Effects of Silviculture on Neotropical Migratory in Central and Southeastern Oak Forests James G. ~ickson',Frank R. Thompson IIP, Richard N. ~onner',and Kathleen E. ~ranzreb~

Abstract - Avian communities that are associated with forest habitat attributes are affected by silvicultural and other stand influences. Some species have specific habitat requirements, whereas others occupy a broad range of vegetative conditions. In general, species richness and density are positively related to stand foliage volume and diversity. Bird density and diversity are usually high in young brushy stands, decrease in dense pole stands as canopies close and shade out understories, and are highest in . older stands with diverse foliage strata. Tree harvesting generally favors early successional species such as the Indigo Bunting, Prairie Warbler, and Yellow-breasted Chat, but some late successional forest species, such as the Black-and-white Warbler, persist with partial cutting. A few forest interior species, such as the Ovenbird, are less abundant in landscapes with cutting and forest fragmentation. Some species may have elevated nest parasitism and nest predation along forest edges. Recommendations for NTMB include: Maintain some large, old-growth stands; manage foresf habitat for NTMB; employ special measures for endangered or sensitive species; implement long-term monitoring; and develop more complete information through research regarding NTMB, population viability, and their forest habitat.

Central hardwood, loblolly-shortleaf pine, longleaf-slash-pine, and bottomland hardwood forests dominate the central and southeastern United States. These forests are managed by both even-aged and uneven-aged silvicukural systems. We review the impacts of silvicultural practices on neotropical migratory birds (NTMB), in forests of this region. We approach this topic by (1) identifying general relationships between birds and forest habitats and landscapes in the region, (2) assessing effects of forest management on the NTMB in each ecosystem, and (3) conclude with some management strategies that extend across forest types.

Bird Habitat increase with plant succession, and vegetation volume and structural diversity (Johnston and Odum 1956, Shugart and Bird species and communities in forest stands have specific James 1973, Shugart et al. 1975). But breeding bird habitat requirements and any changes to stand characteristics densities in regenerating oak-hickory or pine-hardwood influence stand suitability. Avian density and diversity generally stands are often similar to those in mature stands, with densities often lowest in mid-successional pole-sized lJsFs, sournem Fond upedmeot s&aon, Nacogdoches, TX. with little subcanopy (Comer and Adkisson USFS, North Central Forest Expetimenf Station, Columbia, MO. 1975, Comer et al. 1979, Dickson and Segelquist 1979, USFS, Southeast Forest Experiment Station, Clemson, SC. Yahner 1986, Thompson and Fritzell 1990). Tree harvesting drastically alters bird habitat and thus bird Effects of Even-aged Management comnunities. Generally, clearcutting results in a near complete mover of bird species while parlial removal of a forest overstory results in decreases in some species, increases in Regeneration Stands others, and little change in relative abundance of other species (Webb et al. 1977, Crawford et al. 1981, McComb et al. 1989, The first year after harvesting these stands may hav~ mompson et al. 1992). abundant herbaceous ground cover, but they quickly become dominated by tree regeneration from sprouting and advance regeneration resulting in as many as 25,000 stemslha (Gigrich 1971). The first year after clearcutting there is usually a drastic reduction in total bird numbers and a nearly complete turnover in species (Table 1). American Goldfinches and Field Sparrows Landscape Level Impacts often prefer cuts at this age because of the abundant grassffort, vegetation. As tree regeneration dominates the site, NTMB populations are affected by factors at scales larger Yellow-breasted Chats, Indigo Buntings, Prairie Warblers, than habitats or forest stands such as landscape (Freemark et al. Blue-winged warblers, Kentucky Warblers, Common this proceedings). For instance many forest interior migrants are Yellowthroats, White-eyed Vireos, Gray Catbirds, and absent from small forest fragments, and their absence is likely Rufous-sided Towhees occupy the stands. To the east due to lower repmductive success in edgedominated forest Chestnut-sided Wdlers and Hooded W~lersalso may be fragments (Faaborg et al. this proceedings, Robinson et al. this common In shelterwood and seed-tree cuts, and clearcuts with proceedings). While edge-related declines in reproductive residual live trees and snags, some mature-forest, success in fmgmented forests have been documented, the effects canopydwelling species may continue to use the stands. of edges created by timber harvest in predominately forested landscapes is unclear. Yabner and Scott (1988) reported higher nest predation rates of artificial nests in Pennsylvania forest with Sapling Stands clearcutting than with w clmtting. Many forest interior species remain abundant in managed forests (Thompson et al. During age 10 to 20 the stands are dominated by tree 1992), but it is possible that such forests are population sinks saplings with a closed canopy. At age 20 the number of stems where production is insufficient to compensate for adult has been reduced through mortality to 3,400-6,200Jha and the mortality Wlii 1988). Simulated populations of forest larger trees on good sites have reached 18 cm dbh (Gingich interior NTMB were lower in cut than in uncut forest landscape 1971). hhybirds typical of regenenting stands persist at lower (Thompson In Press). densities in these stands. Black-and-white Warblers, In uneven-aged stimds, selection cutting maintains a Wormeating Wahlers, and Kentucky Warblers seem to pprefer mature tree component at all times and does not create a the high stem densities and closed canopies this age class mosaic of Werent aged stands. This may benefit forest provides. Ovenbirds, Wood Thrushes, and Redeyed Vios may interior warblers because large tracts of forest with mature begin using stamds at this age. trees can be maintained. However, edge effects may occur in group selection openings which could be detrimental to local populations, since these openings can be widely dispersed Poletimber Stands throughout forests. Also, selection cutting will not provide habitat for species that require young stands, or a diversity From age 20 to 60 years 90% of the trees will die due to of even-aged stands. competition The canopy remains closed and there is little understory development. As a result, common species tend to be canopy nesters such as Redeyed-Vis, Scarlet Tanagers, Eastern Wood Pewees, and Wood Thrushes, or ground nesters such as Ovenbirds and Black-and-white Warblers.

CENTRAL HARDWOOD FORESTS Mature Stands Oak-hickmy forests dominate the central hardwood forests but give way to mixed hardwoods in the east and oak-pine The structure of matm forests varies widely throughout the fo'orests to the south (Eyre 1980, Sander and Fischer 1989). These region Depending on soils, geology, climate, and geogragl-y; forest have been predominantly managed by clearcutting in the mature stands may have sparse to dense gmundcover and past. However, recently there has been an increase in the use of understory. Decay and deaths of large trees result in cavities, shelterwood and selection cutting, 'particularly on public lands. snags, and tree fall gaps not present in short rotation stands. Table 1. -Abundance of neotropical migratory birds in central However, because of widespread logging, burning, and gmzhg hardwood forests: of this region in the late 1800s and early 1900s much of the Species Stand ~~e~ mature forest in the rkgion is even-aged mging from 60-100 R S PMGT years old. There are no lmown obligate old-growth NTMB in these forests. The most abundant species throughout the region Whip-poor-will UUUU in matwe forests is the Redeyed Vim. Other abundant, or Ruby-throated hummingbil NN?N common species in this age class include Eastern Wood Pewee, Acadian flycatcher CANA Acadian Flycatcher, Blue-gray Gnatcatcher, Ovenbird, Eastern wood-pewee UANA Wormeating Wdler, Scarlet and Summer tanager, and Blue Eastern phoebe UUNU Jay. In oak-pine stands Pine Warblers and Yellow-throated Great-crested flycatcher CCCC Warblers are common Carolina wren UN?? Blue-gray gnatcatcher CCCC Eastern bluebird NNNN Changes in Stand Composition Wood thrush CCUC Gray catbird CCNN?N White-eyed vireo CCNN?N Regeneration cuts do not usually greatly alter tree species Yellow-throated vireo NNNUNU composition in central hardwood stands because they are Red-eyed vireo UUAAUA naturally regenerated largely from advance reproduction and Blue-winged warbler ACNN?N stump sprouts. Small changes in tree species composition have Golden-winged warbler CUNN?N little effect on breeding birds because it is generally believed Northern parula NNUCNC most birds select breeding habitat by vegetation structwe. Past chestnut-sided warbler CCNN?N practices of converting low quality hardwood stands to pine have Yellow-throated warbler NNUUNU been largely abandoned on public lands but may persist on some Pine warbler NNCCNC private lands. Pine plantations genedy support a lower density Prairie warbler ACNN?N and diversity of breeding birds because of their structural Black-and-white-warbler CCCCCC simplicity. Worm-eating warbler UCCCCC Ovenbird UCCCUU Louisiana waterthrush NUCCCC Common yellowthroat AUNN?N Uneven-aged Management Kentucky warbler ACUUAC Hooded warbler CCUUCC Yellow-breasted chat ACNN?N Single and Multi-tree Gaps Orchard oriole UNNNNN Summer tanager CCCACC Canopy gaps resulting from the harvest of single trees or Scarlet tanager UUCAUA groups of trees provide habitat for a variety of migrant birds. Indigo bunting ACUUAC Species such as the Hooded Warbler and Indigo Bunting appear Rufous-sided towhee AUNNCN to be able to make use of small gaps typical of single tree Field sparrow ANNN?N selection while other species such as Yellow-breasted Chats, Brown-headed cowbird ACCCCC American goldfinch UNNNNN Blue-winged Warblers, and Prairie Warblers require large Blue jay CCCCCC openings more typical of clearcuts (E Thompson pea. obsv.). American crow UUUUUU There is a dearth of informalion on the area sensitivity of species requiring early successional forest or gaps. These canopy gaps 'Includes oak-hickory, mixed hardwood, and oak-pine forest types. also may be amactive to cowbirds and predators. Habitat associations based on Conner arid Adkisson 1975, Conner ef a/. 1979, Dickson and Segelqujst 1979, Dickson et a/. 7980, Evans and Kirkman 7987, Yahner 7986, Thompson and Frifzell 7990, Thompson et at. (7993, Whitehead unpubl. data, Robinson unpubl. Change in Stand Structure data. R = regeneration, S = sapling, P = poletimber, M = mature, G = group selection, T = single tree selection. Uneven-aged stands have a well developed understory and A = abundant, C = common or regular, P = present, U = uncommon, sub-canopy because of frequent canopy gaps. The presence of l\r = not present. several well developed vegetation levels and more complex - - - habitat structure than similar aged even-aged stands could result Rotations for central hardwood stands managed for sawtimber in higher within-stand bird species diversity. For instance, in are usually 60-120 years. At age 80 dominant trees will mge Illinois selectively cut stands contained NTMB associated with from 30-46 cm dbh If left undistubed these stands will slowly mahm forest habitats as well as some young second growth become uneven-aged as they age and individual trees die. species. There are indications that uneven-aged stands may be poor Table 2. - Abundance of neotropical migratory birds in habitat for some IMture forest species. There were fewer Loblolly-shortleaf pine forests'. ovenbirds in mature uneven-aged than in even-aged forests; and Species Stand ~ge~ all redeyed vireo males in a selectively cut stand in Missouri were unmated, compared to 80% mated in a neahy uncut mature RSPMO forest (Ziehmer 1992). Whip-poor-will N~NNN? Ruby-throatedhummingbirdu U U U U LOBLOLLY-SHORTLEAF PINE FORESTS Acadiin flycatcher NNUCC Eastern wood-pewee NUPCA I he loblolly-shortleaf pine ecosystem of the rolling porlions Eastern phoebe NNNN? I of the Gulf Coastal Plains, the Piedmont, and portions of Great-crested flycatcher N N U P C Appalachian, Ouachita, and Ozaik Mountains, is characterized Blue-gray gnatcatcher NNUGA by a species composition of at least 50 percent (either Eastern bluebird UUNNP ' loblolly, shortleaf, or a mix). Associated species include oaks, Wood thrush NNUCA ' American robin NUUUU hickories, sweet- blackgum, winged elm, and red maple Gray catbird UUNUU (Garrison et al. 1977). The degree of hardwood inclusion in White-eyed vireo UAPUC these forests is largely determined by past frequency and Yellow-throated vireo NNUAC intensity of natural and prescnied fire. Red-eyed vireo NUCAA Harvesting of loblolly and shortleaf pines and subsequent Blue-winged warbler NNNNN stand regenemtion can be successfully accomplished with both Golden-wingedwarbler N N N N N even- and uneven-aged silvicultural techniques (Baker and Northern parula NNUUC 1 Balmer 1983, Lawson and Kitchens 1983), In modem managed Chestnutsided warbler N N N N , N Pine warbler NNCAA 1 forests prembed fire, intensive mechanical site preparation, I pine seedling planting, and herbicides are used to enhance or Prairie warbler CANNU maintain the pine component. Black-and-white-warbler N U C C C I Worm-eating warbler NNCCC. Chuck-will's-widow UUUUU Ovenbird NNUCC Bird Communities Louisiana waterthrush NNNPP Kentucky warbler NUCPP The loblolly-shortleaf pine ecosystem provides habitat for Hooded warbler NUCAC a vev diverse myof birds, many of which are neotropical Yellow-breasted chat CAUNP migrants. Stand structure and the proportion of pines and Summer tanager NNUCC hardwoods are important determinants of avian communities. Scarlet tanager NNUUU Indigo bunting NAPUP Rufous-sided towhee NPCCP Even-aged Management Field sparrow CUNNN Brown-headed cowbird PCPPN Early succession stands are stmcturally simple with little American goldfinch UUUUU verlical foliage diversity. As stands develop different stages of Blue grosbeak CUNNN are birds. Blue jay UUCAC stand development attractive to different species of American crow UUPCC

Seedling and Sapling Stands Includes oak-hickory, mixed hardwood, and oak-pine forest types. Habitat associations based on Conner and Adkisson ~~di~~season--~learcuttingtoday and over the past (1975), Conner et al. (1979), Crawford et al. (1981), Yahner (1986), Thompson and Fritzell (1990), Thompson et al. (1992), 'eveEd decades has produced habitat conditions similar to Hammel (in Press), Robinson (unpubl. data), Mitehead (unpubl. those cmted by large landscape level disturbances. An array of data). mtropicd migrants (e.g., Prairie wa1b19rs, Field Spmws, Blue Grosbeaks,) are atlracted to the youngest stands during the * R = regeneration, S = sapling, P = poletimber, M = mature, breeding season (Noble and Hamilton 1976; Meyers and O = OklgrO*h- Johnson 1978; Coriner et d. 1979,1983; Dickson and &elquist 1979; Dickson et al. 1980, 1984; Childers et al. 1986). When A = abundant, C = common or regular, P = present, U = un,,,n, un,,,n, = not present. the stands exceed three years old the number of neotropical ,,, migrants using them for breeding sites blooms. Indigo ~un&~s, Painted Buntings, Whiteeyed Vios, Yellow-breasted Chats, the only early succession species that may persist into the more and Common Yellowthroats are some of the more common mature stages of forest growth, depending on the availability of species found in the well developed shrubby vegetation of young open patches where sufficient light has penetrated to stimulate clearcuts. growth of understory foliage for nesting sites. Wood Thrush, As these even-aged stands develop, foliage patchiness in Redeyed Vis, Black-and-white Wahlers, Eastern Wood the 0-3 m layer increases and some trees grow to 4-5 m. A few Pewees, Great Crested and Acadian flycatchers, Pine, Hooded, additional species of neofropical migrants begin to occupy the Kentucky warblers, Summer Tanagers, and Blue-gray older sapling lobloIly-shortleaf stands. In the northern portion Gnatcatchers are now abundant (Noble and Hamilton 1976; of the ecosystem Ovenbirds, Rufous-sided Towhees, Meyers and Johnson 1978; Conner et al. 1979, 1983; Dickson Black-and-white Warblers, and American Redstarts begin to and Segelquist 1979; Dickson et al. 1980, 1984; Cbilders et al. appear. Further south, only Black-and-white Warblers me added 1986). Yellow-throated Warblers, Northern Parulas, and in any numbers. During this later sapling stage of succession Yellow-billed Cuckoos are also present. Pine Warblers are some species, such as Field Sparrows and Blue Grosbeaks, begin attracted to the pine foliage, whe~asmost of the other species to disappear as foliage over-grows bare ground and grasses. are primarily dependant on deciduous foliage that has developed. Winter--Many neotropical mi- use young and older Winter-Matm loblolly-shortleaf pine and pine-hardwood sapling aged loblolly-shortleaf pine stands as wintering habitat forests provide winter cover and food for American Robins, (Noble and Hamilton 1976, Dickson and Segelquist 1977). Hermit Thrush, Goldencrowned and Rubycrowned kinglets, Winter Wrens, Brown Thrashers, American Robins, Hermit Black-and-white Wihlers, and White-throated Sparmws (Noble Thrush, Eastern Bluebirds, Rubycrowned and Goldencrowned and Hamilton 1976, Dickson and Segelquist 1977). kinglets, Pine and Yellow-nunped warblers, Darkeyed Juncos, and Field, Song, Lincoln's, and White-throated spamws all use young pine plantations during winter. Management Activities Affecting Bird Communities

Pole Stands Management activities such as site preparation, hefiicide use, and thinning, sect stand structure and impact the avian Breeding season-When young pines and hardwoods reach fauna. the pole stage (12-25 years) most of the early succession Site preparation--The intensity of mechanical site migrants are no longer found. The taller foliage gradually fonns preparation prior to planting affects the amount of hardwood a canopy that reduces light penetration through to the understory. vegetation that regenerates with the pine seedlings. Intensive A few Whiteeyed Visand Yellow-bl.easted Chats remain in K-G blading (bull-dozer blade) and chopping of roots will open areas, such as wind rows, (Noble and Hamilton 1976; reduce the amount of hardwood regeneration substantiaUy. Less Meyers and Johnson 1978; Comner et al. 1979, 1983; Dickson intensive site prepition, such as prescribed burning, permits and Segelquist 1979; Dickson et al. 1980, 1984; Childers et al. more hardwood vegetation to survive and grow along with the 1986). Kentucky and Black-and-white warblers become fairy pines. Bird species diversity will be reduced as the amount of common Red-eyed Vios, Womeating, Pine, ine, Hooded hardwoods in all pine stands decreases. Herbicides--Hehicides warblers, Yellow-billed Cuckoos, and Summer Tanagers begin sometimes are used to kill hardwoods that compete with pines to appear. Occasionally, Wood Thrush and Brown Thmhers are in young pine plantations. Reduction of hardwood vegetation detected in such stands. would set back stand development and associated bird Winter--Both pine and pine-hardwood pole sized stands communities. used during winter by neotropical migrants, such as Brown Thinning-Thinning opens the canopy, releasing the Creepers, Winter Wrens, Hermit Thrush, Eastern Bluebirds, remaining pines for increased diameter growth, and permits Goldencrowned and Ruby-crowned kinglets, Black-and-white, more light into the understory which promotes understory Pine, and Yellow -rumped warblers, and White-throated vegetation (Blair and Enghardt 1976, Blair 1982). Spmws (Noble and Hamilton 1976, Dickson and Segelquist Thinning is beneficial to some neotropical migrants and has 1977). a significant positive influence on bird abundance and species richness during the breeding season (Chritton 1988). Indigo Buntings, Pine Warblers, and Brown-headed Cowbirds increased Mature LobloIlyShortleaf Pine Stands in abundance following thinning of a loblolly plantation in Texas. White-eyed Vios, Wormdg Warblers, and Hooded Breeding season--After 35 to 50 years the developing stand Warblers were negatively affected, whereas Black-and-white begins to achieve a few characteristics of maturity. h overstoIy Warblers appeared to be unaffected by thinning. canopy is now present, but midstow and understoly foliage can Thinning also affects migrants that winter in the be present also, depending on how much light filters through loblolly-shortleaf pine ecosystem in the South (Chritton 1988). the canopy. A few Whiteeyed Vios and Indigo Buntings are During winter, bird abundance and species richness was higher in thinned pine stands than in unthinned starads. Pine Wdlers, added to the overall bird community. But, cowbird parasitism ~ubycrownedand Goldenrnwned kinglets, and Darkeyed may increase and the few mature forest species that prefer a JWCOS increased in numbers following thirming in Texas. continuous canopy may be negatively impacted. Rotation ages--Longer rotations of 70 to 80 years and longer perrnit some old-growth attributes to develop provide habitat for species of neotropical migrants that prefer mature LONGLEAF, SLASH, AND pine forest, such as Redeyed Vireos, Northern Parulas, and LONGLEAF-SLASH PINE FORESTS Hooded, Pine, and Yellow-throated w&lers. Shorter saw log rotations of 35 to 50 years would provide habitat for some The longleaf-slash pine ecosystem, a subclimax type mature forest species but probably at lower densities then the mahtahed by fire, generally occurs along the lower coastal plain longer rotations. Pulp wood rotations of 20-30 years provide of the southeastern U.S. Agriculhual conversion, intensive habitat for early successional species when the stands are young, timber management, and substantial declines in frequency and but has limited suitability for neotropical migrants requiring intensity of fires have resulted in drastic reduction in the longleaf stands beyond the pole timber stage. pine forests. The natural range of slash pine is more restricted (South Carolina to Central Florida and southeast Louisiana), although it has been planted extensively as far north as North Seed-tree and Shelterwood Harvesting Carolina and west to eastern Texas. Most of the typical longleaklash pine cover type is found in the flatwoods of Seed-tree and shelterwood harvesting leave uncut mature Florida and Georgia Slash pine seedlings im more shade pines to serve as seed sources and in shelterwood as shelter for tolerant than those of longleaf pine; hence, slash pine may begin the developing stands. These large residual pines can be of value as an understory and remain subordinate to the longleaf pine to early and late successional neotropical migrant birds (Hall overstory. Slash pine regenemlion is eliminated by fire and 1987). without fire the encroachment of hardwoods will progress until Breeding Season--Nmtropical migrants using seed-ke and they predominate and exclude the pines. Longleaf pine seedlings shelterwood cuts during the breeding season are: Eastern in the grass stage can tolerate and benefit from prescribed Kingbirds, Acadian Flycatchers, Pine Warblers, Prairie Wdlers, burning for brownspot control. Both slash and longleaf pine Yellow-breasted Chats, Wilson's Warblers, Hooded Wahlers, seedlings growing in height are vulnerable to fire, but both Orchard Orioles, Indigo Buntings, and Chipping and Field species are somewhat immune when they attain a height of about sparmws. 5m Winter--During winter, seed-tree and shelterwood hawesting again provide habitat for both early and late succession bird species. Neotropical migrants using cuts Bird Habitat included American Kestrels, Ruby-crowned Kinglets, Yellow-rumped Warblers, Common Yellowthroats, Red-winged Younger forests tend to have moderate bird species diversity Blackbirds, Purple Finches, Pine Siskins, American Goldfinches, PSD), the initial BSD tends to decrease by the time the forests and Grasshopper, Henslow's, Lark, Field, White-crowned, are 16-20 years old, after which it increases to maimity (Childers White-throated, Swamp, and Song sparrows (Hall 1987). gt d. 1986, Noble and Hamilton 1976, Repenning and Labisky Unfortunately, the benefits for bird species richness gained 1985). by the presence of the residual pines is lost completely when Avifauna in pine forests is substantially influenced by the the residuals are removed following the establishment of the presence and extent of hardwoods (Johnston and Odum 1956, new pine stand. Dickson and Segelquist 1979). Pine stands normally contain numerous hardwood species and removal influences bird community composition Burning to maintain pine forests can Uneven-aged Timber Management have deleterious effects on birds using the hardwood cornpoilent for cover, nesting, or foraging. However, for ground foragen, There is extremely little published information on the bird the effects of buming may be beneficid as fire reduces the litter communities that inhabit stands managed under single-tree or thereby exposing seeds that then become available for food. group selection harvesting, so mntly we can only speculate Slash pine sites in southeastern Georgia that contained based on data from seed-tree and shelterwood cuts, and heavily hardwoods supported 17 species that either were absent or ?binned stands. Species that use early successional stands, small occurred in low densities in the pure slash pine and, therefore, gaps within fo~sts,and "edge" would probably be favoml by bardwoods were important in maintaining the BSD (Johnson single tree selection harvesting. In group selection larger gaps and Landers 1982). up to 1 ha would have a similar but more pronounced effect on Several studies have examined longleaf and slash pine the avian community than single tree selection. Yellow-breasted stands of various ages to determirae how avian community Chats, Indigo Buntings, and Prairie Whlers, will probably be structure may change as the stands mature. In slash pine stands Table 3. -Abundance of neotropical migratory birds in longleaf and slash pine forests'. in southeastern Georgia in relation to stand age and regeneration method, Johnson and Landers (1982) found that bird numbers Species Stand age2 tended to be lowest in the 1-year-old slash pine plantations, RSPMO increased in the 2-6-year-old stands, and then declined again Longleaf Pine until approximately mid-rotation (16 years). Once stands passed Common nighthawk p3puuu mid-rotation ages (16-28yeas), the initiai stand treatments such Chuck-will's-widow UUU Eastern wood-pewee UCC as site preparation or whether naturally regenerated or planted, Acadian flycatcher P P had no detectable effect on the avifauna. Five species of breeding Great-crested flycatcher U P NTMB were found in the fallow areas, with only the Bluegray Eastern kingbird P P Gnatcatcher being abundant, and five species were regularly Purple martin U observed in the regenerating areas. Of the seven species found Barn swallow U in the seedlinglsapling stage, the Indigo Bunting, Common Prairie warbler C Yellowthroat, and Ruby-throated Hummingbird were either Summer tanager UUCC abundant or common The Common Yellowthroat, Bluegray Blue grosbeak P P Gnatcatcher, and Eastern Wood Pewee were the most commonly SIash Pine observed of the 10 NTMB in the pole stage. Yellow-billed cuckoo UUU Also, regenerating longleaf pindslash pine stands in the Ruby-throated hummingbird C C grass forb stage are year round habitat for the Bachman's Eastern wood-pewee C Sparrow, Northern Bobwhite , Mourning Dove and Eastern Acadian flycatcher C Great-crested flycatcher PCC Bluebird; breeding habitat for the Common Nighthawk; and Eastern kingbird P wintering habitat for Henslow's and Leconte's sparrows. The Bewick's wren C sapling stage is suitable habitat for the Common Nighthawk, Blue-gray gnatcatcher P P Common Yellowthat, Indigo Bunting, and a 'number of White-eyed vireo C permanent resident species. Yellow-throated vireo P The effects of prescribed buming on a 20 year-old slash Yellow-throated warbler C pine stand resulted in a drastic decline in ground cover and shrub Prairie warbler UUU foliage in Everglades National Park, Florida (Eden 1976). Common yellowthroat CPPP However no si@cant difference was detected in NTMB or Yellow-breasted chat C foraging guilds after the fire, possibly because of individual Summer tanager C CCC attachments to home range and familiar foraging sites. Blue grosbeak P P Harris et al. (1974) compared site preparation techniques in Indigo bunting CCPUU LongleafSlash Pine three 9-year old slash pine stands with a naturally regenemted Osprey U U slash pine stand and with a mature longleaf pine stand that had American swallow-tailed kite U U been prescribed burned the previous year. The average number Yellow-billed cuckoo P of birds observed per transect sampled was higher in areas that Common nighthawk PUUUU had undergone low intensity site prepamtion than in matwe slash Chuck-will's-widow UUU pine; however, the number of species observed per transect was Ruby-throated hummingbird U significantly lower. There were nine times more birds in mature Eastern wood-pewee UCC longleaf pine stands than in the low intensity site prepared slash Great crested flycatcher P U P P pine stands and 60 times more than in the high intensity treated Eastern kingbird UUUUU site p~paredstands. Purple martin U Repenning and Labisky (1985) compared the avian Barn swallow U community in 3 nahmlly regenerated longleaf pine stands (>50 White-eyed vireo C C Yellow-throated warbler U U U U years) to slash pine plantations of 4 different ages (I-, lo-, 24; Prairie warbler UUUUU and 40-year-old stands) (3 stands per age group). Density of Common yellowthroat UPPCC breeding birds was highest in the IongIeaf pine stands (288 Yellow-breasted chat U U birds/km2) and oldest slash pine stands (149 birds/km2). Summer tanager UUPP Longleaf pine stands supported more breeding birds, a higher Blue grosbeak P P biomass, and greater species richness than any of the slash pine Indigo bunting . CCPUU stands. Cavity and ground nesters increased in both density and From Hamel et at. 119821. Johnson and Landers 11982). number of species from the youngest to oldest stands and were O'Meara el al. (19853, ~ebnnin~and Labisky (1985), zdkson highest in the longleaf pine forests. Of the species of (1991), and Hamel (in press), 7 R = regeneration, S = sapling, P = poletimber, M = mature, 0 = old neotropical migrant bmding birds, 2 species wexc found in growth. 1-year old stands (13 birds per krn2), 4 in 10-year old stands A = abundant, C = common or tegular, P = present, U = uncommon (19 birds per km2), 2 in 24-year old stands (22 birds per km2), 4 in 40-year old (33 bids per lcm2), ad5 in malure longleaf pine The avifa~naof oak-g~mcy~re~~forea is abwand forest (33 bitrls per lun2). Wmtering birds were most numemus in diverse. Habitat suitability for bird communities and the youngest slash pine stands, perhaps the result of the abm depends on stand stmX& and other factos. of seed-producing gmses and fobs (Repenning and Labisky 1985). They concluded that conversion of mature longleaf stands to slash pine plantations did not accommodate the bird community Breeding Birds , associatsd with the mature habitat. O'Meara et al. (1985) and Rowse and Marion (1980) There are a variety and ab- of bnxdhg birds in the examined the same 3 amsof flatwoods in Florida containing slash maburr: bottomland hardwood forests of the South. In a Louisim and scattered longleaf pine tms. The areas had been harvested 35 floodplain fo~stabout half of the number and species of breeding years prior, allowed to regenerate mludly, and fire had been season birds were neotropical mi- and about half were excluded for 20 years. Pinz stands in 2 of the axas were clearcut pennanent residents (Dickson 1978b). The proportion of and were monitored before culling and for 3 years afterwards, neotropical migmt breede~was lower than in molie maberly and Unharvested slash pine 35-years-old, regenemting slash pine seasonally harsher climates. (mnUy clm), cypress and edge ams (< 10 m from the Oak-gumcypress forests are special habitat for many species interhce of 2 other habitats) were compand The Great-cmmd of birds pickson 1978% 1988, Hamel et a1 1982). A survey of Flycatcher, White-eyed Vi,and Common Ydowthmat were the breeding bird censuses from wen mature stands (Dickson et aL NTMB in the 35-year old stands. For the 3 years subsequent to 1980) showed that Yellow-billed Cuckoos, Acadian Flycatchers, clearcuthg, harvested habitats contained low spring and summer and Red-eyed Vireos were consistently abundant in densities with only a few species adapted to early-successional oak-gumcypn=ss habitat Other species regularly inhabit these vegetation Thm NTMB species (Common Nighhawk, Common st.,and some have special aflinities for this habitat (Table 4). Yellowthroat, and Blue Grosbeak) were present at densities less Table 4. - Neotropical migrant breeding bird specvies present in than 7/bm2. Winter densities in clearcut habitat were higher southeastern oakgumcypress forests (from Hame! et resulting from the presem of wintering flocks of American Robins al. 1982 and other sources) and Red-winged Blackbirds and mixed species flocks of spanavs. The findings of this study are similar to that of Repenning and American anhinga Blue-gray gnatcatcher Green-backed heron Whiteeyed vireo Lab& (1985) in .that an hmedi& result of cleamttmg slash Great blue heron Yellow-throated vireo pine was a qkment of birds by species adapted to early Little blue heron Red-eyed vireo successionaI stages, a lower density of breeding bids, reduced Cattle egret1 Black-and-white warbler BSD, and a much larger winkring than bmxbng pophiion. Great egret Prothonotary warbler Snowy egret Swainson's warbler The above studies evaluated the effects of even-aged Tricolored heron Worm-eating warbler management on the avifaum However, little work has been done Blackcrowned night heron Bachman's warbler - in lat ti on to the of birds to uneven-aged h&g Yellow-crowned night heron Northern parula techniques. Single tree selection hawesling opens up the forest Wood stork Black-throated green Glossy ibis warbler canopy, wimg small gaps in the forest floor. In the group selection White ibis Yellow-throated warbler method, cuts are 0.2 to 0.8 ha in size and thus create Hooded merganser Prairie warble? numerous small gaps in the forest In both cases, piesthat use American swallow-tailed kite Ovenbird edge habitats or small gaps will benefit. However, it is not clear if Mississippi kite Louisiana waterthrush Cooper's hawk Kentucky warbler the habitat mentation created by this approach will be Bald eagle Common yellowthroat2 detrimental to birds quiring large tracts of udimrbed forest Osprey Yellow-breasted chat2 Purple gallinule Hooded warbler Common moorhen American redstart Mourning dove1 Eastern meadowlark1 OAK-GUM-CYPRESS FORESTS Yellow-billed cuckoo Red-winged blackbird Chimney swift' Brown-headed cowbird1 Oak-gum-cypress forests (also called bottomland Ruby-throated hummingbird Orchard oriole hardwoods) oocur on mesic to hydric sites along streams or Belted kingfisher Northern oriole from Great-crested flycatcher Summer tanager rivers Viato eastern Texas and up the Mississippi Eastern phoebe' Blue grosbeak2 river to Indiana. Dominant vegetative communities of this Acadian flycatcher Indigo bunting2 complex m closely associated with sites which are determined Eastern wood pewee Painted bunting2 mainly by soils, elevation, and hydroperiod. Oak-gumcypress Barn swallow' Rufous-sided towhee2 forest area has declined, especially in the Mississippi River Delta Purple martiniwood thrush where land was converted to soybeans, cotton, and pasture. ' Assodated wiN, human altered non-forest habitat. Recently there have been some effoa to restore bottomland * Associated with eady successional stands. forests that have been converted to agricultud uses and to protect what remains. Many long-legged waders nest and forage in aquatic woodlands. including Yellow-bellied Sapsucken, American Robins, Hennit Wood Storks, which ax now endangered, nest in tall cypress Thrushes, Ruby-crowned Kinglets, and Orange-crowned and hardwoods and feed in associated aquatic systems. Warblers. Several migratory raptors inhabit bottomland hardwoods, such as the Mississippi and Swallow-tailed kites, Cooper's Hawks, Bald Eagles, and Ospreys. Also, Purple Gallinules an8 Common Moorhens are found in appropriate aquatic habitat. Silviculture and Bird Communities Acadian Flycatchers are strongly associated with forested wetlaads (Shugart and James 1973, Smith 1977), and Wood Although specific studies of bird co-ty changes Thrushes are a common breedmg bird in the rnesic sites. =lad to silvk&ural practices in oak-gumcypress forests are Whiteeyed Vireos are common in this lype in low, shrubby generally lacking, some information may be presented based on foliage and Redeyed Visin canopy foliage. There are many general habitat relationships of bird species and from study warblers in this habitat, some with special affinities. results of the other habitats in the South. Dmtic habitat Prothonotary, Swainson's, Northern Parula, Kentucky, and dteration in oak-gum-cypress forests will influence bird Hooded are strongly associated with this habitat. Prothonotary community composition Conversion of mature bottomland Warblers nest in cavities, often over water. Northern Parulas forest stands to other land uses, of course, will drastically alter construct their nest with Spanish moss in moist woods. The bird communities. Marry bottomland stands and their associated Swainson's WarbIer is primariIy associated with understory bird communities have been lost to reservoirs. This conversion thickets of southern river floodplains and the southern probably only benefits a few aquatic species such as cormorants Appalachian Mountains Meanly 1971). The habitat of the rare and perhaps some cavity nesters in the short term if the trees Bachman's Wahler is bottomlands and headwater swamps are not cleared before inundation and water-killed snags are left. subject to distwbances (Hooper and Harnel 1977). Both Conversion of bottomland forests to agricultural land also resuIts Swainson's and Bachman's warblers are associated with cane in elimination of the forest associated avian community. thickets, which were once extensive in southern bottomland Tree harvesting and regeneration in bottomland forests is a forests (Meanly 1971, Remsen 1986). Kentucky and Hooded habitat alteration that results in changes to the bird communily. warblers are usually found in the moist understory of bottomland The replacement of mature stands of mixed hardwoods by hardwoods @ickson and Noble 1978). Other wa&lers often hardwood plantations alters bid communities. These plantatiois found in mature stands include Black-and-white, Womeating, and natural stands of pure black willow or cottonwood lack Yellow-throated, Ovenbird, American Redsm and LouisiaPla vegetative diveity and support fewer birds ada less diverse Wateahnrsh (Hamel et al. 1982). bird community than natural mixed stands (Wesley et al. 1976). Bird communities are related to age and development of But in areas where the majority of land is in mature mixed forest stands (Shugart and James 1973, Dickson and Segelquist stands and tk plantations repment a small land commitment, 1979). Breeding birds discussed previously have been those the overall beta (landscape) bird diversity of an area could be associated with mature stands of mixed species, and these would increased because of the birds associated with early successional also generally be present in middle-aged stands. Young stands studs that inhabit the plantations. ln Mississippi, Red-winged would have mostly a merent bird composition. In the earliest Blackbirds, Common Yellowthroats, Yellow-breasted Chats, stages of hardwood stand development the Dickcissel and Northern and Orchard orioles, Rufous-sided Towhees, and Red-winged Blackbird would be characteristic species (Weinell Warbling Viswere common in plantations, but not in natural 1989). Neotropical migrant birds typifying the avian community stands (Wesley et al. 1976). in young brushy stands include the Yellow-bmed Chat, Indigo Clearcuts with natural regeneration generally would favor Bunting, Painted Bunting, Prairie Warbler, Common edge species such as Wood Pewees and early successional Yellowthroat, adWhiteeyed Vireo. species such as Indigo Buntings, Prairie Warblers, White-Eyed =reos, and Yellow-breasted Chats (Dickson and Segelquist 1979, McComb et al. 1989, Thompson et al. 1992). Winter Birds Harvest regimes in which some trees are harvested and some left, such as improvement cuts or thhings would have a Winter is an important period for many birds. Mature less drastic effect on bird communities than clearcuts and would oak-gumcypress forests provide critical habitat for wintering favor early successional and edge species. Partial cuts or small birds, and support very abundant wintering bird communities clearcuts usually result in higher bird diversity and most mature (1400 - 2000/ km2, Dickson 1978b). Most of the species are forest associated birds remain in forested stands whe~some permanent or winter residents, such as Common Grackle and mature trees or stands remain- Understory vegetation growth in White-throated Spam>~v,that do not winter in the tropics. But the opened st& would favor umferstory associated species these mature bottomland forests are regular habitat for several such as Kentucky and Hooded ~7art,lers@ickson and Noble species that winter from sothem forests into the mpics, 1978, McComb et al. 1989, Thompson et al. 1992). A few forest interior species associated with closed canopy stands are regenerated, and maintaining both coniferous and forest would dwindle with tree harvest and stand opening. deciduous components of mixed stands. An extensive Studies have shown Ovenbiid and Wood Thrush abundance monitoring program should be implemented which O-acks bird were negatively correlated with stand harvest (Crawford et al. species abundance and viability over the long term. 1981, Webb et al. 1977). Research of NTMB and their forest habitat should be Rotation age also would affect bird ~ommunitycomposition expanded to develop more complete information for in forest stands. Short mtations would favor early successional management of NTMB. The different forested ecosystems, their species, whereas long rotations should favor cavity using species components and function, should be explored more fully. such as the Great-crested Flycatcher and canopy associated Species density may not always be a suitable measure of habitat species such as the Redeyed Vireo, Yellow-throated Vireo, quality (Van Home 1983). A better understanding of species Northern Pazula, and Summer Tanager @ickson and Noble demographics, including productivity, cowbird parasitism, and 1978). nest predation is essential. Moreover, additional information relating avian communities to forest composition, distribution, fragmentation and various silvicdlure pmctices is needed to ensure the future of sensitive NTMB. MANAGEMENT RECOMMENDATIONS

NTMB communities are determined by local habitat factors as well as landscape composition At a landscape level the single LITERATURE CITED most important consideration is to maintain large areas in breeding and wintering forest habitats to provide for large Baker, J. B., & W. E. Balmer. 1983. Loblolly pine. Pp. 149-152 NTMB populations and minimk numbers of cowbirds and irr Silvicultural systems for the major forest types of the predators associated with agricultural, subwban, and urban land United States P.M. Burns, tech compl.). USDA For. Serv. uses. At the habitat level the most basic management step is to Agric. Hdbk No. 445. maintain native ecosystems. Management should promote rare Blair, R. M. 1982. Growth and nonstructural carbohydrate ecosystems and habitats required by threatened or endangersd content of southern browse species as influenced by light species and regional species of high management wmem A intensity. J. Range Manage. 35: 756-760. high priority in southeastern forests is to protect existing Blair, R. M., & H. G. Enghardt 1976. Deer forage and ovektory old-growth stands and corridors, and to allow new old growth dynamics in a loblolly pine plantation. J. Range Manage. 29: stands to develop. Restoration and maintenance of natwal 104-108. ecosystems that have been substantially reduced or altered, such Childers, E. L., T. L. Sharik, & C. S. Adkisson. 1986. Effects as longleaf and oak-gumcypress forests, should be accelerated. of loblolly pine plantations on dynamics in the NTMB species exhibiting popdation declines and identified as Virginia Piedmont. J. Wildl. Manage. 50: 406-413. species of high management concern breed in all stages of forest Chritton, C. A. 1988. Effects of thinning a loblolly pine succession (Hunter In Press, Thompson et al. In Press), so a plantation on nongame bird populations in east Texas, MS. diversity of successional stages should be provided. Unless Thesis. Stephen F. Austin State Univ., Nacogdoches, Texas. specific concerns dictate otherwise, both selection cutting and Conner, R. N., & C. S. Adkisson 1975. Effects of clearcutting even-aged management should be used to create small openings on the diversity of breeding birds. J. For. 73: 781-785. for gap species, large openings for early successional forest Comer, R. N., J. G. Dickson, B. A. Locke, and C. A. Segelquist. mi-, and a balanced age-class distribution to mainbin 1983. Vegetation characteristics important to common sulEient mature forest habitats. This range of opening sizes in east Texas. Wilson Bull. 95: 349-361. more closely imitates the range in size of natural openings or Comer, R. N., J. W. Via, & L D. Phr. 1979. Weds of disturbances in forests (Hunter 1990) than the use of any one pine-oak clearcutting on winter and breeding birds in regeneration practice. Where late successional, area or southwestern Virginia. Wilson Bull. 91: 301-3 16. edge-sensitive NTMBs are a concern (e.g. Redeyed Vireo, Crawford, H. S., R. G. Hooper, & R. W. Titterington 1981. Ovenbird, Pine Whler, etc.) some blocks of mihgmented Songbird population response to silvicultuml practices in forest should be set aside from timber harvest, larger central Appalachian hardwoods. J. Wildl. Manage. 45: regeneration cuts on longer rotations used in even-aged systems, 680-692. and single tree selection favored over group selection Even-aged Dickson, J. G. 1978a. Forest bird communities of the bottomland systems should be used to provide young forest habitats for early hardwoods. Pp. 66-73 in Proceedings of the Workshop: successional species (e-g. Prairie W&ler, Yellow-breasted Chat, Management of Southern Forests for Nongame Birds (R. M Blue-winged Warbler, etc.) because openings created by DeGraff, tech coord.). U.S. For. Ser. Gen Tech Rep. SE-14. selection cutting may be too small for many of these species. Dickson, J. G. 1978b. Seasonal bird populations in a south Other stand level practices that will maintah NTh4I3 community central Louisiana bottomland hardwood forest. J. Wi. viability include retaining live cavity trees and snags when Manage. 42: 875-883. Dickson, J. G. 1988. Bird communities in oak-gum-cypress Hunter, W. C. In Press. Species and habitats of special forests. Pp. 51-62 in Bird Conservation 3. International concern within the southeast region h Proceedings of a Council for Bird Preservation (J. A. Jackson,. Ed.). workshop on the stzitus and management of neotropical University of Wisconsin Press, Madison. migratory birds. Rocky Mountain Forest and Range Dickson, J. G. 1991. Birds and mammals of pre-colonial Experiment Station General Technical Report. southern old-growth forests. Natural Areas J. 11: 26-33. Johnson, A. S., & J. L. Landers. 1982. Habitat relationships Dickson, J. G., R. N. Conner, & J. H. Willianpon. 1980. of summer resident birds in slash pine flatwoods. J. Wildl. Relative abundance of breeding birds in forest stands in Manage. 46: 416-428. the southeast. South. J. Appl. For. 4:174-179. Johnston, D. W., & E. P. Odum. 1956. Breeding bird Dickson, J. G., R N. Conner, & J. H. Williamson 1984. Bird populations in relation to plant succession on the Piedmont community changes in a young pine plantation in east of Georgia. Ecology 37: 50-62. Texas. So. J. Appl. For. 8: 47-51. Lawson, E. R., & R. N. Kitchens. 1983. Shortleaf pine. Pp. Dickson, J. G., & R. E. Noble. 1978. Vertical distribution of 157-161 in Silvicultural systems for the major forest types birds in a Louisiana bottornIand hardwood forest. Wilson of the United States (I2.M. Burns, tech. compl.). USDA Bull. 90: 19-30. For. Serv. Agric. Hdbk No. 445. Dickson, J. G., and C. A. Segelquist. 1977. Winter bird McComb, W. C., P. L. Groetsch, G. E. Jacoby, & G.A. populations in pine and pine-hardwood stands in east McPeek. 1989. Response of forest birds to an improvement Texas. Proc. Annu. Conf. S.E. Assoc. Fish Wildl. cut in Kentucky. Proc. SE Assoc. Fish and Wild. Agencies Agencies. 31: 134-137. 43: 313-325. Dickson, J. G., & C. A. Segelquist. 1979. Breeding bird Meanly, B. 1971. Natural history of the Swainson's Warbler. populations in pine and pine-hardwood forests in Texas. J. North Am. Fauna No. 69. wild Manage. 43: 549-555. Meyers, J. M., & A. S. Johnson 1978. Bird communities Emlen, J. T. 1970. Habitat selection by birds following a associated with succession and management of forest fire. Ecology 51: 343-345. loblolly-shortleaf pine forests. Pp. 50-65 in Proceedings of Evans, K.E. and RA. Kirkman. 1981. Guide to bird habitat the workshop Management of southern forests for nongame of the Ozark Plateau U.S. For. Serv. Gen. Tech Rep birds. USDA For. Serv. Gen Tech Rept. SE-14. NC-68. Noble, R. E., & R. B. Hamilton. 1976. Bird populations 'in Eyre, F. H. (Ed.) 1980. Forest cover types of the United States even-aged loblolly pine forests of southeastern Louisiana. and Canada. Society of American Foresters, Washington, Proc. Southeast Assoc. Game Fish Comm. 29: 441-450. DC. O'Meara, T. E., E. A. Rowse, W. R. Marion, & E. D. Harris. Garrison, G. A., A. J. Bjungstad, D. A. Duncan, M. E. Lewis, 1985. Numerical responses of flatwoods avifauna to & D. R. Smith. 1977. Vegetation and environmental clearcutting. Florida Scientist 48: 208-219. features of forest and range ecosystems. U.S. Dept Agric., Pulliam, H. R. 1988. Sources, sinks, and population Agric. Handbook 475. regulation Amer. Nat. 132: 652-661. Gingrich, S. F. 1971. Management of young and intermediate Remsen, J. V., Jr. 1986. Was Bachman's Warbler a bamboo stands of upland hardwoods. U.S.D.A. For. Sew. Res. specialist? Auk 103: 2 16-2 19. Paper NE-195. Repenning, R W., & R. F. Labisky. 1985. Effects of even-age Hall, S. B. 1987. 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Greenville, Mississippi. Hatnilton 1992. Bmdhg bird populations in Missouri Ozatk Yahner, R. H. 1986. Structure, seasonal dynamics and habitat fo& with and without clmtting. J. WildL Manage. 5q1): relationships of avian communities in small even-aged 23-30. forest stands. Wilson Bull. 98: 61-82. Thompson, F. R, III., S. J. Lewis, J. Ckn, and D. Ewert InPress. Yahner, R. H., and D. P. Scott. 1988. Effects of forest Status of neompical migrant bids in the midwest ide-ing fragmentation on depredation of Uicial nests. J. Wild. species of management concern in Pme&n@ of a wodcslmp Manage. 49: 508-513. on the status and management of mtropical migratory birds. Ziehmer, R. L. 1992. Effects of uneven-aged timber Rocky Mountain Forest and Range Experiment Station General management of forest bird communities. M.S. Thesis, Technical Report. Univ. of Missouri, Columbia.