <3 m tall and may require 7–8 snags/ha to ensure adequate nest and roost sites, particularly in the presence of inter-specific
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
4/10/2020
Brown-headed Nuthatch Sitta pusilla
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project competition for cavities. McNair (1984) determined mean snag and cavity entrance heights were 3.2 (± 0.3) m and 2.2 ( ±
0.2) m respectively, with 92% (22 of 24) of the nest cavities located in the top half of the snags and were not equally distributed on the nest snags (n = 24, X 2 = 16, P < 0.001) using museum records, suggesting that BHNU preferred the top portion of short nest snags. Variation in density of small and large pine trees, density of pine snags, and % cover of hardwood shrubs, fire history, and hydrological conditions at 141 BHNU nest sites affected productivity in the pine rocklands of south FL (Lloyd and Slater 2006). The number of (large) pine snags ≥15 cm dbh surrounding the nest site had a strong positive effect on productivity (model-averaged β = 1.6 fledglings/nest, 95% CI = 0.1–3.0) and the number of small pines surrounding nest sites had a weak positive effect on productivity (model-averaged
β
= 1.3, 95% CI = 0.4–2.2).
McNair (1984) found a modal cavity height for all BHNU nest-site categories was 1.2 m and the median cavity height was
1.5 m, with 90% of all cavity heights recorded <3.66 m. Nest-sites in pine stumps, deciduous stumps, unidentified stumps, posts and nest boxes, with nesting cavities located at mean heights of <3 m were used most frequently. Dornak et al
(2004) monitored 24 BHNU nests, and found nest snags were shorter (3.2 ± 0.3 m) than control snags (12.0 ± 1.8 m), had less dense midstories (density class=1.9 ± 0.1) and less leaf litter (29.3 ± 2.0%) than control sites (density class = 2.5 ± 0.2 and 36.7 ± 2.8%, respectively). A minimum number of snags necessary to support primary cavity-nester populations can be calculated by multiplying the # of individuals desired in a stand able to use each snag size class, by 4 snags/individual
(McComb et al 1986). Where BHNU occur, ≥ 91 snags/40 ha (>12.7 cm) must be present in pine and pine-hardwood stands to maintain “average” population levels for all primary cavity-nesters in FL pineland communities (McComb et al
1986). Only 77snags/40 ha are needed when BHNU are absent (difference = 14 snags/40 ha attributable to BHNU).
Hence ≥212 snags/40 ha should be available for primary cavity-nesters, distributed by size class as follows: 120 snags
≥12.7 cm dbh; 84 snags ≥25.0 cm dbh; and 8 snags ≥50 cm dbh per 40 ha. Additional snags are needed by secondary cavity-nesters, to replace those that fall, and to provide foraging sites that may be necessary to maintain high population densities and to assure optimal survivorship. The LMJV (2009) included snag density in a BHNU habitat suitability index model, assuming a suitability index score of 0 when ≤8 snags of any size were present (Dornak and others 2004).
Based on color-banded adults and nestlings, Cox and Slater (2007) found nests are constructed low in snags in the final stages of decay, i.e. in the softest wood that will last the 33+ days it takes to complete a nesting cycle (Tall Timbers
Research Station 2009). BHNU avoid short rotation pine forests (less than 80 years) because older pine trees often have dead limbs present (Connor and Dickson 1997, Hunter et al. 2001, Barnhill and Imm 2006). In loblolly-shortleaf pine, highest BHNU densities occurred in stands 45-60 years old (Meyers and Johnson 1978). Lloyd and Slater’s (2006) pine rocklands study in south FL found snag density was lowest in areas that had not burned in the previous 5 years, and thus the absence of fire a gradual decline in BHNU productivity was predicted. Fire, an important slash pine mortality agent may be the most effective tool for increasing both snag recruitment (Menges and Deyrup 2001), and favoring the understory conditions the BHNU prefers.
The LMJV (2009) BHNU habitat suitability model testing revealed a positive relationship (r = 0.58; P ≤ 0.001) between average HSI score and mean BHNU abundance across 37 of 88 BBS route subsections within the Central Hardwood and
West Gulf Coastal Plain regions, made even stronger (r = 0.80) when subsections without BHNU detections were removed from analysis. Allen (2001) and Allen et al (2006) presented a multiple linear regression relative abundance model for occupied habitat BHNU at Fort Bragg, NC with good predictive abilities. Lloyd and Slater (2006) also created a 10 candidate model set from linear combinations of variables believed to reasonably explain variation in BHNU productivity based on BHNU ecology knowledge for south FL pine rockland populations.
Landscape/Spatiality-Temporality (Mobility, Area Sensitivity, Insularity-Connectivity Considerations) :
Territories of breeding males in GA were 3.2 hectares (Norris 1958) and estimated winter ranges were 16.7 hectares in LA
(Morse 1970). BHNU mean territory size for (5) BHNU breeding groups studied at an urbanized east TX college campus
(Herb and Burt 2000) was 8.0 ha (range = 4.4-16.2 ha/territory). Nearest neighbor distances average 200 m, suggesting a territory density of 1 territory/7 acres in the south GA and north FL Red Hills region of (Cox and Slater 2007, Tall
Timbers Research Station 2009). Scattered blocks or strips of pine or pine-hardwood can be allowed to mature to provide habitat for the wildlife community associated with BHNU and RCW (McComb et al 1986). Incubation (by the female) lasts 13-15 days and then young are tended by both parents that may be assisted by offspring “helpers” from previous broods. Fledglings leave the nest at about 18 days, but are fed by parents for 24-26 days more (The Nature Conservancy
2000). The best predictor of fledgling production in south FL was the date on which incubation began (Lloyd and Slater
2006), with the model predicting a steep nest productivity decline as the breeding season progressed. Adult BHNU’s are sedentary (high site fidelity) and long-lived (Cox and Slater 2007, Tall Timbers Research Station 2009). Juvenile dispersal spanned short distances and was farther for females than males. First time breeding males often established
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
4/10/2020
Brown-headed Nuthatch Sitta pusilla
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project territories within 300 m of their parent’s territory (i.e., generally the nearest neighboring nest to the natal territory), and this behavior often produced small neighborhoods that of closely related individuals. Suitable habitat within the historic range may remain unoccupied due to fragmentation, complicated by the birds' site tenacity and weak flight (The Nature
Conservancy 2000, Cox and Slater 2007).
Kilgo et al (2002) used US Forest Service stand type, age and condition data and bird point counts to test BIRDHAB performance for predicting BHNU presence-absence for 60 forest stands near Aiken, SC. BIRDHAB tended toward overpredicting BHNU presence (false-presence error rate = 43.9%) but performed well in predicting BHNU absence (2% false absence error rate), yielding a marginally useful 54.1% overall accuracy rate for BHNU presence-absence.
Howell et al (2006) constructed and tested BHNU habitat models for GA, comparing variables from 4 “nested” spatial scales (home range = 5.76 ha; community = 144 ha; Reserve = 3,600 ha; and Region = 90,000 ha). Evergreen forest was an important predictor for BHNU presence at both the home-range (5.76 ha) and community (144 ha) scales, and variables from all 4 spatial scales were selected for all model comparisons correctly classified BHNU presence/absence 68.9% of the time. Landscape attributes (for 3 nested spatial scales: 300 m; 900 m; and 1,500 m radii) were related to distributions for BHNU at Fort Bragg, NC (Allen 2001) and were positively associated with less landscape patch richness density; landscape proportion of open pine habitat, and negatively correlated with increasing upland hardwood landscape proportion and density. The occurrence (presence/absence) of BHNU at Fort Bragg, NC was associated more strongly with landscape than microhabitat features, and BHNU relative abundance indices increased with distance from the drains dissecting the longleaf pine dominated sand hills and suggested BHNU sensitivity to forest fragmentation. Lloyd and
Slater (2006) excluded landscape-level factors (e.g., patch size or distance to habitat edge) that may have influenced productivity of their south FL BHNU populations. Most of the variation in landscape-level features in their two study areas arose from the naturally patchy distribution of plant communities in the ecosystem that they believed was not susceptible to modification by management. Their results may not apply to small and highly fragmented pine rockland patches remaining within the Miami-Dade County network of parks where ongoing surveys indicate that BHNU are absent. BHNU were entirely absent from pine-hardwood forest patches < 10 ha in a northeast GA piedmont study area
(McIntyre’s 1995), suggesting that BHNU abundance increased as pine-hardwood patch size increased above 10 ha.
Withgott and Smith (1998) concluded that area sensitivity does not appear to be an issue for BHNU, which is not an acceptable host for the brown-headed cowbird (LMJV 2009).
Highest BHNU densities and frequencies in western Arkansas occurred in the year immediately following a burn (Wilson et al. 1995), but results from north FL suggested a first-year post-burn peak is followed by a decline and then a higher peak in the fifth post-burn year (Engstrom et al. 1984). BHNU in central GA pine stands were present at 68% (mean =
2.99 detections/count/site; SE = ± 0.81) of breeding season points in mature (>60 year old) pine stands within 3 years after burning, but only at 38% (mean 1.23/count/site; SE = ±0.49) of points in mature pine stands unburned for ≥ 20 years
(White et al 1999). In south FL BHNU populations, 1–2 year fire-return intervals may be useful to restore long-unburned pine rockland and reduce fuel loads, but over the long-term such short fire-return intervals may reduce snag abundance
(Lloyd and Slater 2006). Fires in stands that remain unburned for longer periods (e.g., 6–8 yr) will be significantly hotter and thus recruit the greatest number of new snags (Menges and Deyrup 2001, Platt et al. 2002) and likely will increase snag retention time. The mid- and understory vegetation density increases with fire-return interval. A compromise allowing for spatial and temporal heterogeneity in fire-return interval may be useful to create and maintain high-quality
BHNU habitat. Determining the range of fire-return intervals that optimizes mid- and understory conditions and the number of large snags requires additional information about the role of fire in snag population dynamics, and better understanding of how hydrological conditions and other disturbances (e.g., hurricanes or insect outbreaks) mediate fire effects in pine rocklands (e.g., Lockwood et al. 2003). The appropriate regime for a given area depends upon local climatic conditions, aspect, and site history, affects on the rate of understory regeneration. Data suggest that burns be limited in size to create a mosaic of understory development (Kerpez and Stauffer 1989), and snag retention (Cox and
Slater 2007) but no information is available on the ideal patch size (The Nature Conservancy 2000).
(Specific and Measurable Habitat Parameters)
Mature (≥ 60 years old), large (≥ 25 cm mean dbh), pine-dominated or mixed pine-hardwood (≥ 50% pine) forest
(Engstrom 1984, Repenning and Labisky 1995, White 1996, Withgott and Smith 1998, Wilson and Watts 1999, Herb and
Burt 2000, Conner 2002). Open canopy (25-75% total canopy cover), mid- and understory conditions (≤25 hardwood stems or ≤ 3.0 m 2 /ha hardwood basal area), with abundant herbaceous (≥ 40% grass-forb cover) ground cover (Engstrom
1984, Wilson and Watts 1999, Herb and Burt 2000, Allen 2001, Conner et al 2002, Provencher et al 2002a and 2002b,
Dornak et al 2004, Allen et al 2006). Abundant (≥ 5/ha or 2.5/acre), large diameter (≥ 13 cm dbh), standing dead snags ≥
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
4/10/2020
Brown-headed Nuthatch Sitta pusilla
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
2 m in height are present (McNair 1984, McComb et al 1986, O’Halloran and Conner 1987, Conner and Dickson 1997,
Withgott and Smith 1998, Wilson and Watts 1999, Weiss 2003, Dornak et al 2004, Lloyd and Slater 2006, Cox and Slater
2007) . Population management units supporting 100-200 breeding pairs as a self-sustaining population should generally be ≥ 350 ha each consisting of suitable mature pine habitat patches ≥ 5 ha separated by ≤ 300 m of unsuitable habitat
(McIntyre 1995, Herb and Burt 2000, Cox and Slater 2007, Tall Timbers Research Station 2009).
SMART Objectives
Maintain/increase pine-dominated forest canopy cover at/to 25-70%, at least 50% of which is in pines ≥ 25 cm dbh, as measured within 5 years post treatment (Engstrom 1984, Repenning and Labisky 1995, White 1996,
Withgott and Smith 1998, Wilson and Watts 1999, Herb and Burt 2000, Conner 2002, LMJV 2009).
Maintain/increase standing dead snags ≥ 13 cm dbh and ≥ 2 m in height to ≥ 5/ha (2.5/acre) as measured by the end of the first post-treatment growing season (McNair 1984, McComb et al 1986, O’Halloran and Conner 1987,
Conner and Dickson 1997, Wilson and Watts 1999, Herb and Burt 2000, Weiss 2003, Dornak et al 2004, Lloyd and Slater 2006, Cox and Slater 2007, Tall Timbers Research 2009, LMJV 2009).
Maintain/reduce woody shrub and hardwood cover ≤ 35% (or ≤ 25 hardwood stems/ha, or ≤ 3.0 m 2 /ha hardwood basal area) as measured within 36 months post treatment (Engstrom 1984, Wilson and Watts 1999, Conner et al
2002, Provencher et al 2002a and 2002b, Dornak et al 2004, Allen et al 2006, LMJV 2009).
Maintain/increase grass-forb ground cover at/to 25-80%, as measured by the end of the first post-treatment growing season (Engstrom 1984, Wilson and Watts 1999, Herb and Burt 2000, Allen 2001, Conner et al 2002,
Provencher et al 2002a and 2002b, Rutledge and Conner 2002, Dornak et al 2004, Allen et al 2006).
Monitoring for Treatment Success – Recommended (FIREMon) Protocols
Mature pine canopy % cover/basal area
Large (≥ 13 cm dbh) snag density (#/ha)
Woody shrub hardwood cover/stocking (%,
#/ha).
Live bunch grass-forb understory % cover
Vertical Photo point, Densitiometer, and Tree Data
Photo point, Tree Data, or belt transects
Photo point, line intercept, and Tree Data
Photo point, point intercept, line intercept, or Brown’s Fuel
Transects
References Cited :
Allen, J.C. 2001. Species-habitat relationships for the breeding birds of a longleaf pine ecosystem. M.S. Thesis.
Virginia Polytechnic Institute and State University, Blacksburg, VA. 239 pp.
Allen, J.C., S.M. Krieger, J.R. Walters, and J.A. Collazo. 2006. Associations of breeding birds with fireinfluenced and riparian–upland gradients in a longleaf pine ecosystem. The Auk 123(4):1110–1128.
Barnhill, L. M. and D. Imm. 2006. Pine Savannah Bird Group. South Carolina. [Online] Available at http://www.dnr.sc.gov/cwcs/pdf/PineSavannahbirds.pdf
Accessed 4/28/2009. 10 pp.
Connor, R.N. and J.G. Dickson. 1997.
Relationships between bird communities and forest age, structure, species composition and fragmentation in the west gulf coastal plain. Texas Jour. Science 49(3) Supplement: 123-138.
Connor, R.N., C.E. Shackelford, R.R. Schaefer, D. Saenz, D.C. Rudolph. 2002.
Avian community response to southern pine ecosystem restoration for red-cockaded woodpeckers. Wilson Bulletin, 114(3):324-332.
Cox, J.A. and G.L. Slater. 2007.
Cooperative breeding in the brown-headed nuthatch . Wilson Journal of
Ornithology 119(1):1–8.
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
4/10/2020
Brown-headed Nuthatch Sitta pusilla
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
Dornak, L. L., D. B. Burt, D. Coble, R. N. Conner. 2004.
Relationships between habitat and snag characteristics and the reproductive success of the Brown-headed nuthatch ( Sitta pusilla ) in eastern Texas. Southeastern
Naturalist 3(4):683-694.
Engstrom, R.T., R.L. Crawford, and W. W. Baker. 1984.
Breeding bird populations in relation to changing forest structure following fire exclusion:A 15-year study. Wilson Bulletin 96(3):437-450.
Haas, S.E. 2007.
Fine-scale spatial genetic structure in the brown-headed nuthatch ( Sitta pusilla ). MS Thesis,
University of Florida, Gainesville, FL. 46 pp.
Harlow, R.F., and D.C. Guynn, Jr. 1983.
Snag densities in managed stands of the South Carolina coastal plain.
Southern Journal of Applied Forestry. 7(4):224-229.
Harrison, H.H. 1975.
A Field Guide to Birds' Nests. Houghton Mifflin Co., Boston, MA. 257 pp.
Headstrom, R. 1965.
Birds' Nests. Ives Washburn, Inc., New York, NY. 128 pp. Hamel 1992
Herb, A. and D.B. Burt. 2000.
Influence of habitat use patterns on cooperative breeding in the brown-headed nuthatch. Bull. Of the Texas Ornithological Society 33(3):25-36.
Howell, J.E., J.T. Peterson and M.J. Conroy. 2006.
Building Hierarchical Models of Avian Distributions for the State of Georgia. Journal of Wildlife Management 72(1):168-178.
Hunter, W.C., L. Peoples and J. Collazo. 2001.
Partners In Flight Bird Conservation Plan for the South Atlantic Coastal Plain (Physiographic Area 3). American Bird Conservancy. The Plains, Virginia. 166 pp.
Kerpez, T.A., and D.F Stauffer. 1989. Avian communities of pine-hardwood forests in the southeast: characteristics, management, and modeling. U.S.D.A., Forest Service, Southeastern Forest Experiment Station. General
Technical Report SE-58:156-169.
Kilgo, J.C., D.L. Gartner, B.R. Chapman, J.B. Dunning, K.E. Franzreb, S.A. Gathreaux, C.H.
Greenberg, D.J. Levey, K.V. Miller and S.F. Pearson. 2002.
A test of an expert-based bird-habitat relationship model in South Carolina. Wildlife Society Bull. 30(3):783-793.
Lloyd, J.D. and G.L. Slater. 2006. Environmental factors affecting productivity of brown-headed nuthatches.
Journal of Wildlife Management 71(6):1968-1975.
Lockwood, J. L., M. S. Ross, and J. P. Sah. 2003.
Smoke on the water: the interplay of fire and water flow on
Everglades restoration. Frontiers in Ecology and the Environment 1:462–468.
Lower Mississippi Valley Joint Venture. 2009.
Landbird Habitat Suitability Index Model: Brown-headed nuthatch Sitta pusilla [Online] Available: http://www.lmvjv.org/hsi_model/ accessed 04/09/2009.
McComb, W.C., S.A. Bonney, R.M. Sheffield, N.D. Cost. 1986.
Snag resources in Florida: Are they sufficient for average populations of primary cavity- nesters? Wildlife Society Bulletin, 14(1):40-48.
McIntyre, N.E. 1995.
Effects of forest patch size on avian diversity. Landscape Ecology 10(2):85-99.
McNair, D.B. 1984.
Clutch-size and nest placement in the brown-headed nuthatch. Wilson Bulletin 96(2):296-301.
Menges, E. S., and M. A. Deyrup. 2001.
Postfire survival in south Florida slash pine: interacting effects of fire intensity, fire season, vegetation, burn size, and bark beetles. International Journal of Wildland Fire 10:53–63.
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
4/10/2020
Brown-headed Nuthatch Sitta pusilla
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
Meyers, J.M., and A.S. Johnson. 1978.
Bird communities associated with succession and management of loblolly shortleaf pine forests. USDA Forest Service, Southeast Forest Experiment Station, Gen. Tech. Rep. SE-14:50-65.
Miller, K.E., and G.A. Jones. 1999.
Nesting phenology and cooperative breeding of the brown-headed nuthatch in north Florida pinelands. Florida. Field Naturalist 27:89-94.
Morse, D. H. 1970.
Ecological aspects of some mixed-species foraging flocks of birds. Ecological Monographs
40:119-168.
Norris, R.A. 1958.
Comparative biosystematics and life history of the nuthatches Sitta pygmaea and Sitta pusilla .
University of California Publications of Zoology 56:119-300.
O’Halloran, K. A., and R. N. Conner. 1987.
Habitat use by brown-headed nuthatches. Bull. of the Texas
Ornithological Society 20:7–13.
Platt, W. J., B. Beckage, R. F. Doren, and H. H. Slater. 2002.
Interactions of large-scale disturbances: prior fire regimes and hurricane mortality of savanna pines. Ecology 83:1566–1572.
Provencher, L.D., N.M. Gobris and L.A. Brennan. 2002a.
Effects of hardwood reduction on winter birds in northwest florida longleaf pine sandhill forests. The Auk 119(1):71–87.
Provencher, L.D., N.M. Gobris, L.A. Brennan, D.R. Gordon, and J.L. Hardesty. 2002b.
Breeding bird response to mid-story hardwood reduction in Florida sandhill longleaf pine forests. Journal of Wildlife Management.
66(3):641-661.
Repenning, R.W. and R.F. Labisky. 1985.
Effects of even-age timber management on bird communities of the longleaf pine forest in northern Florida. Journal of Wildlife Management. 49(4):1088-1098.
Rutledge, B.T., and L.M. Conner. 2002.
Potential effects of groundcover restoration on breeding bird communities in longleaf pine stands. Wildlife Society Bulletin, 30(2):354-360.
Tall Timbers Research Station. 2009. Brown-headed Nuthatch: Research Update. [Online] Available at http://www.talltimbers.org/ve-bhnh.html
. accessed 3/2/2009. 3 pp.
Taylor, B.T. 2003.
Arthropod assemblages on longleaf pines: a possible link between the red-cockaded woodpecker and groundcover vegetation. MS Thesis, Virginia Polytechnic Institute and State Univ., Blacksburg, VA. 106 pp.
The Nature Conservancy. 2000.
Species Management Abstract: Brown-headed nuthatch. The Nature
Conservancy, Arlington, VA. Available at http://conserveonline.org/library/bhnu.doc/view.html
accessed 04/09/2009.
Thill, R.E., D.C. Rudolph and N.E. Koerth. 2003.
Shortleaf pine-bluestem restoration for red-cockaded woodpeckers in the Ouachita Mountains: implications for other taxa. Red-cockaded Woodpecker Symposium, Savannah,
GA, Jan. 27-31. 2003. pp. 657-671.
Weiss, V.A. 2003.
Nest sites used by brown-headed nuthatches in the Virginia coastal plain. The Raven 74(1):3-10.
White, D.H., C.B. Kepler, J.S. Hatfield, P.W. Sykes, Jr., and J.T. Seginak. 1996.
Habitat associations of birds in the Georgia piedmont during winter. Journal of Field Ornithology, 67(1):159-166.
White, D.H., B.R. Chapman, J.H. Brunjes, IV, R.V. Raftovich, Jr., and J.T. Seginak. 1999.
Abundance and reproduction of songbirds in burned and unburned pine forests of the Georgia piedmont. Journal of Field
Ornithology, 70(3):414-424.
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
4/10/2020
Brown-headed Nuthatch Sitta pusilla
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
Wilson, C. W., R. E. Masters, and G. A. Bukenhofer. 1995.
Breeding bird response to pine-grassland community restoration for red-cockaded woodpeckers. J. Wildlife Management 59(?):56-67
Wilson, M. D., and B. D. Watts. 1999.
Response of Brown-headed nuthatches to thinning of pine plantations.
Wilson Bulletin 111(1). pp. 56-60.
Wilson, M. D., and B. D. Watts. 2000.
Breeding bird communities in pine plantations on the coastal plain of
North Carolina. The Chat. 64(1):1-14.
Withgott, J.H. and K.G. Smith. 1998.
Brown-headed nuthatch ( Sitta pusilla ). Pp. 1-23 In Poole, A. and
Gill F. (eds.). The Birds of North America No. 349.
23 pp.
Tall Timbers Research Station 2009
Fire Ecology Profile for Conservation Design #4 A Strategic Habitat Conservation Project
4/10/2020