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International Journal of Wildland Fire International Journal of Wildland Fire Society
Journal of the International Association of Wildland Fire
RESEARCH ARTICLE

Positive effects of fire on birds may appear only under narrow combinations of fire severity and time-since-fire

Richard L. Hutto A C and David A. Patterson B
+ Author Affiliations
- Author Affiliations

A Division of Biological Sciences, 32 Campus Drive #4824, University of Montana, Missoula, MT 59812, USA.

B Department of Mathematical Sciences, 32 Campus Drive #0864, University of Montana, Missoula, MT 59812, USA.

C Corresponding author. Email: hutto@mso.umt.edu

International Journal of Wildland Fire 25(10) 1074-1085 https://doi.org/10.1071/WF15228
Submitted: 22 July 2015  Accepted: 26 May 2016   Published: 11 July 2016

Abstract

We conducted bird surveys in 10 of the first 11 years following a mixed-severity fire in a dry, low-elevation mixed-conifer forest in western Montana, United States. By defining fire in terms of fire severity and time-since-fire, and then comparing detection rates for species inside 15 combinations of fire severity and time-since-fire, with their rates of detection in unburned (but otherwise similar) forest outside the burn perimeter, we were able to assess more nuanced effects of fire on 50 bird species. A majority of species (60%) was detected significantly more frequently inside than outside the burn. It is likely that the beneficial effects of fire for some species can be detected only under relatively narrow combinations of fire severity and time-since-fire. Because most species responded positively and uniquely to some combination of fire severity and time-since-fire, these results carry important management implications. Specifically, the variety of burned-forest conditions required by fire-dependent bird species cannot be created through the application of relatively uniform low-severity prescribed fires, through land management practices that serve to reduce fire severity or through post-fire salvage logging, which removes the dead trees required by most disturbance-dependent bird species.

Additional keywords: Black-backed Woodpecker, conifer forest, ecological integrity, fire severity, mixed-severity fire, restoration, salvage logging, wildfire.


References

Agresti A (2002) ‘Categorical data analysis’, 2nd edn. (John Wiley and Sons: Hoboken, NJ).

Apfelbaum SI, Haney A (1985) Changes in bird populations during succession following fire in northern Great Lakes Wilderness. In ‘Proceedings – National wilderness research conference: current research’. (Ed. RC Lucas) USDA General Technical Report INT-212, pp 10–16. (Ogden, UT)

Baker WL (2009) ‘Fire ecology in Rocky Mountain landscapes’. (Island Press: Washington, DC)

Baker WL (2012) Implications of spatially extensive historical data from surveys for restoring dry forests of Oregon’s eastern Cascades. Ecosphere 3, 23
Implications of spatially extensive historical data from surveys for restoring dry forests of Oregon’s eastern Cascades.Crossref | GoogleScholarGoogle Scholar |

Baker WL (2015a) Are high-severity fires burning at much higher rates recently than historically in dry-forest landscapes of the western USA? PLoS One 10, e0136147
Are high-severity fires burning at much higher rates recently than historically in dry-forest landscapes of the western USA?Crossref | GoogleScholarGoogle Scholar | 26351850PubMed |

Baker WL (2015b) Historical northern spotted owl habitat and old-growth dry forests maintained by mixed-severity wildfires. Landscape Ecology 30, 655–666.
Historical northern spotted owl habitat and old-growth dry forests maintained by mixed-severity wildfires.Crossref | GoogleScholarGoogle Scholar |

Baker WL, Williams MA (2015) Bet-hedging dry-forest resilience to climate-change threats in the western USA based on historical forest structure. Frontiers in Ecology and Evolution 2, art88
Bet-hedging dry-forest resilience to climate-change threats in the western USA based on historical forest structure.Crossref | GoogleScholarGoogle Scholar |

Baker WL, Veblen TT, Sherriff RL (2007) Fire, fuels and restoration of ponderosa pine-Douglas fir forests in the Rocky Mountains, USA. Journal of Biogeography 34, 251–269.
Fire, fuels and restoration of ponderosa pine-Douglas fir forests in the Rocky Mountains, USA.Crossref | GoogleScholarGoogle Scholar |

Bechtoldt CL, Stouffer PC (2005) Home-range size, response to fire, and habitat preferences of wintering Henslow’s Sparrows. The Wilson Bulletin 117, 211–225.
Home-range size, response to fire, and habitat preferences of wintering Henslow’s Sparrows.Crossref | GoogleScholarGoogle Scholar |

Blackford JL (1955) Woodpecker concentration in burned forest. The Condor 57, 28–30.
Woodpecker concentration in burned forest.Crossref | GoogleScholarGoogle Scholar |

Bond ML, Siegel RB, Hutto RL, Saab VA, Shunk SA (2012) A new forest fire paradigm: the need for high-severity fires. The Wildlife Professional 6, 46–49.

Breininger DR, Smith RB (1992) Relationships between fire and bird density in coastal scrub and slash pine flatwoods in Florida. American Midland Naturalist 127, 233–240.

Brown S, Clarke M, Clarke R (2009) Fire is a key element in the landscape-scale habitat requirements and global population status of a threatened bird: the Mallee Emu-wren (Stipiturus mallee). Biological Conservation 142, 432–445.
Fire is a key element in the landscape-scale habitat requirements and global population status of a threatened bird: the Mallee Emu-wren (Stipiturus mallee).Crossref | GoogleScholarGoogle Scholar |

Brown JK, Smith JK (2000) Wildland fire in ecosystems: effects of fire on flora. USDA Forest Service, Rocky Mountain Research Station, General Technical Report RMRS-GTR-42-volume 2. (Ogden,UT).

Buckland ST, Anderson DR, Burnham KP, Laake JL, Borchers DL, Thomas L (2001) ‘Introduction to distance sampling: estimating abundance of animal populations.’ (Oxford University Press: New York, NY.)

Chalmandrier L, Midgley GF, Barnard P, Sirami C (2013) Effects of time since fire on birds in a plant diversity hotspot. Acta Oecologica 49, 99–106.
Effects of time since fire on birds in a plant diversity hotspot.Crossref | GoogleScholarGoogle Scholar |

Conway CJ, Kirkpatrick C (2007) Effect of forest fire suppression on Buff-breasted Flycatchers. Journal of Wildlife Management 71, 445–457.
Effect of forest fire suppression on Buff-breasted Flycatchers.Crossref | GoogleScholarGoogle Scholar |

Covert-Bratland KA, Block WM, Theimer TC (2006) Hairy woodpecker winter ecology in ponderosa pine forests representing different ages since wildfire. Journal of Wildlife Management 70, 1379–1392.
Hairy woodpecker winter ecology in ponderosa pine forests representing different ages since wildfire.Crossref | GoogleScholarGoogle Scholar |

DellaSala DA, Karr JR, Schoennagel T, Perry D, Noss RF, Lindenmayer D, Beschta R, Hutto RL, Swanson ME, Evans J (2006) Postfire logging debate ignores many issues. Science 314, 51–52.
Postfire logging debate ignores many issues.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD28XhtVKhsLvK&md5=9ceb36d4d92289d6fe82bc73e2c89a83CAS | 17023633PubMed |

DellaSala DA, Bond ML, Hanson CT, Hutto RL, Odion DC (2014) Complex early seral forests of the Sierra Nevada: what are they and how can they be managed for ecological integrity? Natural Areas Journal 34, 310–324.
Complex early seral forests of the Sierra Nevada: what are they and how can they be managed for ecological integrity?Crossref | GoogleScholarGoogle Scholar |

Duckworth RA (2008) Adaptive dispersal strategies and the dynamics of a range expansion. American Naturalist 172, S4–S17.
Adaptive dispersal strategies and the dynamics of a range expansion.Crossref | GoogleScholarGoogle Scholar | 18554143PubMed |

Duckworth RA (2009) Maternal effects and range expansion: a key factor in a dynamic process? Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 364, 1075–1086.
Maternal effects and range expansion: a key factor in a dynamic process?Crossref | GoogleScholarGoogle Scholar | 19324612PubMed |

Duckworth RA (2010) Evolution of personality: developmental constraints on behavioral flexibility. The Auk 127, 752–758.
Evolution of personality: developmental constraints on behavioral flexibility.Crossref | GoogleScholarGoogle Scholar |

Duckworth RA (2012) Evolution of genetically integrated dispersal strategies. In ‘Dispersal Ecology and Evolution’. (Ed. J Clobert) pp 83–94. (Oxford University Press: Oxford, UK).

Duckworth RA (2014) Human-induced changes in the dynamics of species coexistence. In ‘Avian urban ecology: physiological and behavioural adaptations to the urban habitat’. (Eds D Gil, H Brumm) pp 181–191. (Oxford University Press: Oxford, UK).

Duckworth RA, Badyaev AV (2007) Coupling of dispersal and aggression facilitates the rapid range expansion of a passerine bird. Proceedings of the National Academy of Sciences of the United States of America 104, 15017–15022.
Coupling of dispersal and aggression facilitates the rapid range expansion of a passerine bird.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD2sXhtVyqt7nI&md5=becec0de668e6fa10a58b69e8fafd1d0CAS | 17827278PubMed |

Dudley JG, Saab VA, Hollenbeck JP (2012) Foraging-habitat selection of Black-backed Woodpeckers in forest burns of southwestern Idaho. The Condor 114, 348–357.
Foraging-habitat selection of Black-backed Woodpeckers in forest burns of southwestern Idaho.Crossref | GoogleScholarGoogle Scholar |

Fontaine JB, Kennedy PL (2012) Meta-analysis of avian and small-mammal response to fire severity and fire surrogate treatments in U.S. fire-prone forests. Ecological Applications 22, 1547–1561.

Ganey JL, Block WM, Boucher PF (1996) Effects of fire on birds in Madrean forests and woodlands. In ‘Effects of fire on Madrean Province ecosystems’. (Eds PF Ffolliott, LF DeBano, MB Baker, GJ Gottfried, G Solis-Garza, CB Edminster, DG Neary, LS Allen, RH Hamre) USDA Forest Service General Technical Report RM-GTR-289, pp. 146–154. (Fort Collins, CO)

Gentry DJ, Vierling KT (2007) Old burns as source habitats for Lewis’s Woodpeckers breeding in the Black Hills of South Dakota. The Condor 109, 122–131.
Old burns as source habitats for Lewis’s Woodpeckers breeding in the Black Hills of South Dakota.Crossref | GoogleScholarGoogle Scholar |

Hannah KC, Hoyt JS (2004) Northern Hawk Owls and recent burns: does burn age matter? The Condor 106, 420–423.
Northern Hawk Owls and recent burns: does burn age matter?Crossref | GoogleScholarGoogle Scholar |

Hanson CT, North MP (2008) Postfire woodpecker foraging in salvage-logged and unlogged forests of the Sierra Nevada. The Condor 110, 777–782.
Postfire woodpecker foraging in salvage-logged and unlogged forests of the Sierra Nevada.Crossref | GoogleScholarGoogle Scholar |

Hardy CC (2005) Wildland fire hazard and risk: problems, definitions, and context. Forest Ecology and Management 211, 73–82.
Wildland fire hazard and risk: problems, definitions, and context.Crossref | GoogleScholarGoogle Scholar |

Hargrove WW, Pickering J (1992) Pseudoreplication: a sine qua non for regional ecology. Landscape Ecology 6, 251–258.
Pseudoreplication: a sine qua non for regional ecology.Crossref | GoogleScholarGoogle Scholar |

Hessburg PF, Salter RB, James KM (2007) Re-examining fire severity relations in pre-management era mixed conifer forests: inferences from landscape patterns of forest structure. Landscape Ecology 22, 5–24.
Re-examining fire severity relations in pre-management era mixed conifer forests: inferences from landscape patterns of forest structure.Crossref | GoogleScholarGoogle Scholar |

Heyerdahl EK, Lertzman K, Wong CM (2012) Mixed-severity fire regimes in dry forests of southern interior British Columbia, Canada. Canadian Journal of Forest Research 42, 88–98.
Mixed-severity fire regimes in dry forests of southern interior British Columbia, Canada.Crossref | GoogleScholarGoogle Scholar |

Hobson KA, Schieck J (1999) Changes in bird communities in boreal mixedwood forest: harvest and wildfire effects over 30 years. Ecological Applications 9, 849–863.
Changes in bird communities in boreal mixedwood forest: harvest and wildfire effects over 30 years.Crossref | GoogleScholarGoogle Scholar |

Holmes AL, Robinson WD (2013) Fire mediated patterns of population densities in mountain big sagebrush bird communities. Journal of Wildlife Management 77, 737–748.
Fire mediated patterns of population densities in mountain big sagebrush bird communities.Crossref | GoogleScholarGoogle Scholar |

Hoyt JS, Hannon SJ (2002) Habitat associations of Black-backed and Three-toed woodpeckers in the boreal forest of Alberta. Canadian Journal of Forest Research 32, 1881–1888.
Habitat associations of Black-backed and Three-toed woodpeckers in the boreal forest of Alberta.Crossref | GoogleScholarGoogle Scholar |

Hudak AT, Robichaud PR, Evans JB, Clark J, Lannom K, Morgan P, Stone C (2004) Field validation of Burned Area Reflectance Classification (BARC) products for post fire assessment. In ‘Remote sensing for field users; proceedings of the 10th Forest Service remote sensing applications conference’. 5–9 April 2004, Betheseda, MD. (Ed. JD Greer) American Society of Photogrammetry and Remote Sensing. (CD-ROM) (Salt Lake City, UT).

Hudak AT, Morgan P, Bobbitt MJ, Smith AMS, Lewis SA, Lentile LB, Robichaud PR, Clark JT, McKinley RA (2007) The relationship of multispectral satellite imagery to immediate fire effects. Fire Ecology 3, 64–90.
The relationship of multispectral satellite imagery to immediate fire effects.Crossref | GoogleScholarGoogle Scholar |

Huot M, Ibarzabal J (2006) A comparison of the age-class structure of black-backed woodpeckers found in recently burned and unburned boreal coniferous forests in eastern Canada. Annales Zoologici Fennici 43, 131–136.

Hutto RL (1995) Composition of bird communities following stand-replacement fires in northern Rocky Mountain (U.S.A.) conifer forests. Conservation Biology 9, 1041–1058.
Composition of bird communities following stand-replacement fires in northern Rocky Mountain (U.S.A.) conifer forests.Crossref | GoogleScholarGoogle Scholar |

Hutto RL (2008) The ecological importance of severe wildfires: some like it hot. Ecological Applications 18, 1827–1834.
The ecological importance of severe wildfires: some like it hot.Crossref | GoogleScholarGoogle Scholar | 19263880PubMed |

Hutto RL (2016) Should scientists be required to use a model-based solution to adjust for possible distance-based detectability bias? Ecological Applications , In press.

Hutto RL, Belote RT (2013) Distinguishing four types of monitoring based on the questions they address. Forest Ecology and Management 289, 183–189.
Distinguishing four types of monitoring based on the questions they address.Crossref | GoogleScholarGoogle Scholar |

Hutto RL, Gallo SM (2006) The effects of postfire salvage logging on cavity-nesting birds. The Condor 108, 817–831.
The effects of postfire salvage logging on cavity-nesting birds.Crossref | GoogleScholarGoogle Scholar |

Hutto RL, Young JS (2002) Regional landbird monitoring: perspectives from the northern Rocky Mountains. Wildlife Society Bulletin 30, 738–750.

Hutto RL, Pletschet SM, Hendricks P (1986) A fixed-radius point count method for nonbreeding and breeding season use. The Auk 103, 593–602.

Hutto RL, Conway CJ, Saab VA, Walters JR (2008) What constitutes a natural fire regime? Insight from the ecology and distribution of coniferous forest birds in North America. Fire Ecology 4, 115–132.
What constitutes a natural fire regime? Insight from the ecology and distribution of coniferous forest birds in North America.Crossref | GoogleScholarGoogle Scholar |

Hutto RL, Bond ML, DellaSala DA (2015) Using bird ecology to learn about the benefits of severe fire. In ‘The ecological importance of mixed-severity fires: nature’s phoenix’. (Eds DA DellaSala, CT Hanson) pp 55–88. ( Elsevier: New York, NY).

Hutto RL, Keane RE, Sherriff RL, Rota CT, Eby LA, Saab VA (2016) Toward a more ecologically informed view of severe forest fires. Ecosphere 7, e01255

Imbeau L, Savard J-P, Gagnon R (1999) Comparing bird assemblages in successional black spruce stands originating from fire and logging. Canadian Journal of Zoology 77, 1850–1860.
Comparing bird assemblages in successional black spruce stands originating from fire and logging.Crossref | GoogleScholarGoogle Scholar |

Johnson DH (2008) In defense of indices: the case of bird surveys. The Journal of Wildlife Management 72, 857–868.
In defense of indices: the case of bird surveys.Crossref | GoogleScholarGoogle Scholar |

Kirkpatrick C, Conway C, Jones PB (2006) Distribution and relative abundance of forest birds in relation to burn severity in southeastern Arizona. The Journal of Wildlife Management 70, 1005–1012.
Distribution and relative abundance of forest birds in relation to burn severity in southeastern Arizona.Crossref | GoogleScholarGoogle Scholar |

Koivula MJ, Schmiegelow FKA (2007) Boreal woodpecker assemblages in recently burned forested landscapes in Alberta, Canada: effects of post-fire harvesting and burn severity. Forest Ecology and Management 242, 606–618.
Boreal woodpecker assemblages in recently burned forested landscapes in Alberta, Canada: effects of post-fire harvesting and burn severity.Crossref | GoogleScholarGoogle Scholar |

Koplin JR (1969) The numerical response of woodpeckers to insect prey in a subalpine forest in Colorado. The Condor 71, 436–438.
The numerical response of woodpeckers to insect prey in a subalpine forest in Colorado.Crossref | GoogleScholarGoogle Scholar |

Kotliar NB, Hejl SJ, Hutto RL, Saab VA, Melcher CP, McFadzen ME (2002) Effects of fire and post-fire salvage logging on avian communities in conifer-dominated forests of the western United States. Studies in Avian Biology 25, 49–64.

Kotliar NB, Saab VA, Hutto RL (2005) Fire on the mountain: birds and burns in the Rocky Mountains. USDA Forest Service, General Technical Report PSW-GTR-191, pp 1090–1092. (Albany, CA)

Kotliar NB, Kennedy PL, Ferree K (2007) Avifaunal responses to fire in southwestern montane forests along a burn severity gradient. Ecological Applications 17, 491–507.
Avifaunal responses to fire in southwestern montane forests along a burn severity gradient.Crossref | GoogleScholarGoogle Scholar | 17489255PubMed |

Kotliar NB, Reynolds EW, Deutschman DH (2008) American three-toed woodpecker response to burn severity and prey availability at multiple spatial scales. Fire Ecology 4, 26–45.
American three-toed woodpecker response to burn severity and prey availability at multiple spatial scales.Crossref | GoogleScholarGoogle Scholar |

Lee DE, Bond ML, Siegel RB (2012) Dynamics of breeding-season site occupancy of the California Spotted Owl in burned forests. The Condor 114, 792–802.
Dynamics of breeding-season site occupancy of the California Spotted Owl in burned forests.Crossref | GoogleScholarGoogle Scholar |

Lentile LB, Morgan P, Hudak AT, Bobbitt MJ, Lewis SA, Smith AMS, Robichaud PR (2007) Post-fire burn severity and vegetation response following eight large wildfires across the western United States. Fire Ecology 3, 91–108.
Post-fire burn severity and vegetation response following eight large wildfires across the western United States.Crossref | GoogleScholarGoogle Scholar |

Lindenmayer DB, Blanchard W, McBurney L, Blair D, Banks SC, Driscoll DA, Smith AL, Gill AM (2014) Complex responses of birds to landscape-level fire extent, fire severity and environmental drivers. Diversity and Distributions 20, 467–477.
Complex responses of birds to landscape-level fire extent, fire severity and environmental drivers.Crossref | GoogleScholarGoogle Scholar |

MacKenzie DI, Nichols JD, Royle JA, Pollock KH, Bailey LL, Hines JE (2006) ‘Occupancy estimation and modeling: inferring patterns and dynamics of species occurrence’. (Elsevier Academic Press: Burlington, MA).

Margolis EQ, Swetnam TW, Allen CD (2011) Historical stand-replacing fire in upper montane forests of the Madrean Sky Islands and Mogollon Plateau, southwestern USA. Fire Ecology 7, 88–107.
Historical stand-replacing fire in upper montane forests of the Madrean Sky Islands and Mogollon Plateau, southwestern USA.Crossref | GoogleScholarGoogle Scholar |

Marlon JR, Bartlein PJ, Gavin DG, Long CJ, Anderson RS, Briles CE, Brown KJ, Colombaroli D, Hallett DJ, Power MJ, Scharf EA, Walsh MK (2012) Long-term perspective on wildfires in the western USA. Proceedings of the National Academy of Sciences of the United States of America 109, E535–E543.
Long-term perspective on wildfires in the western USA.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC38XksVyhu7g%3D&md5=dd45aa898bccea2f2f637621b0f082d3CAS | 22334650PubMed |

Murphy EG, Lehnhausen WH (1998) Density and foraging ecology of woodpeckers following a stand-replacement fire. The Journal of Wildlife Management 62, 1359–1372.
Density and foraging ecology of woodpeckers following a stand-replacement fire.Crossref | GoogleScholarGoogle Scholar |

Nappi A, Drapeau P (2009) Reproductive success of the Black-backed Woodpecker (Picoides arcticus) in burned boreal forests: are burns source habitats? Biological Conservation 142, 1381–1391.
Reproductive success of the Black-backed Woodpecker (Picoides arcticus) in burned boreal forests: are burns source habitats?Crossref | GoogleScholarGoogle Scholar |

Nappi A, Drapeau P (2011) Pre-fire forest conditions and fire severity as determinants of the quality of burned forests for deadwood-dependent species: the case of the black-backed woodpecker. Canadian Journal of Forest Research 41, 994–1003.
Pre-fire forest conditions and fire severity as determinants of the quality of burned forests for deadwood-dependent species: the case of the black-backed woodpecker.Crossref | GoogleScholarGoogle Scholar |

Nappi A, Drapeau P, Saint-Germain M, Angers VA (2010) Effect of fire severity on long-term occupancy of burned boreal conifer forests saproxylic insects and wood-foraging birds. International Journal of Wildland Fire 19, 500–511.
Effect of fire severity on long-term occupancy of burned boreal conifer forests saproxylic insects and wood-foraging birds.Crossref | GoogleScholarGoogle Scholar |

Odion DC, Hanson CT, Arsenault A, Baker WL, DellaSala DA, Hutto RL, Klenner W, Moritz MA, Sherriff RL, Veblen TT, Williams MA (2014) Examining historical and current mixed-severity fire regimes in ponderosa pine and mixed-conifer forests of western North America. PLoS ONE 9, e87852
Examining historical and current mixed-severity fire regimes in ponderosa pine and mixed-conifer forests of western North America.Crossref | GoogleScholarGoogle Scholar | 24498383PubMed |

Perry DA, Hessburg PF, Skinner CN, Spies TA, Stephens SL, Taylor AN, Franklin JF, McComb B, Riegel G (2011) The ecology of mixed severity fire regimes in Washington, Oregon, and Northern California. Forest Ecology and Management 262, 703–717.
The ecology of mixed severity fire regimes in Washington, Oregon, and Northern California.Crossref | GoogleScholarGoogle Scholar |

Pons P, Clavero M (2010) Bird responses to fire severity and time since fire in managed mountain rangelands. Animal Conservation 13, 294–305.
Bird responses to fire severity and time since fire in managed mountain rangelands.Crossref | GoogleScholarGoogle Scholar |

Pyke GH, Saillard R, Smith J (1995) Abundance of Eastern Bristlebirds in relation to habitat and fire history. Emu 95, 106–110.
Abundance of Eastern Bristlebirds in relation to habitat and fire history.Crossref | GoogleScholarGoogle Scholar |

R Core Team (2014) R: A language and environment for statistical computing. (R Foundation for Statistical Computing: Vienna, Austria). Available at http://www.R-project.org/ [Verified 7 June 2016]

Ralph CJ, Sauer JR, Droege S (1995) Monitoring bird populations by point counts. USDA Forest Service, General Technical Report PSW-GTR-149, pp 1–181. (Albany, CA)

Raphael MG, Morrison ML, Yoder-Williams MP (1987) Breeding bird populations during twenty-five years of post-fire succession in the Sierra Nevada. Condor 89, 614–626.
Breeding bird populations during twenty-five years of post-fire succession in the Sierra Nevada.Crossref | GoogleScholarGoogle Scholar |

Rush S, Klaus N, Keyes T, Petrick J, Cooper R (2012) Fire severity has mixed benefits to breeding bird species in the southern Appalachians. Forest Ecology and Management 263, 94–100.
Fire severity has mixed benefits to breeding bird species in the southern Appalachians.Crossref | GoogleScholarGoogle Scholar |

Saab VA, Dudley JG (1998) Responses of cavity-nesting birds to stand-replacement fire and salvage logging in ponderosa pine/Douglas-fir forests of southwestern Idaho. USDA Forest Service, Research Paper RMRS-RP-11, pp 1–17. (Ogden, UT)

Saab VA, Dudley J, Thompson WL (2004) Factors influencing occupancy of nest cavities in recently burned forests. The Condor 106, 20–36.
Factors influencing occupancy of nest cavities in recently burned forests.Crossref | GoogleScholarGoogle Scholar |

Saab VA, Russell RE, Dudley J (2007) Nest densities of cavity-nesting birds in relation to post-fire salvage logging and time since wildfire. Condor 109, 97–108.
Nest densities of cavity-nesting birds in relation to post-fire salvage logging and time since wildfire.Crossref | GoogleScholarGoogle Scholar |

Saab VA, Russell RE, Dudley JG (2009) Nest-site selection by cavity-nesting birds in relation to postfire salvage logging. Forest Ecology and Management 257, 151–159.
Nest-site selection by cavity-nesting birds in relation to postfire salvage logging.Crossref | GoogleScholarGoogle Scholar |

Saracco JF, Siegel RB, Wilkerson RL (2011) Occupancy modeling of Black-backed Woodpeckers on burned Sierra Nevada forests. Ecosphere 2, art31
Occupancy modeling of Black-backed Woodpeckers on burned Sierra Nevada forests.Crossref | GoogleScholarGoogle Scholar |

Schieck J, Song SJ (2006) Changes in bird communities throughout succession following fire and harvest in boreal forests of western North America: literature review and meta-analyses. Canadian Journal of Forest Research 36, 1299–1318.
Changes in bird communities throughout succession following fire and harvest in boreal forests of western North America: literature review and meta-analyses.Crossref | GoogleScholarGoogle Scholar |

Sherriff RL, Platt RV, Veblen TT, Schoennagel TL, Gartner MH (2014) Historical, observed, and modeled wildfire severity in montane forests of the Colorado Front Range. PLoS One 9, e106971
Historical, observed, and modeled wildfire severity in montane forests of the Colorado Front Range.Crossref | GoogleScholarGoogle Scholar | 25251103PubMed |

Siegel RB, Tingley MW, Wilkerson RL, Howell CA, Johnson M, Pyle P (2016) Age structure of Black-backed Woodpecker populations in burned forests. The Auk 133, 69–78.
Age structure of Black-backed Woodpecker populations in burned forests.Crossref | GoogleScholarGoogle Scholar |

Smucker KM, Hutto RL, Steele BM (2005) Changes in bird abundance after wildfire: importance of fire severity and time since fire. Ecological Applications 15, 1535–1549.
Changes in bird abundance after wildfire: importance of fire severity and time since fire.Crossref | GoogleScholarGoogle Scholar |

Stephens JL, Ausprey IJ, Seavy NE, Alexander JD (2015) Fire severity affects mixed broadleaf-conifer forest bird communities: results for 9 years following fire. The Condor 117, 430–446.
Fire severity affects mixed broadleaf-conifer forest bird communities: results for 9 years following fire.Crossref | GoogleScholarGoogle Scholar |

Swanson ME, Franklin JF, Beschta RL, Crisafulli CM, DellaSala DA, Hutto RL, Lindenmayer DB, Swanson FJ (2011) The forgotten stage of forest succession: early-successional ecosystems on forest sites. Frontiers in Ecology and the Environment 9, 117–125.
The forgotten stage of forest succession: early-successional ecosystems on forest sites.Crossref | GoogleScholarGoogle Scholar |

Taylor DL (1973) Some ecological implications of forest fire control in Yellowstone National Park, Wyoming. Ecology 54, 1394–1396.
Some ecological implications of forest fire control in Yellowstone National Park, Wyoming.Crossref | GoogleScholarGoogle Scholar |

Taylor DL (1979) Forest fires and the tree-hole nesting cycle in Grand Teton and Yellowstone National Park. In ‘Proceedings of the First Conference on Scientific Research in the National Parks, vol. 1’, 9–12 November 1976, New Orleans, LA. (Ed. RM Linn) USDI National Park Service Transactions Series 5, pp. 509–511. Available at https://archive.org/details/proceedingsoffir00linn [Verified 16 June 2016]

Taylor DL, Barmore WJ (1980) Post-fire succession of avifauna in coniferous forests of Yellowstone and Grand Teton National Parks, Wyoming. In ‘Workshop proceedings: management of western forests and grasslands for nongame birds’. (Ed. RM DeGraaf) USDA Forest Service, General Technical Report INT-86, pp 130–145 (Ogden, UT).

Tingley MW, Wilkerson RL, Bond ML, Howell CA, Siegel RB (2014) Variation in home-range size of Black-backed Woodpeckers. The Condor 116, 325–340.
Variation in home-range size of Black-backed Woodpeckers.Crossref | GoogleScholarGoogle Scholar |

Van Tyne J (1926) An unusual flight of Arctic Three-toed Woodpeckers. The Auk 43, 469–474.
An unusual flight of Arctic Three-toed Woodpeckers.Crossref | GoogleScholarGoogle Scholar |

Veblen TT, Romme WH, Regan C (2012) Regional application of historical ecology at ecologically defined scales: forest ecosystems in the Colorado front range. In ‘Historical environmental variation in conservation and natural resource management’. (Eds JA Wiens, GD Hayward, HD Safford, CM Giffen) pp 149–165. (John Wiley and Sons, Ltd.: Oxford, UK)

Vierling KT, Gentry DJ (2008) Red-headed Woodpecker density and productivity in relation to time since fire in burned pine forests. Fire Ecology 4, 15–25.
Red-headed Woodpecker density and productivity in relation to time since fire in burned pine forests.Crossref | GoogleScholarGoogle Scholar |

Vierling K, Lentile L (2008) Indirect effects of fire severity on avian communities in ponderosa pine and aspen forests in western North America: a review. Fire Ecology 4, 133–149.
Indirect effects of fire severity on avian communities in ponderosa pine and aspen forests in western North America: a review.Crossref | GoogleScholarGoogle Scholar |

Villard P, Beninger CW (1993) Foraging behavior of male Black-backed and Hairy woodpeckers in a forest burn. Journal of Field Ornithology 64, 71–76.

Villard MA, Schieck J (1997) Immediate post-fire nesting by Black-backed Woodpeckers, Picoides arcticus, in northern Alberta. Canadian Field Naturalist 111, 478–479.

Welsh AH, Lindenmayer DB, Donnelly CF (2013) Fitting and interpreting occupancy models. PLoS One 8, e52015
Fitting and interpreting occupancy models.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3sXhtlCqtL0%3D&md5=3b3a4aff73a75e85cb3a7935b744d326CAS | 23326323PubMed |

Wickham H (2009) ‘ggplot2: elegant graphics for data analysis.’ (Springer: New York, NY).

Wickham H (2014) ‘tidyr: Easily tidy data with spread() and gather() functions. R package.’ Version 0.2.0. Available at http://CRAN.R-project.org/package=tidyr [Verified 7 June 2016]

Wickham H, Francois R (2015) ‘dplyr: a grammar of data manipulation. R package.’ Version 0.4.1. Available at http://CRAN.R-project.org/package=dplyr [Verified 7 June 2016]

Williams MA, Baker WL (2012a) Comparison of the higher-severity fire regime in historical (A.D. 1800s) and modern (A.D. 1984–2009) montane forests across 624,156 ha of the Colorado Front Range. Ecosystems 15, 832–847.
Comparison of the higher-severity fire regime in historical (A.D. 1800s) and modern (A.D. 1984–2009) montane forests across 624,156 ha of the Colorado Front Range.Crossref | GoogleScholarGoogle Scholar |

Williams MA, Baker WL (2012b) Spatially extensive reconstructions show variable-severity fire and heterogeneous structure in historical western United States dry forests. Global Ecology and Biogeography 21, 1042–1052.
Spatially extensive reconstructions show variable-severity fire and heterogeneous structure in historical western United States dry forests.Crossref | GoogleScholarGoogle Scholar |

Williams MA, Baker WL (2014) High-severity fire corroborated in historical dry forests of the western United States: response to Fulé et al. Global Ecology and Biogeography 23, 831–835.
High-severity fire corroborated in historical dry forests of the western United States: response to Fulé et al.Crossref | GoogleScholarGoogle Scholar |

Woinarski JCZ, Brock C, Fisher A, Milne D, Oliver B (1999) Response of birds and reptiles to fire regimes on pastoral land in the Victoria River District, Northern Territory. The Rangeland Journal 21, 24–38.
Response of birds and reptiles to fire regimes on pastoral land in the Victoria River District, Northern Territory.Crossref | GoogleScholarGoogle Scholar |

Yocom-Kent LL, Fulé PZ, Bunn WA, Gdula EG (2015) Historical high-severity fire patches in mixed-conifer forests. Canadian Journal of Forest Research 45, 1587–1596.
Historical high-severity fire patches in mixed-conifer forests.Crossref | GoogleScholarGoogle Scholar |

Yunick RP (1985) A review of recent irruptions of the Black-backed Woodpecker and Three- toed Woodpecker in eastern North America. Journal of Field Ornithology 56, 138–152.

Zhao Q, Azeria ET, Le Blanc M-L, Lemaître J, Fortin D (2013) Landscape-scale disturbances modified bird community dynamics in successional forest environment. PLoS ONE 8, e81358
Landscape-scale disturbances modified bird community dynamics in successional forest environment.Crossref | GoogleScholarGoogle Scholar | 24282585PubMed |