Fire and Birds in the Central Hardwood Landscape
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Fire and Birds in the Central Hardwood Landscape R. Todd Engstrom1 Abstract largely shaped edaphic conditions (Quarterman and others 1993), but fire played an important role to arrest succession Fire is often viewed as destructive of wildlife and wildlife and control brush and saplings as soils developed (Deselm habitat, because it can cause dramatic change. The and Murdock 1993). Lightning-started fires played a role in perception of catastrophic fire that burns up forests and kills creating some of these habitats, but Native Americans wildlife has been fostered to enlist public support for undoubtedly used fire frequently and pervasively to create suppressing fires. A message that is much more difficult to the open habitats found by early European colonists convey is that when fire is removed from the landscape (Bonnicksen and others 1999; Askins 1999). Timber harvest, important wildlife habitat can be slowly and subtly eliminated fire suppression, and plant succession have dramatically from ecosystems that depend on the frequent occurrence of changed the composition of some of these plant fire. communities, and, in the process, the avian comunities. Excellent descriptions of some of the grassland preserves in Many fire-dependent communities—shortleaf pine forests, the Central Hardwoods region (e.g., Tennessee) can be barrens, glades, prairies, balds, and oak savannas—are found at: http://www.tnc.org/infield/State/Tennessee/ embedded in the extensive and diverse central hardwood landscape. Approximately 232 species of birds (119 The purpose of this paper is to provide a brief overview of breeding, 44 wintering, and 69 year-round residents) reside the avifauna of the region, including species that have in this landscape. Some of the rarer bird species in the become extinct or extirpated, to explore the bird species region are closely tied to fire-dependent habitats. The status composition of three general vegetation communities that of the Red-cockaded Woodpecker (Picoides borealis) is were shaped by frequent fires—shortleaf pine (Pinus tenuous in pine forests of Kentucky and Oklahoma where fire echinata) forests, grassland/shrubland communities (balds, has been suppressed for much of the century. Efforts to barrens, and glades), and open oak woodlands and reintroduce fire to restore the open-structured, pine- savannas—and to discuss the effects on birds of use of dominated forests are hopeful signs for the species. prescribed fire in the management and restoration of these Grassland birds are one of the most rapidly declining groups communities. of species in North America. Habitat conversion has eliminated many of the grassland communities within the The Central Hardwoods Avifauna Central Hardwoods region, but plant succession in the absence of fire has also degraded habitat suitability for many Hicks (1998) uses four of Braun’s (1950) forest types to grassland birds. Numerous preserves are being established define the Central Hardwoods region: the mixed mesophytic, to retain the rare biota of these grasslands. Many oak- western mesophytic, oak-hickory, and oak-chestnut forests. dominated forests and oak woodland- savanna mosaics are This large region extends over all or part of 35 states (Figure slowly shifting to greater proportions of trees more 1). I obtained a list of the bird species within this region by associated with mesic conditions, such as maples (Acer sp.), comparing distribution maps of bird species (Dunn 1987) to and canopy closure. These shifts in structure and the map of the Central Hardwoods. Bird species whose composition bring attendant changes in the avian winter or breeding season distributions overlapped with the community. Fire—with proper consideration of the potential outline of the Central Hardwoods region were included; effects on small populations of birds and invertebrates and migrant and transient birds were not. the tradeoffs in bird species that will be favored—is an essential tool for habitat maintenance for some of the rare Approximately 232 bird species winter or breed within the and declining birds in the Central Hardwoods region. Central Hardwoods region (Appendix); all scientific names follow the American Ornithologists’ Union (1998). Sixty-nine (30%) of these species occur year-round, 44 (19%) occur in Introduction the winter, and 119 (51%) occur in the breeding season only. A diverse array of vegetation communities that have Four bird species that were found in the Central Hardwoods experienced frequent fire for millennia are scattered historically, Passenger Pigeon (Ectopistes migratorius), throughout the extensive Central Hardwoods forest region Carolina Parakeet (Conuropsis carolinensis), Ivory-billed (Hicks 1998; Braun 1950; Braun 1950; Hicks 1998; Anderson Woodpecker (Campephilus principalis), and Bachman’s and others 1999). Some of these communities were Warbler (Vermivora bachmanii), have become extinct. The fragmented and highly dispersed (e.g., balds) and some Passenger Pigeon and Bachman’s Warbler may have been were extensive (e.g., the Kentucky barrens). The structure affected by changes in fire regimes in the Central and plant composition of some of these communities was Hardwoods region, although it seems likely that outright habitat loss was more important in their decline than habitat degradation through fire exclusion. For example burning in 1Tall Timbers Research Station, 13093 Henry Beadel Road, forests by Native Americans kept forest structure open and Tallahassee, FL 32312-0918 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 75 Figure 1.—Central Hardwoods region from Braun (1950). parklike, which may have favored mast producing species is that the Bewick’s Wren expanded eastward in the early and facilitated foraging by Passenger Pigeons on the 19th century in response to agricultural expansion, but was ground. Bucher (1992) has hypothesized that destruction of then out-competed by the House Wren as it expanded its forest breeding habitat and the lack of social facilitation in range (Kennedy and White 1997). Although the Bewick’s foraging were the major factors that contributed to the Wren was not found in the region prior to 1821, C. Hunter extinction of the species. Bachman’s Warbler may have (pers. comm.) has suggested that it might have been present narrowly specialized on canebrakes (Arundinaria spp.) in in the oak-savanna mosaic of the Central Hardwoods. Loss bottomland hardwoods for breeding habitat (Remsen 1986). of this habitat to agriculture and modification of the fire Fire played a critical role in maintenance of these large regime may have degraded the best habitat for the Bewick’s stands of cane (Hughes 1966). Wren. Several bird species have been nearly extirpated from the Pine Woodland Specialists Central Hardwoods landscape. The distribution within the in the Central Hardwoods Region region of Greater Prairie Chicken, Red-cockaded Woodpecker, Bewick’s Wren, and Brown-headed Nuthatch Some of the rarest and most threatened bird species of the have all receded within the last 200 years. The Greater Central Hardwoods landscape are associated with pine Prairie Chicken formerly occurred in Ohio, Kentucky, woodlands along the southern edge of the region. The Red- Arkansas, and Missouri, and the eastern seaboard from cockaded Woodpecker, a federally endangered species Massachusetts and Virginia, but disappeared from hunting since 1970, is highly associated with open pinelands in the pressure and habitat destruction (Schroeder and Robb southeastern United States (Jackson 1994). Historically, the 1993). Lumsden (1966) suggested that the Greater Prairie distribution of the woodpecker within the Central Hardwoods Chicken may have colonized the eastern U.S. in response to was primarily the Ouachita Highlands of Arkansas, Missouri, grassland openings created when Native Americans burned and Oklahoma, and the Cumberland Plateau of Kentucky woodlands. and Tennessee (fig. 1). This species was typically associated with stands of shortleaf pine (Pinus echinata) on dry slopes Causes for the decline of the Bewick’s Wren in the Central and ridges. Interestingly, the distribution of Red-cockaded Hardwoods is far from clear. The most common explanation 76 Proceedings: Workshop on Fire, People, and the Central Hardwoods Landscape GTR-NE-274 Woodpecker clusters in the Daniel Boone National Forest Red-headed Woodpecker, Prairie Warbler, Indigo Bunting, has striking overlap with the distribution of Native American Bachman’s Sparrow) in response to management for Red- archeological sites (L. Perry, pers. comm.), which suggests cockaded Woodpeckers in Arkansas. (Hines 1999) found that Native American use of fire may have been important in similar response of the avian community in Kentucky. In both maintaining habitat for the woodpecker. studies some “interior” species (Black-and-white Warbler, Ovenbird) were found more frequently in control plots than in The Red-cockaded Woodpecker was apparently extirpated treatments. from Missouri by 1946 (Eddleman and Clawson 1987) and from Tennessee by 1994 (Nicholson 1997). Only one Avian Communities of Savannas, Barrens, individual native to Kentucky remains, although 16 and Glades in the Central Hardwoods Region individuals have been translocated to Kentucky from larger populations to support the recovery effort there (L. Perry, Savannas, barrens, and glades were important components pers. comm.). In each of these states the loss or decline of of the Central Hardwoods landscape before European