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Frank Renfrow theirfirst nesting records of the speciesbe- Red-breasted Nuthatches in these lower-ele- tween 1996 and 2003, mostly in mixed vationareas tends towards white pine-hem- 61'1South O'Fallon Avenue forestsbest describedas white pine-hem- 1ock•leciduousforests, which suggests that lock-deciduousforests. In pre-twentieth- recentmaturation of this foresttype within Bellevue,Kentucky 41073 centuryliterature, there are hints of the theseareas is a keyfactor in rangeexpansion species'prior occurrence in suchareas, and in this species.Recent degradation of the (email:Frank [email protected]) it may be that,after extensive logging in the spruce-firforests of thehigher elevations, as early twentiethcentury, changes in habitat well asthe mid-twentieth-century decline of andforest maturity have begun to drawthe the American Chestnut (Castanea dentara), Abstzact speciesback to formerrange. It is alsopossi- may have spurred the species'breeding The association of Red-breasted Nuthatch ble thatthe introductionof pathogenssuch rangeexpansions into lower elevations. Out- (Sittacanadensis) with spruce-firforests in asthe Balsam Woolly Adelgid may have con- sidethe spruce-fir forests, the natural range the AppalachianMountains is well known; tributedto shiftingRed-breasted Nuthatch of EasternWhite Pine (Pinus strobus) may be duringthe mid-twentiethcentury, however, populationsout of higher-elevationspruce- thekey factor delimiting the historic as well thespecies was also found nesting and sum- fir forests and into lower-elevation areas. as the recentlyexpanded range of Red- meringat lowerelevations, primarily in as- Forestconservation and preservationstrate- breasted Nuthatch within the southeastern sociation with hemlock forests. More recent- giesmust be consideredif populationsof UnitedStates. An understandingof the his- ly,breeding populations have been found at thisspecies are to thrivein themountains of toric and current environmental factors driv- still lowerelevations in the Appalachianre- the southeastern United States. ing theserange and population changes will gion;three southeastern states documented be importantin determiningan adequate Introduction planfor habitatmanagement for thepreser- Red-breastedNuthatch (Sitta canadens•s)is a vation of Red-breastedNuthatch popula- permanentresident of thenorthern conifer- tionswithin the Appalachianregion of the ous forests of North America. with a breed- southeastern United States. ing rangethat extendssouthward in close associationwith spruce-firforests of the Breedingrange of the Red-breasted higherelevations of theAppalachian Moun- Nuthatchin the southeasternUnited tains in the southeastern United States. Dur- 5tates in historiccontext ingthe mid-twentieth centu• Red-breasted Up until the mid-twentiethcentury, the Nuthatches were also found in summer breedingrange of Red-breastedNuthatch in wxthinmiddle elevations, primarily in asso- the southeasternUnited States was thought ciationwith hemlock forests. In recentyears, to belimited to spruce-firforests within the therehave been multiple records document- higherelevations of theAppalachian Moun- ing breedingof this speciesin 1ower-eleva- tains and to extend no farther south than tmn areasof the Appalachianregion. These easternTennessee a! Roane,Great Smoky, includesmall, disjunctpockets ol occur- and Unicoi Mountains and in western North rencealong the Cumberland Plateau in Ken- Carolina at Black Mountain and Mount ucky and Tennessee,as well as moresub- Mitchell (A.O.U. 1957). A few earlierrecords •tantialpopulations along the Blue Ridge fromthe late nineteenth and early twentieth Escarpmentin Georgiaand South Carolina. centuries,however, hint that breedingout- There have alsobeen multiple records of sideof thatrange may have occurred. Most breedingrange expansion into lower- and notablewas one record for OconeeCounty, •, middle-elevation areas in the Al- SouthCarolina on 10June1910 (Sprunt and leghenyMountains of WestVir- Chamberlain1949), the samelocation where ginia and Virginia,in the Blue nestingwas verified in 2003(Renfrow 2003). Ridge Mountains of Virginia, Veryclose to thearea of recentrange expan- North Carolina. Tennessee,and sionwas a late-springrecord at LakeRabun, Georgia,and indications of possi- Rabun County, Georgia 11 May 1947 ble expansioninto the Ridgeand (Burleigh1958), althoughthis recordmay Valleysection of Tennesseeand pertainto a late-departingtransient. Ihere Georgia. seemsto havebeen no earlyevidence to in- Red-breastedNuthatch, Rabun County, Georgia, 2 June 2004. Habitatpreference for breeding dicatebreeding of thisspecies on the Cum-

ß516 NORTH AMERICAN BIRDS berlandPlateau, which is located along a line running approximately150 km west and north of the spruce-firforests of the Ap- palachians.A record of a femalecollected 16 July 1886at Bardstown,Nelson Count}; Ken- tucky (elevation 198 m, Beckham1886), some 140 km west of Kentuck;• Cumber- land Plateaupopulation, ma} pertainto an isolatedpost-irruption summering individ- ualor post-breedingtransient. In NorthCar- olina, Brewster(1886) found a presumed breedingpair in late May 1885 near High- lands,Macon County in old-growthhemlock forestat middle elevation(1219 m) Ihe firstclear indications of a breeding population in the southernAppalachians outsideof thespruce-fir forests came in the 1940sduring several intensive field studies undertakento censusbreeding bird popula- tions in lniddle-elevation areas of eastern Tennessee and western North Carolina. In the Highlandsarea of North Carolina,Holt (1974) begana seriesof studiesin 1946in Figure1. Malefled-breasted Nuthatch inOconee County, South Carolina, 14 June 2000. whichthis specieswas again found present Allphoto•rapl•s an•lr•ap grapl•c• •j,the autl•or. in summer between 1160 and 1250 m. In the Great Smoky Mountains National Park, firstSouth Carolina nest was photographed 18June 2005 alongParched Corn Creek,5 Kendeighand Fawver (1981) conducted and documentedat the ChattoogaRecre- km distant from Rock Bridge (Renfrow fieldworkduring the summersof 1947and ation Area in Oconee County. A second 1996,1999a, 2001, Renfrow,unpubl. data). 1948and found this species in a broadrange South Carolina nest was found near Burrell's Farther south along the Cumberland of elevationswithin study areas between 873 Ford in OconeeCounty on 15 April 2005 Plateau, a male Red-breasted Nuthatch was m and 1795m elevation.Stupka (1963) de- (Renfrow2003, Renfrow,in press;Renfrow, locatedon 26 May 2002 nearBandy Creek notes a record of a Red-breasted Nuthatch at unpubl.data). (400 m) at theBig South Fork NationalRiv- 884 m at CadesCove 12 June 1958 as con- There are indications that Red-breasted er andRecreation Area in ScottCounty, Ten- stitutingthe lowest-elevationbreeding sea- Nuthatchmay nest in additionalscattered nessee.Subsequent attempts to relocatethis son record for this speciesin the Great locations within the Chattahoochee Nation- speciesin the summerof 2002 wereunsuc- SmokyMountains National Park. al Forest in northernGeorgia, with two cessful.On 7 June 2003, an active Red- It wasnot untilthe latter part of thetwen- recordsfor June 1996at the CooperCreek breasted Nuthatch nest was observed and tieth century,however, that the breeding ScenicArea (734 m) in Union Countyand photographedin the samelocation as the rangeof the specieswas found to extend severalrecords at Jack• River (330 m) in the 2002 sighting. Red-breastedNuthatches intoSouth Carolina and Georgia. On 26 May CohuttaWilderness Area in FanninCounty wereagain observed exhibiting behavior in- 1986, one of two Red-breasted Nuthatches for the summer of 2000. There are also sum- dicativeof breedingin the BandyCreek sightedwas observedinspecting cavities at mer records from 1995-2000 for Rabun vicinityon 7 and 11July 2005 in two sepa- the ChattoogaRecreation Area (elevatmn Countyfrom one higherelevation location, rate locations (Renfrow and Stedman 2003, 780 m) in Sumter National Forest, Oconee RabunBald (1436 m) (¸berle and Haney Renfrow,unpubl_ data). County, South Carolina (Post and Gau- 1997. Beatonet al. 2003: G. Beaton.pers. In WestVirginia, nesting was considered threaux1989; W. Post,pers. comm.). In the comm.).The species has been found in sum- to be limitedto the highermountains and mid-1990s, summer records were obtained mer alongPersimmon Ridge Road (608 m) mountainvalleys by Hall (1983); very few in OconeeCounty as well as on the other in Greenville County, South Carolina publishedrecords of breedingbirds come sideof the ChattoogaRiver within Chatta- (Carter 1993). from elevations below 900 m. West Vir- hoocheeNational Forest in RabunCounty, Nestingof Red-breastedNuthatch on the ginia's BreedingBird Atlas project also Georgia(Oberle and Forsythe1995, Oberle CumberlandPlateau was not suspectedun- foundthis specieslimited to the higherar- and Haney 1997). In 2000, Red-breasted til this specieswas observedat the Rock eas of the Allegheny Mountains, with Nuthatches were found to be well distrib- Bridgearea (300-340 m) of the RedRiver recordsfrom Tucker, Grant, Randolph, Poc- uted alongthe ChattoogaRiver. In the 28 GorgeGeological Area, Daniel BooneNa- ahontas, Greenbrier, Webster, and Nicholas river-kilometers between Burrell• Ford (ele- tionalForest, Wolfe County, Kentucky dur- Counties (Buckdew and Hall 1994). In vation 650 m) to the north and Earrs Ford ing the summerof 1996. An inactivenest 1997, breedingRed-breasted Nuthatches (elevation490 m) to the south(Figure 1), a hole coated with pine resin was pho- were found to be well distributed at a lower- total of 14 individuals were observed at 10 tographedat thislocation on 2 July,and an elevationsite in PocahontasCounty, with separatelocations 12-15 June 2000. In May adult male was observedfeeding recently nestingconfirmed at 850 m on 24 June 2002, a firstGeorgia nest was photographed fledgedyoung on 4 July.Evidence of a very (Renfrow,unpubl. data). and documented at the Nicholson Tract of smallpopulation of threeto six pairswas In Virginia,Kain 0987) describedRed- the ChattahoocheeNational Forest; two ad- gatheredhere duringsubsequent breeding breastedNuthatch as a locallycommon sum- ditionalRabun County nests were located in seasonsencompassing 1997-2005. An ac- mer residentat Mr. Rogersand a locallyun- the vicinityduring the 2002 breedingsea- tive nestwas photographed and document- common to rare summer resident elsewhere son. Another nest was located at the Nichol- ed at Rock Bridgein May 1999. Another in the higher elevations During the sonTract on 19April 2005. In April 2003, a Wolfe Countynest record was obtained on 1984-1989Virginia Breeding Bird Atlas proj-

VOLUME 59 (2005) NUMBER 3 517 oct,breeding evidence was report- '• ,. stituung the only breeding ed in Grayson,Highland, Wise, recordfor North Carolina'spied- Rockbridge,Botetourt, and Au- , mont (Burroughs 1976), oc- gustaCounties, with breeding curredsubsequent to minorwin- confirmedat Whitetopand Old •' terirruptions inassociation with RagMountains (R. Clapp,pers. • bird feeders in wooded residen- comm.). In 2002, two lower-ele- t, tial areaswith pines. vation nests were located in Hatrapand Quinn (1995) de- MontgomeryCounty, the first at scribe some variabihty of the 634m on 27April and the second southernboundary of thebreed- at 616 m on 2 June 2002 (R. ing rangeof Red-breastedNut- Mays, unpubh data) In May hatchin the Appalachians,w•th 2004, two additionalnests were greaternumbers breeding on the locatedin this samevicinity (R. southernperiphery in yearsfol- Mays, pers. comm.). A Red- lowinggood cone crops. This is breastedNuthatch at Big Black alsoindicated by Mays(unpubl. Mountain (1265 m) in Harlan data)in referenceto thebreeding County,Kentucky on 3July 1982 population in Montgomery (Palmer-Bafl2003) wasmost like- Figure2.A white pine-hemlock-deciduous forestinWolfe County,Virginia, which does not ly a wanderingindividual from County,Kentucky. Note the dominance ofEastern White Pine. seemto bepresent on anannual the nearbyWise County,Virginia basis.Assumptions of sporadic breedingpopulation, as there is virtuallyno post-irruptivebreeding records may be lo- occurrencemay on occasionbe incorrect,as coniferouselement on Kemucky'sBig Black cated within urban and suburban areas breedingRed-breasted Nuthatches can easily Mountain. amongplantings ofornamental conifers. Ex- be overlooked.Peterjohn (1989) described Thisseries of newbreeding records in the amplesinclude a nestingat a cemeteryin summeringRed-breasted Nuthatches at southeasternUnited States roughly coincided HamiltonCounty, Ohio in 1982and a nest- HockingCounty, Ohio as remaining for only with an earlier established trend towards ingattempt at a CampbellCounty, Kentucky oneor twoyears and then disappearing. In- southwardrange expansion of thespecies in cemeteryin 2001(Renfrow 1982, 2001). A tensivescrutiny of theplanted pine areas in otherstates just to thenorth. However, in HockingCounty has since revealed con- contrast to the recordsfrom southern ar- sistentnesting activity 1998-2005 (Ren- eas-documentedin naturallyregenerat- frow 1998, 1999b, 2002, Renfrow,un- ing mixedforests--the northern states' publ.data). These birds are often present recordshave been documented primarily atone isolated pine stand in oneyear and in maturingconifer plantings. In Pennsyl- at anotherstand the next year, so all suit- vania,maturation of coniferplantations ablehabitat must be thoroughlycovered establishedduring the 1930sprovided to assessannual occurrence accurately. habitat for nesting Red-breasted [naccessibihtyof their habitats and secre- Nuthatchesby theearly 1960s. Expansion tivebehavior during the breeding season in Pennsylvaniacontinued well into the mayaccount for these populations being 1990s,with firstcounty nest records for overlooked until the latter half of the PhiladelphiaCounty in 1995and Alleghe- twentiethcentury, as was the case with ny Countyin 1996 (McWilliamsand Swainsoh'sWarbler (Limnothlypis swain- Brauning 2000; R. Leberman,pets. sonii),which also breeds in someof the comm.). In southeasternOhio, summer- sameareas (Renfrow and Stedman2003). ing Red-breastedNuthatches were ob- [n North Carolina.Holt (1974) described servedintermittently during the 1980s thedifficulties for fieldstudies posed by (Peterjohn1989), and nestingwas con- impenetrabletangles of rhododendronin firmedin summer1998. Furthernesting thevirgin hemlocks, which she traversed evidencewas gathered during the subse- with greateffort on handsand knees. quentbreeding seasons of 1999-2005, but AlongOhio's Allegheny Plateau, Hicks onlya totalof oneto threepairs could be (1935) emphasizedthe inconspicuous- locatedin anygiven year. Although a few ness and choice of secluded habitats as nativepine species do occurin Hocking prinmryfactors contributing to the ex- Countyin mixedassociation and although tremedifficulty of locatingRed-breasted hemlock-deciduous forest is found in sev- Nuthatchesduring the breeding season. eral deep gorges,the Red-breasted Nuthatcheshere were only found in areas Figure]. An addgid-kifiM 01d-gr0wth Eastern Heml0

;18 NORTH AMERICAN BIRD clearingfor subsistenceagriculture and the (Georgeand Bazzaz 2003). A similar cycle of renewal has taken subsequentregeneration of mixed lorest Manyplots of landcleared for agricultural placeon the CumberlandPlateau. Thomas Ihe introductionof pathogensresulted in a useby nineteenthcentury settlers in theAp- (1926) describeda vast forest of the finest host of catastrophictree diseasesthat ad- palachianswere abandoned during the agri- whitepine at WolfeCounty, Kentucky just verselyaffected the higher-elevationsforests culturaldepressions of the 1890sas well as prior to the culminationof extensivelog- in particular.Changes in thebreeding range duringthe GreatDepression of the 1930s. gingof thearea. Ihese forestshave now re- of Red-breastedNuthatch may have resulted Theseabandoned fields were quicklysuc- turnedto Wolfe County,with the greatest fromthe conflictinginfluences of expansion ceededby standsof naturallyregenerating densitieso[ EasternWhite Pineoccurring and recoveryof EasternWhite Pinein the EasternWhite Pine, renewing the ecological in the RockBridge area (Figure 2). Ihe his- lower-elevation forests coincident with the cycle(Walker 1990). Ihis is particularlyevi- toric White Pine Church (less than one km lossof spruce-firhabitat occurring in the dentin theareas where breeding Red-breast- fromthe Scott County, Iennessee nest loca- higherelevations. tion) may predatethe ex- Landuse practices tn the •' , tensivelogging operations EasternUnited States dur- • o[ the early twentiethcen- ing the nineteenthcentury tury(Manning 1993) and eliminated Eastern Hemlock ,, (Tsugacanadensis) and East- ingnow second-growthstands withinamatur-forest ern White Pine from many - againdominated byEast- areas of the deciduous for- ernWhite Pine (Renfrow est, particularlythe north- andStedman 2003). ern hardwood and central broad-leavedregions. Ihe useof hemlockbark by the sion of the white tanningindustry, along with loggingand fires,severely forests in the lower eleva- restricted the distribution of pine-hemlock-deciduouscoverytionDuring ofand thethispossible Appalachians,periodexpan-ofre- bothconifer species (Yahnet therewas aprecipitous de- 2000). In the southernAp- cline in the health of the palachians,the remoteness spruce-fir forests in the of the areadelayed the de- higherelevations. The Bal- structionof theforests. Log- Figure4. EasternWhite Pine resin applied to Red-breastedNuthatch sam Woolly Adelgid gingoperations for Eastern nestentrance Oconee County, South Carolina, 2 June 2003. (Adelgespiceae), inadver- White Pine started in 1875 tently broughtinto North on the North Carolina section of what was to ed Nuthatchesreach their highest densities Americaat the beginningof the twentieth becomethe Great Smoky Mountains Nation- in northeasternGeorgia, at the Nicholson century,was found to havereached the en- al Park,where this pine species comprised 20 tract of the Chattahoochee National Forest as demicFraser Fir (Abiespaseri) population per cent of the bottomlandforests (Stupka well as in South Carolina's Sumlet National of the southernAppalachians during the 1964). Fire was also a factor: for instance, Forestat the RussellFarmstead, located just mid-1950s(Carlin 1984). Duringthe latter EasternWhite Pinewas negatively affected acrossthe ChattoogaRiver from the old half of the century,a majorityof mature bywildfires that consumed over 50,000 acres Nicholsonfarmstead (Renfrow 2002). FraserFirs succumbedto this adelgid.At within CaldwellCounty, North Carolinain Beginningin 1911, much of the land in Mount Collins in the Smokies, Fraser Fir 1894 (Ashe 1895). the ChattoogaRiver watershed was pur- hasbeen virtually eliminated and canopy Ihe first of a seriesof catastrophictree chasedby the U.S.Forest Service, resulting coverreduced to half of its previouslevel diseasesbegan in 1904 with the accidental in a periodof recoveryand renewalfor the (Rabenold et al. 1998). The death of the firs introduction into North America of Endoth- region'sforests (Boyd 2001). Despitethe in- causedweakening in the anchoringof the ia parasitica,the fungusresponsible for tervening period of settlementactivity, intertwinedroots of adjacentRed Spruces chestnutblight. In the southernAppalachi- there is a remarkablesimilarity between (Picearubens) (Gove 1996). Subsequent ans,the American Chestnut once comprised early accountsof the forestand the condi- vulnerabilityto windstormsalong with oth- an estimated25 per centof the mixedde- tion of the forestalong the BlueRidge Es- er possiblefactors, including drought, ice ciduousforest. By the 1940s,the chestnuts carpmenttoday. In a 1776 descriptionof storms, and acid rain have resulted in the werealmost entirely extirpated, leaving only the habitatof the southernBlue Ridge Es- lossof nearly50 percent of theRed Spruce the stubbornlyregenerating sprouts still carpment,William Bartramdescribed the treesat highelevations from New England presenttoday. Large openings created by the approachto "OcconneMountain" as "a to North Carolina in the 1980s and 1990s lossof the chestnutsresulted in the expos- magnificentlandscape infinitely varied" (Ayerset ah 1998). ing of mineralsoils particularly favorable to with a speciesof rhododendron"foremost Furthercomplicating this alreadycom- the regenerationof EasternWhite Pine in the assemblyof mountainbeauties"; after plex seriesof catastrophicevents affecting (Walker 1990, Yahnet 2000). Loss of the relatinga seriesof steeprocky ascents, Bar- theAppalachian forests, the Hemlock Wool- American Chestnut has also been linked to tram lists Eastern White Pine first in a ros- ly Adelgid(Adelges tsugae) was imported an increase in RosebayRhododendron ter of treespecies present (Doren 1955). In into North America in the mid-twentieth (Rhododendronmaximum) and Mountain thissame area, over two hundred years lat- century. During the 1990s, widespread Laurel(Kalmia latiJolia), now estimatedto er, Gaddy(2000) describesEastern White deathof hemlocksbegan to occurin the coveran estimated2.5 millionha in theAp- Pineas one of the mostcommonly encoun- ShenandoahNational Park (Figure 3), and palachians.In turn,the dense cover of Rose- teredevergreens on slopesthat are often evidenceof this adelgidhas since been bay Rhododendronhas beenlinked to in- dominatedby a shrublayer of heathspecies foundin thesouthern Appalachians, casting creasedregeneration of EasternHemlock includingthree species of rhododendron. doubton the long-termsurvival of Eastern

VOLUME 59 (2005) NUMBER 3 519 Hemlockin manyareas (Ayers et al. 1998, of 11pairs per 40 ha,as well as in pineheath tions of Appalachian forests as being LeGrand2004). Hemlockmortality in the forests between 1219 m and 1349 m at con- sharplysegregated by slopeposition and northeastern United States has been linked centrationsof 7.3 pairsper 40 ha. orientation,these forests are broadlyinte- to declinesin populationsof Black-throated At elevationsbelow 860 m, breeding grated,with EasternWhite Pine often dom- Green Warbler (Dendroicavirens), Acadian populationsof Red-breastedNuthatch are inatinginto the upperslopes, and w•th a Flycatcher (Empidonaxvirescens), and foundin mixedforest invariably containing considerableintermingling of Pitch,Short- BlackburnianWarbler (Dendroicafnsca) and a strongelement of EasternWhite Pine, leaf,and Virginia Pines into lower-slopear- anincrease in somespecies birds not associ- Eastern Hemlock, Red Maple (Acer eas.This forest type seems not to havebeen ated with conifersin the East, including rubrum), Sweet Birch (Betnla lenta), and givena specificdesignatton in earlierlitera- White-breasted Nuthatch (Sitta carolinensis) variousoaks (Qnercns spp.), as well asa sig- ture and might be best categorizedas a (Tingleyet al. 2002).In thesame study, data nificantcomponent of "hard"pine species whitepine-hemlock-deciduous forest of a on Red-breastedNuthatch did not showsig- suchas Pitch (P rigida),Virginia (P virgini- distinctlysouthern character. In the past, nificanttrends (Tingley et al. 2002).There ana), and Shortleaf (P echinata). Understo- the characterization of forests as "hem- are indicationsthat breedingRed-breasted ry vegetationin thesemixed forests is quite lock-deciduous"may havecaused the ele- Nuthatchesin thelower-elevation areas may dense,consisting of a denseheath layer of ment of Eastern White Pine within this for- be successfullyadapting to theloss of hem- oneor morespecies of Rhododendron,Rose- est type to be overlooked.For example, lock in areas where other conifers are Kendeighand Fawver(1981) failedto present.[n 2002, breedingwas con- mention Eastern White Pine in the firreedin MontgomeryCounty, Virginia contextof theirGreat Smoky Mountain at 634 m in an area with Eastern White NationalPark studyareas, including Pine listed first in order of abundance; those in hemlock-deciduous forest, in this vicinity,only a few declining, whereas Holt (1974) listed Eastern adelgid-in[estedhemlocks were present White Pine as second in abundance to (Mays, unpubl. data). Red-breasted Eastern Hemlock in hemlock-hard- Nuthatches were also found in associa- woodforests within her studyareas at tion with Eastern White Pine in areas Highlands,North Carolina. with standingdead hemlocks in eastern In the lower- to middle-elevation AugustaCounty, Virginia in April 2004 forestsoften described as pure hemlock (Renfrow,unpubl. data). or hemlock-deciduous,some element of an additionalconifer species is most Habitat selection in likely required for Red-breasted 11ppalachianpopulations Nuthatchbreedtng, as the thtnwscostty oflled-breastedNuthatch of EasternHemlock resin is apparently Habitattypes favored by Red-breasted figure5. lhena•al •tn•e0f [isle•n•hite Pine,showin• 10r•i0ns 0f not suitablefor nestentrance prepara- Nuthatchesin the higherelevations of re{entl•dis{0•ered Red-breasted •ulhtl{h nest re{0rds in the (umberland tion:this species regularly coats the out- theAppalachians include spruce-fir for- Pl•eauan• BlueRidge [srarpmenl. stde entrance of the nest hole w•th the est, maturespruce forest, and mixed moreviscous resin of spruce,fir, or pine spruce-hardwoodforest (Simpson1976, bay Rhododendronin particular, with (Harrapand Quinn 1995),and access to at Hall 1983). In the Smokies,Kendeigh and MountainLaurel present as well. Theseba- leasta few such resin-producingconifers Fawver (1081) found this species tn sichabttat elements are all presenttn lower- nmybe a minimalrequirement to providead- spruce-firdilnax forestsbetween 1760 m elevation Red-breasted Nuthatch nest site equatehabitat for nesting.At elevationsbe- and 1790m at concentrationsof 20 pairsper locationsin Kentucky,Tennessee, West Vir- low thoserequired for fir or spruce,Eastern 40 ha, but this specieswas not detectedin ginia,Virginia, North Carolina,South Car- White Pine is most often found in close as- seralspruce-fir forests above 1705 m. This olina,and Georgia. Extensive breeding-sea- sociation with Eastern Hemlock. The resin of speciescan occasionally be found nesting in son searches in lower-elevation areas thispine species has been the most frequent high-elevationnorthern hardwood forests, adjacentto nest areas(specifically, areas sourcefor nestpreparation in the Kentucky, with thepresence of evena smallamount of that lack one or more of these habitat ele- SouthCarolina, and Georgianests of Red- sprucebeing sufficient to producea nesting ments) have been conducted from 1996 breastedNuthatch observed by this author population(Hall 1083).In North Carolina, through2005; these searches have failed to (Figure4)_ Pitch, Virginia, and Shortleaf Pine Red-breasted Nuthatches also nest where find any summering Red-breasted also have viscous resin suitable for the nest. RedSpruce ranges well below that of Fraser Nuthatches(Renfrow, unpubl. data). Shortleaf Pine is listed as one of three south- Fir, in a transitionalzone of spruce-hem- These lower-elevation Appalachian ern pine specieswith resinchemistry suit- lock-deciduous forest between 1372 m and forests with breeding Red-breasted ablefor useby Red-cockadedWoodpeckers 1676m in the southernBlue Ridge (Carlin Nuthatches combine elements from several (Picoidcsborealis) for producingresin-barri- 1984, Renfrow 2003). foresttypes, including mixed mesophytic, ers that prowdenest defenseagainst rat Belowthe rangeof Redbpruce, summer- hemlock-whitepine-northern hardwoods snakes(Elaphc spp.) (Conneret al. 2003). ing Red-breastedNuthatches have been and southernoak-hickory-pine. Braun Ghalamborand Martin (1999) suggestthat foundin old-growthhemlock, hemlock-de- (1950) describeda forestof this type at Red-breasted Nuthatches construct similar ciduous,hemlock-white pine, and white Kemucky• Red River Gorgeas a mtxed resinbarners for reductionof bothcompeti- pine forestsat elevationsbetween 1097 m mesophyticcommunity like the hemlock- tiveand predatory interactions at thenest. and 1341 m in the southernBlue Ridge whitepine-northern hardwoods forest but In lower- to middle-elevation situations, (Holt 1974,Simpson 1976). In theSmokies, with an understo• stratumof rhododen- treesutilized for nestingby Red-breasted Kendeighand Fawver(1981) found this dron,magnolia (Magnolia spp.), and holly Nuthatchfall into twogeneral categories. In speciesin hemlock-deciduousforests be- (Ilex spp.) that lends the habitat a more somecases, nesting birds use a standing tween 873 m and 1356 m at concentrations southernaspect. Contra• to somedescrip- deadconifer trunk, usually with thetop por-

520 NORTH AIVIERICAIq BIRD Figure6. Resultsofprescribed burnat Wolfe County, Kentucky, 12May 2004. Note brown-needled Eastern White Pine saplings. tion and most of the limbs broken off but smaller lichen-encrusted branches of oaks Red-breastedNuthatch range with somebark still intact(particularly at possiblybeing more frequently probed and expansioninrelation to the nestcavity entrance). Nest trees in such inspectedthan thoseof the surrounding EasternWhite Pine distribution cases have included Pitch Pine. Eastern conifersin thesemixed-forest environments, The discoveryof breedingRed-breasted White Pine, Shortleaf Pine, and Eastern especiallyduring the nestingseason (Ren- Nuthatches on the Cumberland Plateau calls Hemlock. Other nests have been found in a frow,unpubl. data). into questionits generaldesignation as a deadportion of a livingdeciduous tree, often During searchesconducted in lower-ele- "northern-affinity"species. These disjunct a RedMaple or SweetBirch. These nests are vationareas of theAppalachians, areas pos- pocketsof breedingoccurrence are consider- usuallyexcavated fresh in a new location sessingall of theabove-mentioned forest el- ablyremoved from the breedingranges of eachyear. ements-but lacking a dense heath otherpassefine species considered to pos- The seeds extracted from cones of the understorystratum--have nol been found sessnorthern affinities. Haney (1999) lists EasternWhite Pineare an importantsource to containRed-breasted Nuthatches, for rea- Red-breastedNuthatch with Brown Creeper of food for this nuthatch, as are insects sons that are not clear. The same is not true (Certhia americana), Winter Wren gleanedfrom lichen-encrusted twigs and in planted pine areasfarther north. Al- (Troglolytestroglodytes), and Golden- branches.In winter,this species can often be thoughthis species forages readily on the crownedKinglet (Regulus satrapa) as species observedforaging in the deepergrooves, groundin clearingsassociated with trails, forminga recurringgroup that exploits splits,and irregularities in thebark of these trampledopenings, and even grass)- areas, it coniferouscover and complex structure typ- pines,especially at the circularwhorls of can occasionallybe found foraginglow icalof oldernatural forests across broad ge- branchjunctures with the maintrunk. The amongthe rhododendronfoliage. Close as- ographicaldomains. These other specres do loose,flaky barks of Pitch,Virginia, and sociationof RosebayRhododendron with extendtheir breeding ranges from higher el- ShortleafPines also serve as favored foraging Eastern White Pine and Eastern Hemlock evationsinto the Blue RidgeEscarpment substrates.Both Eastern Hemlock and East- within their rangesin the southeastern Gorges of South Carolina but have no ernWhite Pinetend to producehea W cone UnitedStates may account for thisapparent knownbreeding populations on the Cum- cropsin whichthe cones quickly open upon inconsistency,as might the apparentclose berland Plateau (Stedman and Stedman ripening;the latterspecies produces cones associationof the lower-elevationAp- 2002, Palmer-Ball 2003, Renfrow 2003). biennially,thus limiting the availabilityof palachian populations of Red-breasted Blue-headedVireo (Vireo solitarius) and thesefood sources during many seasons. On Nuthatch with those of Swainsonk Warbler Black-throatedGreen Warbler, two Neotrop- the otherhand, the conesof Pitch,Virginia, (Renfrowand Stedman2003). Old-growth ical migrantspecies that sharethis same andShotdeaf Pine are persistent and slow to hemlock forests in both middle- and lower- habitatand are sometimesthought of as open(Holmes 1959), providing a morede- devatronareas are notablefor their prodi- northern-affinityspecies, both have breed- pendablefood source from year to year.Ad- gious understory stratum of Rosebay ing rangesthat extendfar to the south,well ditionally,the deciduouselement of the for- Rhododendron,as noted by Brewster intothe Piedmont region of Georgia(Beaton est providesforaging substrates, with the (1886), Holt (1974), and Renfrow (2003). et al. 2003).

OLUME 59 (2005) NUMBER 3 Outsideof therange of thespruce-fir for- on the increase;for instance,numbers sur- cana), Black-throated Green Warbler, est,the naturalrange of EasternWhite Pine veyedalong Willis Knob Roadin Rabun Swainsong Warbler, Black-and-white War- (Figure5) maywell be the key component County,Georgia were found to be compara- bler(Mniotilta varia), Worm-eating Warbler delimitingthe historicas well asthe recent- ble to thosealong Clingman• Dome Road at (Hcbnithcrosvermivorus), Ovenbird (Sciun•s ly expandedrange of Red-breastedNuthatch Great SmokyMountains National Park in aurocapillus),Louisiana Waterthrush (Seiu- within the southeastern United States. This June2004 (Renfrow, unpubl. data). •us •notacilla), and Hooded Warbler (Wilso- naturalrange extends along the Appalachian A considerableportion of the breeding nia cit•na): the avifaunal richness of this Mountains southwardthrough the Blue habitat of Red-breasted Nuthatch in the lower-elevationAppalachian habitat is clear. RidgeMountains of South Carolinaand southeastern United States occurs on lands Concernsregarding loss of biodiversity Georgia,with one major lobe extending designatedas NationalForest, and thesear- withinthe herbaceous layer and changes in westwardthrough the Ridgeand Valley into eascontain an estimated 25 percent of hem- forestsuccession due to the predominance Tennesseeg Cumberland Plateau and a dis- lock-whitepine forests in the SouthernAp- of RosebayRhododendron and Mountain continuoussection encompassing the north- palachians(Hunter et al. 1999).Management Laurelin the understorystratum (George ern portionof the CumberlandPlateau in for increasedprotection of the mixedwhite and Bazzaz 2003) need to be balanced with Kentucky(kirtle 1970). pine-hemlock•leciduousforest type would concernsfor protectionof existingbird bestrongly indicated for the protection of the habitat in these areas. Tentative conclusions: Red-breastedNuthatch. At present,the Hem- Red-breastedNuthatch range lockWoolly Adelgid presents a serious threat Acknowledgments expansionand habitat change to theseforests (LeGrand 2004), and the im- I wouldlike to especiallyacknowledge and Known Red-breastedNuthatch breeding plementationof adelgidcontrol measures is thankRyan Mays for providingdetailed in- populationswithin lower-elevationareas of of criticalimportance. The United States For- formation on Red-breasted Nuthatch breed- the southeasternUnited States are relatively estService has initiated one adelgid control inghabitat and behavior in westernVirginia_ fewat present,but further expansion into ar- measure,namely the releaseof the coccinel- The followingpersons also provided infor- eas of maturingmixed white pine-hemq lid beetlePscudoscy•nmls tsugae in selected mationreferenced in thispaper: Gift Beaton, [ock•leciduousforest might be expected locations within the Chattahoochec National Roger Clapp, Steve Cole, Andy Gaston, with the continued degradation of Forestin 2004.This beetle is a highlyspe- GeorgeHall, KathrynHuie, RobertLeber- spruce-firforests in higher-elevationareas. cializedprcdator, feeding almost exclusively man,Harry LeGrand,Brainard Palmer-Ball, Kendeighand Fawver(1981) cite overflow on theHemlock Woolly Adelgid and possibly Jr., Will Post,Stephen Stedman, and Bill fromnearbv spruce-fir forests as a possible othersimilar adelgid species (Jensen 2004; S. Whan.Finally, I offera specialthanks to my explanationfor breedingpresence of Red- Cole, pers.comm.). Further expansion of wife,Veronica Rcnfrow, for pointingout the breastedNuthatch, Brown Creeper, Winter useof thiscontrol measure could help to pre- obvious birds that I ohen overlook such as Wren, and Golden-crownedKinglet in the servethe remaininghemlock constituent of the first June record of a Red-breasted hemlockcommunities of the GreatSmoky theAppalachian torests. Nuthatchin Kentucky Mountains National Park. Other concernswith regard to Red- Askins (2000) lists Red-breastedNuthatch breasted Nuthatch habitat include the ef- Literature cited amongavian speciesprimarily associated fectsof fireand of logging.Recent planning Ashe,W. W. 1895. Forest fires:their destruc- with undisturbedspruce and fir forests.At strategiesutilizing prescribed burns in Na- tive work, causesand prevention.North present,it is not clearto whatextent the se- tional Forestareas, such as a prescribed CarolinaGeological Su•xey Bulletin 7. vere diseasein southernspruce-fir forests burn in 2004 (Figure6) at the RockBridge Askins,R. A. 2000.Restoring North Amo•cak will affectpopulations of this species,but area in Kentucky (B. Palmer-Ball, pers. bi•rls:lessons [•om lamlscapeecology. Yale populationdeclines have already been de- comm.;K. Huff, pers.comm.) and several University Press,New Haven, Connecti- tectedin someareas. During the late 1990s. recentprescribed burns within the Chatta- cut. Red-breastedNuthatches were reported to be hoocheeNational Forest in RabunCounty, Ayers,H., J. Hager,and C.E. Little.1998. An scarcem theirusual stronghold in thespruce Georgia(A. Gaston,pers. comm.), could re- AppalathkmTragedy. Sierra Club Books, belt of the Cheat Mountains (Hall 1997). duceRed-breasted Nuthatch populations lo- San Francisco,Califonua. Rabcnold et al. (19c•8) list Red-breasted cally Thisspecies appears to havea strong Beaton.G.. P W. Sykes,and J. w. Parrish. Nuthatch with Black-cappedChickadee affinityfor structural features of old-growth 2003Annotated Checklist of GeorgiaBirds. (Poecileatricapillus), Brown Creeper, Winter forestsand mature, undisturbed forests, is Fifth edition. Occasional Publication No. Wren, and Golden-crownedKinglet as adverselyaffected by forestfragmentation, 14, GeorgiaOrnithological Society. specieshaving declined mcasurab b where and appearsto shun heavilyburned areas Boyd,B. A. 2001. TheChattooga' wild and adelgidinfestations have causeddramatic (Ghalamborand Martin IC)9C),Haney 1999, scenicriver. Fern Creek Press,Clayton, changesin the spruce-firforest structure Askins 2000). Georgia. within Great SmokyMountains National Commercialpressure to rcsulneor ex- Beckhain,C. W 1886 The Red-breastedNut- Park. Brewster (1886) listed Red-breasted pandlogging in whitepine-hemlock-decid- hatch in Kentucky in summer. Auk 3: 489. Nuthatch with Winter Wren and Golden- uous forests likewise continues to be a con- Braun,E. L. 1050.Deciduous Forests of the cro•ncd Kinglet as the most abundant cern.The useof prescribedburns, selective Easto• United States. Hafner Press, New speciesin thespruce forests of • estcrnNorth loggingof conifers,or clearcuttingcould be York. Carolina: the call of the nuthatch is described detrimentalto Red-breastedNuthatch pop- Brewster,W 1886.An ornithologicalrecon- as almostomnipresent in theseareas. As in ulations,as could any management practice naissance in western North Carolina. Auk Brewster's time, Red-breasted Nuthatches that inhibitsgrowth of EasternWhite Pine 3: 04-112, 173-170. can still sometimes be heard three or four at anti RosebayRhododendron. Other passer- Buckdew,Jr., A. R., and G. A. Hall. 1094 a time in higher-elevationareas, but such ine speciesthat breedin significantnum- TheWest Virginia Breeding Bird Atlas. Uni- abundancenow seemsto be the exception bers in these mixed forests include Blue- versityof PittsburghPress, Pittsburgh, ratherthan the rule. Conversely, populations headed Vireo, Red-eyed Vireo (Vireo Pennsylvania. in somelower-elevation areas appear to be olivaccus),Northern Parula (Panda m•1oi- Burleigh,T. D. 1958.Georgia Birds. Univer-

NORTH AMERICAN BIRDS sity of OklahomaPress, Norman, Okla- tion Plan for the SouthernBlue Ridge --. 1998. Another look at the birds of homa. (PhysiographicArea 23). Version 1.0. HockingHills. Ohio Cardinal 22: 44-46. Burroughs,E. B_1976. Apparent nesting of AmericanBird Conservancy,The Plains, --. 1999a. An active nest of the Red- Red-breastedNuthatches in Rockingham Virginia. breastedNuthatch at theRed River Gorge_ County,N.C. Chat40: 45. Jensen, D. W. 2004. Decisionmemo: Hemlock KentuckyWarbler 75:53-55 Carter,R. M. 1993. FindingBirds in South WoollyAdelgid control, fiscal years 2004- ß 1999b. Red-breasted Nuthatches nest- Carolina.University of South Carolina 2006. USDA Forest Service, Chatta- ing at HockingHills: the first recordsfor Press,Columbia, South Carolina. hoochee-OconeeNational Forests,Tallu- southeastern Ohio. Ohio Cardinal 22: 131- Carlin,D. T. 1984.A NaturalistkBlue Ridge lab Ranger District, Rabun County, 133. Parkway.University of KnoxvillePress, Georgia. --. 2001. BreedingRed-breasted Nut- Knoxville, Tennessee. Kendeigh,S.C., and B J. Fawver. 1981. hatchesin Kentucky:a nestingattempt in Conner, R. N., R. H. Johnson, D.C. Breedingbird populationsin the Great CampbellCounty and a Wolfe County Rudolph,and D. Saenz.2003. Do Red- SmokyMountains, Tennessee and North update.Koitucky Warbler 77: 64-68. cockadedWoodpeckers select cavity trees Carolina. Wilson Bulletin 93: 218-242. ---. 2002. HockingHills 2001 - A pine basedon the chemicalcomposition of keGrand,H. E. 2004. Stateof the Region: woodsodyssey: Red-breasted Nuthatch pineresin? Wilson Bulletin 115: 397-402. Southern Atlantic. North American Birds fiedgingdetails and other recordsßOhio Doren,M. V. 1955. Frayelsof WilliamBar- 58: 513-514. Cardinal 25: 186-190. tram.Dover Publications, Inc., New York. kirtle, E. k. 1970.Endemic, disjunct, and -----. 2003. Red-breasted Nuthatch and Gaddy,L. L. 2000.A NaturalistkGuide to the northerntrees in thesouthern Appalachi- Golden-crownedKinglet: the first nest SouthernBlue Ridge Front. University of ans.In: The DistributionalHisto.ry of the records for South Carolina and other South Carolina Press, Columbia, South Biotaof theSouthern Appalachians, Part II: Chattoogarecords. Chat 67: 102-109. Carolina. Flora. ResearchDivision Monograph 2. --. in press.Georgia's first nesting records George,L. O., and E A. Bazzaz.2003. The Virginia PolytechnicInstitute and State for the Red-breastedNuthatch. Oriole 67. herbaceouslayer as a filter determining University,Blacksburg, Virginia. Renfrow,E, and S. J. Stedman.2003. Red- spatialpattern in foresttree regeneration. Manning,R. 1993. TireHistoric Cumberland breastedNuthatches in the Big South In: TireHerbaceous Layer in theForests of Plateau:an explorerkguide. University of Fork River and Recreation Area: first nest- Eastern North America (E S. Gilliam and TennesseePress, Knoxville, Tennessee. ing recordfor Tennessee'sCumberland M. R. Roberts,eds.) OxfordUniversity McWilliams,G. M., and D. W.. Brauning Plateau.Migrant 74: 1-5. Press,New York. 2000. FireBirds of Poresylvania.Cornell Simpson,M. B. 1976.Breeding season habi- Ghalambor,C. K., and T. E. Martin. 1999. UniversityPress, Ithaca, New York. tat and distribution of the Red-breasted Red-breastedNuthatch. The Birds of North Oberle,M. W., and D. M. Forsythe.1995. Nuthatchin the southernBlue Ridge America, No. 459 (A. Poole and E. Gill, Possible breeding by Red-breasted province.Chat 40: 23-25. eds.). The Birds of North America, Inc., Nuthatchand Golden-crowned Kinglet in Sprunt,A., and E. B. Chamberlain.1949. Philadelphia,Pennsylvania. Georgiaand South Carolina.Oriole 60: SouthCarolina Bird Life. Universityof Gove,D. 1996.Spruce-firSelf-guiding Nature 52-55. South Carolina Press, Columbia, South Trail. Great Smoky MountainsNatural Oberle,M. W..,andJ. C. Haney.1997. 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