P0c0sinbreeding bird fauna

Widely recognized for their unique botanical nature, little study has been done on pocosin avifauna.

David S. Lee systemsand/or adjacent to estuarine from an Algonquin Indian term systems.Such mixed areas often pro- HE"poquosin"WORD POCOSIN and is oneORIGINATED of the few vide a rich mosaic of wetland habitats Algonquin words used by European involvingbroad zonesof transitionand settlers. Pocosin habitats are defined complex successionalpatterns. Exten- vath difficulty since considerablecon- siveareas called pocosins by naturalists fusion persistsin the use of the term. and environmentalistsare, in fact, often Nevertheless,it is obviouslycritical to composedof swamp forest, hardwood definethese habits accuratelyif mean- or evergreenshrub bog have been used forest, and marshes.Although there lngful habitat comparisonsare to be to describepocosin vegetation types. seemsto be no comprehensivebotanical made. Tooker (1899) provided detailed The term "bay" is particularly con- definition of pocosinmost researchers discussionon the origin, meaning,and fusing becauseit refers to a number agreethat pocosinsare characterizedby usageof the word, and somerecent au- of moderately advanced successional Pond Pine and denseevergreen shrub thors have credited him as translating stages of Southeastern wetlands that vegetationgrowing on deep peat or pocosin as "swamp on a hill." While support several speciesof bay trees sandypeat soilswith protractedhydro- th•s is an interestinginterpretation of (Sweetbay,Redbay, and Loblolly-bay), periods. the word, it was never defined as such whereasthe term Carolina bay, partly Although pocosins are currently by Tooker.In tracingout the earlyusage named for the presenceof bay trees,re- communities of major interest to en- of the term, both.by Native Americans fers to elliptical depressionsthat may vironmentalistsand are widely recog- and early settlers,Tooker found that support pocosin vegetation. Carolina nizedfor theirunique botanical nature, "pocosin"referred to a wide variety of bays, which are permanent geological few havebeen intensivelystudied flor- low wet areas extending from New features,are often specificallynamed istically.Wells (1946) provideda gen- England through the Carolinas. It was sites(i.e., Wolf Bay in BladenCounty), eralbotanical analysis of Holly Shelter's locally usedinterchangeably with "dis- but bay forestsare successionalstages pocosins(Pender County, North Car- mals" and "galls"by Europeansettlers of wetlands.Likewise, there are many olina)and Kologiski (1977) investigated to describeswampy thickets. areason mapsnamed pocosin (i.e. Light the vegetative communities of the Geologists, hydrologists, botanists, Ground Pocosinin Pamlico County), Green Swamp (Brunswick County, ecologists,and other investigatorshave but visitsto thesesites reveal that many North Carolina),including several types hkewise used the word to describe a va- outgrew,long ago, the pocosinvegeta- ofpocosin, savannaand related succes- riety of low, woodedwetland habitats, tion stage.Definition is further ham- sional communities. Buell and Cain and in many instancesthe terms bay, pered by the fact that many pocosins (1943) describedthe successionalrole bayhead,stream head forest, shrub bog, are situatedwithin extensivepalustrine and ecologicalrequirements of Atlantic

Volume 40, Number 5 1263 ties in various seral stages,some also contain sizable lakes, ponds, marshes, bogs, and swamps. In a large number of bays,natural fire hasbeen suppressed for so long that the plant commumt•es in them are now mature bay forests Little field work has been conducted to survey birds within or adjacent to pocosinhabitats. In fact, the only no- table studiesin pocosinareas have been by Brooke Meanley on the Dismal Swamp fauna of northeasternNorth Carolina and southeastern Virg•ma (Meanley 1968, 1969, 1976, 1979), but none of these was in typical pocos•n habitats. Authors have commented on the lack of information on vertebrates associated with pocosins and succes- sionally related southeasternCoastal Plain habitats(e.g., Wilbur 1981,Shar- itz and Gibbons 1983). However, sev- eral reports on the bird life of succes- sional stages of pocosin-associated communities have recently appeared Lynch (1981) reported on a six-hour surveyof birdshe conductedin a young bay forest in North Carolina and Ter- williger and Rose (1984) compared breedingbirds associatedwith Atlantm White Cedar and maple-gumforest •n the Dismal Swamp. The only actual publishedstudy on the birds of a true pocosincommunity is that of Clark and Potter (1982). This profile of various plant communities on the North Car- olina Biological Survey study site at McCain, Hoke County, North Carohna, provides a partially annotated list of breeding birds of the six major plant communitiesoccurring on the property One of theseis a five-acreCarolina Bay that is dominated by high shrub po- cosin,from which 21 speciesof nesting birdsis reported.With theseexceptions, Figure 1. Pocosinareas in North Carolina (modifiedfrom Richardson1981). published information on the awan fauna of pocosinsystems and associated successionalstages is unavailable. Pocosinhabitats are widespreadgeo- graphically and can be found from White-cedar forests in southeastern cosincommunities. Although this study southeastVirginia to northernFlorida North Carolina. White-cedar forests does not cover Carolina bays per se, and are estimatedto have originallyoc- and savannas are both closely allied severalspecific types of vegetativecom- cupiedover 20% (2.25 million acres)of with pocosins. Additionally Wells, munitiesthat I studiedwere growingin the Coastal Plain of North Carolina (1932), Woodwell(1956), and Sharitz thesebays. Carolina bays vary in size and Gibbons (1983) provided good from only a few acresto many hundreds alone (Fig. 1). Wells (1946) noted that overviewsof pocosin vegetation;and of acres.An exposedsand rim of varying there were over 300 squaremiles ofpo- Wellsand Whitford (1976) presentedan width normally occursaround the pe- cosin in just three southeasternNorth outline of the serial development and rimeter. Thesedepressions are naturally Carolina counties. Since that time a successionalfate of stream head swamp wetter at all seasons than most sur- high.percentage of theseareas has been forests, pocosins, and savanna com- rounding areas, contrastingmarkedly drained, or partially drained, and munities. with the dry sand rims which support clearedfor agricultural or silvicultural Carolina bays must also be men- xeric plant communities.While many purposes;some have been dammed for t•oned in that they often support po- Carolina bays host pocosincommuni- mill ponds. Other areashave been pro-

1264 American Birds, Winter 1986 Protracted tectedfrom fire for solong that the plant communities, through natural succes- :•:• =POCOSIN COMMUNITIES sion, have outgrown pocosincommu- nity serialstages. There is some uncertainty about the extent of the loss of these habitats and the needfor concern.Reports by Heath (1975) and Richardson et al. (1981) provide a generaland gloomy summary of the projectedfuture of theseCoastal Plain wetlands. Most other studies have drawn on these sources as the basis for major concern for vanishing pocosin habitats.Richardson (1983) statedthat only 31% of North Carolina pocosins remain in a natural state. McMullan (1984) hassuggested that the reasonsfor concern may be lessserious than pre- viously stated, owing to faulty data sourcesand incomplete or nonexistent inventories. McMullan (1984) also demonstrated,in an analysisof a 300- year historical study of land use on the Albemarle-Pamlico Peninsula of North Carolina, that pocosin communities have persistedin spiteof a long history of rearing and draining, and that many ComparativelyShort presentday pocosinshave developed (or TIME redeveloped)on abandonedfarm lands. Assumingthat the answeris somewhere recent infrequent suppressed fire or fire fire betweenthe extremesexpressed in these disturbance reports, it appearsthat the original es- Figure 2. Proposedvegetation development of pocosinsand associatedplant communitiesas timatesof habitat lossare too high, but related to disturbance,time, and hydroperiod. that concernfor lossof pocosinhabitats isjustified, although perhaps overstated. Likewise, sincelittle information on the vertebrate fauna associatedwith poco- sins is available, definitive statements have not been investigatedin terms of Community development made by previous authors concerning their bird faunaanywhere. Although the and structure wildlife values of pocosins were pre- information presented here pertains Various environmental factors dic- mature. For example, Lee and Potter only to North Carolina, I suspectthat tate the type ofpocosincommunity that (1984) and Clark et al. (1985) demon- thesefindings would generallyapply to developson a site. The most conspic- strate that the bird and mammal asso- other areasin the Southeast.My efforts uous factors are surface and sub-surface ciates are mostly ubiquitous forms to date have been focused on species soil types, hydroperiod,and fire. The and that for mammals, densityis gener- inventoriesof a large number of differ- importance of the regularity and inten- ally low. ent shrub dominated communities sity of fire asit relatesto season,hydro- An overview of the breeding bird throughoutthe North Carolinal Coastal period, wind, and the accumulation of fauna of these communities is overdue. Plain. While I consider the findings combustiblevegetation cannot be over- Currently,discussions about the unique more than preliminary, problems as- stated.Natural fire and fires set by Na- biological value of pocosinsand con- sociatedwith samplingthe wide array tive Americans for game exploitation siderationfor their use in agribusiness, of pocosin communities and the influ- and later by Europeans for livestock silviculture, peat mining, and waste ence of geographicaldistributions on range managementwere all important disposalare commonplace;but in most portions of the fauna at any particular for long-term maintenance of various cases detailed information on which to site make it impractical to provide serialstages ofpocosins. Fire prevention basemanagement decisions is lacking. elaborate comparisons of relative policies were detrimental to certain For the last four years I have been sur- abundance and density of speciesin communities (particularly savannas), veying the birds of pocosinsand their specifichabitat typesat different local- but recentunderstanding of the impor- associatedplant communities in the ities. Furthermore, the impenetrable tance of consistentcontrolled burning North Carolina Coastal Plain. nature of the plant communities makes in certainsoutheastern vegetation types With exception of the previously it difficult to censushabitats by estab- for gameand habitat enhancementand mentioned studies,pocosins and their lished standardspot mapping or tran- wildfire control has, in part, alleviated successionallyassociated communities secttechniques. this problem.

Volume40, Number 5 1265 The characteristicand conspicuous plants of pocosinsand successionally relatedcommunities are comparatively few. In most instanceseach speciesoc- curs in a majority of the vegetative communitytypes and only their relative abundance or growth forms change. These variations in relative composi- tion, however, may be dramatic, both visually and ecologically,and are ob- viously important influences on the spedescomposition of the breedingbird fauna. It is generally understoodthat birds are recruitedby growth form, not plant species,as far as territorial needs are concerned. This is perhaps dem- onstratedmore dramatically in pocosins and their later serial stagesthan any- where else because the major plant communitiesvary more in growthform than in speciescomposition. The major plant associates(alphabetically by ge- nus, see Appendix 1) are Red Maple, Wire Grass,Atlantic White-cedar, Titi, Loblolly-bay, Sweet Gallberry, Ink- berry, Fetterbush, Sweet-bay, Black- gum, Redbay, Pond Pine, "Bamboo" or Laurel-leavedGreenbrier, Pond Cy- press, and Honey-Cup. Species less uniformly distributed include, Giant Cane, sedges, rushes, Sheep-Laurd, Longleaf Pine, Loblolly Pine, and blue- berrys.These latter speciesare, however, occasionallythe dominant vegetation on certain sites. Major pocosincommunity typesin- clude shrub bogs with scatteredPond Pine overstory, mixed conifer-hard- wood pocosins,and shrub-pine savan- nas. Early successionalstages of all of these types appear to be grass-sedge communities and later ones mature pine-hardwoodand evergreenbay for- ests.With long-termabsence of fire, all eventuallybecome deciduousbay for- ests. In these later stagescypress and Blackgumemerge on the siteswith pro- tracted hydroperiodsand Sweet Gum and pineson dryer ones.Thus, pocosins can be viewed as intermediate succes- sional communities, often maintained in a subclimaxstage by fire and hydro- period, with the mature vegetative stagesbeing suppressed for long periods on the wettestsites, but developingrel- atively quickly on dryer ones.Figure 2 depictsa generalsuccessional model for pocosincommunities discussed.Figure 3 showsvarious examples of commu- Figure 3. Pocosinrelated communities (allphotos/David S. Lee): a) ShortleafPine shrubbog. nities discussed. b) LongleafPine savannawith shrubbog developing in low wet area c) LongleafPine/Wire Grasssavanna. d) YoungAmerican White-cedar stand growingfiom a burnedout shrubpocosin. The developmentof Atlantic White- e) MatureAmerican White-cedar.forest showing winter wind and icestorm damage f) Carolina cedar forestsis unusual as this species bays with exposedsand rims. needs fire or other disturbance to re-

1266 American Birds, Winter 1986 movecompetitive vegetation so that the seedlingscan develop. However, ex- tremelyhot firesdestroy the peat soil and cedar forestsdo not appear. Low- intensity,fast-moving fires on the other hand,do not destroyenough of the root stocksof competitiveshrubs for thisce- dar to become well established. Even when established,Atlantic White-cedar isextremely fire susceptible, and forests persistonly in the totalabsence of fire. Young cedar forestsare usually pure standsof nearly even-agetrees and the densityof suchforests often inhibits the establishmentof other tree speciesfor about40 years.After that time thetrees beginto thin outand the nature of their crowns changes,permitting light to penetrateto theforest floor. Terwilliger and Rose (1984) commented on the managementproblems of maintaining advanced-agecedar forest. At thisstage bay forestsdevelop rapidly, although individual cedarsin these forestsmay persistfor long periods. The open sa- vanna community, on the other hand, requirescontinual policing by fire.If fire is suppressedfor severalconsecutive years, many characteristicsavanna plantsvanish. The precedinganalysis is summarized from Wells (1946), Buell and Cain (1943), Kologiski (1977), Wells and Whitford (1976), and per- sonal observations.

Methods

The avifauna of pocosinsand asso- ciated communities was studied and inventoried in portions of Bladen, Brunswick, Dare, Hoke, and Pender counties of North Carolina between April 1980 and June 1984. Inventories(species list and species dominance)were compiled for all sites and habitatssurveyed. Density of the plant covermade it impracticalto sur- vey many of the breedingbird com- munitiesby traditionalspot mapping or transect methods, and a more ex- pedientcomparative method was nec- essary.In someareas roads or otherav- enues of access allowed transects throughmonotypic habitats. The major surveyproblem was obtaining compar- ative data while excludingedge effects. Countsof singingmales were made for each habitat studied from walked transects(not alwaysstraight) 0.5 to 2

Volume40, Number 5 1267 miles in length, ones driven vath 0.25 to pocosln habitats. Becausemost are jacent communities,and land drainage mile, 5-minute stops (up to 8 miles), ubiquitousin distribution,no segment operations also affect the amount of and from scattered specific stations of the fauna can be used successfullyto standingwater. vathin pure stands of certain habitat characterizepocosins. Some species ap- Althoughthe effectof fire in pocosins types.Additionally, at a few siteseco- pear to exist normally in low densities is logicallyan important factor in gov- tonal areas were so narrow that it was within true pocosinsand becomecom- erning the breedingbird fauna, its over- possibleto censustwo habitats simul- mon only in disturbed areas or ones all influence is certainlymore positive taneouslyby recording singing males with temporaryvegetative shifts caused than negative. Plants associatedwith from each habitat while walking tran- by fires or storms.Birds characteristi- pocosinsrespond quickly to burning; sectsor from listing stationsstraddling cally foundin one or more community thus the abundant new growth of her- two habitat types. Each major com- typesare listedin Table 1. baceous species and the temporary munity was studied at severaldifferent Natural ecotones,edges caused by opening of the shrub layers generally sites(usually areas separated geograph- land-use practices,and openingswere produce an increasein birds. icallyby distancesof 50-150 miles)and far more diverse than interiors of bay The order of successionalpatterns of were revisitedat different times during or cedar forests,and many birds in- the plant/breedingbird community re- the nesting season(late April to late cluded some edgeas part of their ter- lationships have been interpreted as June). To be surethat the total diversity ritories. This can be attributed to gen- follows(birds listed in approximateor- and relativecomposition of eachmajor erally good cover in ecotonalareas, to der of abundance).It shouldbe empha- community type was recorded,I arbi- a richer diversity of insect food, and sized that thesecategories do not rep- trarily determinedthat 1000 encounters perhapsto slightchanges in topography resent total faunal lists,but are the spe- w•th singingmales were necessarybe- that provide temporary refuge from ciesmost regularlyencountered in each fore the surveyof any particularcom- seasonalflooding for ground-nesting major community type. The relative munity was completed. In fact, after species. abundanceof each speciesis expressed about 200-300 encountersthe species In many areasthe plantcommunities as its percent of the total fauna of that composition of the fauna was well es- are located on hard subsoils that form community. tablished,and the number of additional natural basinsretaining surfacewater Early Stages--sedge/grass/rush(often speciesdiscovered was extremely low. in the organictopsoils. Root systemsof scattered pines, but canopy and Speciesadded after 200-300 encounters many of the bay foresttrees do not usu- shrubsremoved by fire or man) and were birds that representedone-half of ally penetratethe subsoilsand the lim- open pine savanna: one percentof the total avifauna or less. ited supportoffered by the shallowpeat Characteristic(>50%): PineWarbler Therefore, in some of the less typical soilsmakes larger trees extremely vul- (14%), Great CrestedFlycatcher (9%), pocosin-associatedcommunities, I did nerableto strongwinds and ice storms. Brown-headedNuthatch (7%), East- not live up to my ambitious attempt to This is magnifiedby the fact that many ern Meadowlark (7%), Common tally 1000 individuals. Non-vocal en- suchtrees are also "crown heavy" as a Yellowthroat (6%), Eastern Wood- counterswith somebirds (e.g., Turkey result of early competition with the Pewee(5%), Northern Bobwhite (5%) Vulture, hawks,woodpeckers) were also normally denseunderstory vegetation. Regular(2-5%): Red-headedWood- included. For crude comparisons of A visit to Carolina bays in Bladen pecker, American Crow, Tufted Tit- density in different communities, the County and to Atlantic White-cedar mouse, Prairie Warbler, Rufous- number of birds encountered/minute forestsin BrunswickCounty duringthe sided Towbee. duringprime surveyhours (0600-0800) spring of 1983 revealedextensive up- Remarks: A total of 38 species of and seasons(May throughthe firstweek rootingand limb breakage(particularly birds was encountered, 14 of which of June) were averaged. Red Maples,Redbay and cedars) caused were seen no more than three times by late March snow and ice storms. (i.e., eachless than 1%of population). Hurricanes and tornados would cer- SedgeWrens were found in altered tainlycause even greater damage. Buell open areas as well as in natural early and Cain (1943) observedareas where successionalstages, exhibiting a note- the weightof Smilax climbinginto the Results worthy distributional pattern. Most canopiesof cedarforests caused trees to surveywork wasdone in savannas,and uproot. Thus, natural openingsin ad- For the most part, the characteristic birds of early sedge/grasscommunities vanced successionalstages are com- avifauna of pocosinsand related com- are not included in the percentages munitiesare wide-ranging,common monplaceand theseopenings provide numerous sites for shade intolerant shown here or in Table 1. Figures and species(Table 1). Within the North relative densities are based on a savanna plantsand early successionaland eco- Carolina Coastal Plain only a few spe- study site in the Green Swamp in tonal faunas.Around suchopenings an ciesare geographicallyrestricted to the BrunswickCounty (Table 1). increasedavian diversity is apparent. extent that they would not be expected Intermediate Stages--shrub bog and wherever pocosin habitats exist, and Degree and duration of flooding of other pocosintypes: none of theseis a major component of pocosincommunities is extremelyvari- Characteristic (>50%): Common the fauna.Most of the true pocosinbirds able.Generally, areas with organicsoils Yellowthroat (16-27%), Rufous- discussedhere are opportunisticspecies have protracted hydroperiodswhereas sided Towhee (12-19%), Prairie that exploit early and intermediate thosewith mineral soilshave compar- Warbler (15-16%), Northern Bob- successionalstages of many plant com- atively short hydroperiods.Local to- white (5%), White-eyed Vireo (4- munitiesand are certainlynot limited pography,the natureof soiltypes ofad- 7%).

1268 American Birds, Winter 1986 Table 1. Relative abundanceindex for birds regularly encounteredin pocosinsand associatedhabitats. Numbers represent abundance (percentto thousandth)for eachof six specificcommunities surveyed. Abundance exceeding 5% and mostconspicuous components of fauna in boldfaced type.

Pocosin

Opengrass Shrub Low shrub High shrub White-cedar Evergreen and savanna savannah pocosin pocosin forest bayforest

No. of encounters 355 175 1190 1095 354 1057

Approx. number hourssurveyed 6.75 3.5 7.5 8.5 3 20.5

Individuals per minute 1.15 .53 3.39 2.89 2.75 1.34 Species Northern Bobwhite .051 .093 .049 .029 .011 .015 Mourning Dove .012 .021 .051 .017 .023 Red-headed Woodpecker .022 .006 Red-bellied Woodpecker .004 .022 .006 Downy Woodpecker .003 .002 .009 .003 .013 Hairy Woodpecker .001 .001 .003 Northern Flicker .019 .018 .016 .013 .006 .004 Eastern Wood-Pewee .054 .009 .015 Acadian Flycatcher .006 .018 Great Crested Flycatcher .090 .056 .021 .009 .028 .029 EasternKingbird .009 .014 .023 Blue Jay .003 .010 .003 .006 American Crow .045 .009 .002 .011 .014 .024 F•sh Crow .016 .009 .001 .006 .006 .009 Carolina Chickadee .003 .013 .012 .050 .040 Tufted Titmouse .022 .093 .013 .014 .079 .048 Brown-headed Nuthatach .070 .009 .013 .003 .011 Blue-grayGnatcatcher .003 .003 .006 .007 Carolina Wren .016 .046 .027 .036 .038 Gray Catbird .047 .044 .006 .021 Brown Thrasher .003 .006 .001 White-eyed Vireo .003 .037 .036 .073 .037 .072 Northern Parula .001 .022 .105 Black-throated Green Warbler .001 .001 .147 .018 Yellow-throated Warbler .006 .006 .003 .023 Pine Warbler .141 .012 .001 .011 .055 Prairie Warbler .041 .205 .147 .163 .113 .032 Kentucky Warbler .037 .005 .015 .017 .010 Prothonotary Warbler .004 .093 .084 Worm-eating Warbler .002 .003 .003 .040 Swainson's Warbler .002 .022 .010 Common Yellowthroat .058 .242 .270 .160 .037 .016 Hooded Warbler .014 .004 .028 .070 Yellow-breasted Chat .006 .010 .011 Northern Cardinal .009 .046 .013 .024 .056 .030 Rufous-sided Towbee .022 .074 .190 .123 .042 .050 Eastern Meadowlark .070 .002 Brown-headed Cowbird .009 .004 .004 .039

Total all other birds .167 .009 .029 .102 .093 .041

Total species 38 16 40 51 40 44

Volume40, Number5 1269 Regular (2-5%): Gray Catbird, Crow, Carohna Chickadee, Tufted th•s woodpecker.S•ngle clans were en- Mourning Dove, Great CrestedFly- Titmouse, Carolina Wren, Gray countered on study sites in Dare and catcher, Carolina Wren, Northern Catbird, Worm-eatingWarbler, Yel- Pender counties. The birds were also Flicker, Eastern Kingbird, Eastern low-throated Warbler, Prairie War- found in savannas,typical habitat for Wood Pewee, Carolina Chickadee, bler, Northern Cardinal, Rufous- the species,in BrunswickCounty and Tufted Titmouse, Brown-headed sided Towhee. on sandridges adjoining Carolina Bays Nuthatch, Hooded Warbler, North- Remarks: Bay forest supports the in Bladen County. Combined the spe- em Cardinal. most interestingand diverseassem- ciesmade up lessthan 0.10% of the total Remarks:The five characteristicspe- blage of birds with most of the 44 fauna of communities censused. Th•s ciesmake up slightlymore than 70% speciesencountered being regular wasthe only EndangeredSpecies found of the breedingfauna. Of 40 breeding componentsof the community. Sev- during the study, and pocosinsprovide speciesencountered, over one-half eral peripheral as well as semi-dis- only sub-marginal habitat for th•s made up less than 1% of the fauna. junct bird populationsseem locally woodpecker. Informationfrom studysites in Holly dependent on these communities. Northern Rough-wingedSwallow: Th•s Shelter, Pender County and Dare Bay forestscontain a remarkabledi- swallowhas expanded its rangeinto po- County, including both short and versityof breedingParulidae, with no cosinareas by usingbanks of drainage high shrub pocosins.House Wrens fewer than 13 speciesrepresenting ditchesas nestsites. At presentit is un- (1.1%) were found at both thesesites over 57% of faunal density,and sev- eral others were noted in transitional common and restrictedto specificsites and habitattypes as well asin similar in southeastern North Carolina. habitatsin BrunswickCounty. These communities. Bay forestswere sur- birds are south and east of their doc- veyedat four sitesin BladenCounty. House Wren: Prior to thesestudies, the umented range and appear to be Only 16 of the 44 speciesrecorded House Wren was not recognizedas oc- confined to this habitat in eastern eachmade up lessthan 1%of the total curring in the nestingseason on the North Carolina. population. Most birds included North Carolina coastal plain. Nev- AdvancedStages--Atlantic White-ce- portionsof edgesor openingsin their ertheless,it is likely that thesehabitats dar forest(mature denseforest) territories. havebeen used for someyears and been Characteristic (>50%): Black- simply overlooked.In July 1934, Bur- Other breedingbirds encountered but throated Green Warbler (15%), Prai- leigh (1937) found wrensnear Stumpy considered transient, atypical or rare rie Warbler (11%), Prothonotary Point in Dare County, and at Beaufort, componentsof the fauna (followed by Warbler (9%), Tufted Titmouse(8%), Carteret County, but theserecords have percent relative dominance combined Northern Cardinal (6%), Carolina been neglected.Both towns are in po- for all habitats): Green-backed Heron Chickadee(5%).' cosinareas, and it appearsthat pocos•ns (0.05%), Wood Duck (0.05%), Black Regular (2-5%): Red-bellied Wood- form the only sites in eastern North Vulture (0.21%), Turkey Vulture pecker, Great Crested Flycatcher, Carolina in which this wren lives. Birds (0.88%), Red-shouldered Hawk White-eyed Vireo, Swainsoh'sWar- presentlyinhabit a few shortshrub po- (0.24%), American Kestrel (0.03%), bler, Northern Parula, Common cosinsin Dare (Potterpers.comm.) and Yellow-billed Cuckoo (0.62%), Barred Yellowthroat, Hooded Warbler, Brunswick counties. Owl (0.03%), Chuck-will's widow Brown-headed Cowbird, Rufous- sided Towhee. (0.03%), Chimney Swift (0.27%), Ruby- Sedge Wren: Although no positive throatedHummingbird (0.03%), Belted Remarks: Forty speciesof birds were evidenceof nestingwas found, Elmse Kingfisher (0.05%), Red-cockaded encountered,14 of which each made Potter&ers. comm.)found Sedge Wrens Woodpecker (0.08%), Pileated Wood- up lessthan 1% of the fauna. White- on territory from April 4 to June 2 •n pecker(0.38%), Purple Martin (0.03%), breastedNuthatches (up to 2.5% at early grass-sedgesuccessional pocosin Northern Rough-winged Swallow one site) replaced Brown-headed development in Dare County. Th•s Nuthatches in this habitat. Individ- (0.24%), Barn Swallow(0.05%), White-' representsa slight southwardrange ex- breasted Nuthatch (0.24%), House uals of many specieswere restricted tension from the scattered records of Wren (0.38%), Sedge Wren (0.11%), to areas of the forest where part of coastalMaryland and Virginia. Eastern Bluebird (0.43%), Wood their territorycontained forest edges. Thrush (0.03%), European Starting Wood warblers made up 50% of the Wood warblers:As a group and as in- (0.13%), Yellow-throated Vireo total fauna density and were repre- dividual speciesthese comprised the (0.13%), Red-eyed Vireo (0.13%), sentedby 11 species.Atlantic White- dominant component of all habitats Ovenbird (0.21%), Summer Tanager cedar forests were surveyed in surveyed.For example,69 singingmale (0.08%), Indigo Bunting(0.32%), Bach- Common Yellowthroats were tallied •n Bladen, Brunswick, and Dare coun- ties. man's Sparrow (0.16%), Red-winged a censusstrip one-half mile longand 40 Blackbird (0.05%), Common Grackle yards wide in a low shrub pocosin •n --Evergreen bay forest: (1.61%) and Orchard Oriole (0.34%). BrunswickCounty. Additionally, sev- Characteristic(>50%): Northern Pa- The total of all miscellaneous birds in eral speciesthat were either unknown rula (10%), Prothonotary Warbler all habitatswas 7.66% of faunaldensity. from the Coastal Plain, or at least con- (8%), White-eyed Vireo (7%), sideredvery rare, were found asregular Hooded Warbler (7%), Pine Warbler Notes on selectspecies components of tall shrub pocosinsas (5%). well as bay and white-cedarforest. No Regular (2-5%): Mourning Dove, Red-cockadedWoodpecker: Low shrub fewer than 15 of the 16 speciesknown Great Crested Flycatcher, American pocosinssupported small numbers of to breed on the Coastal Plain of North

1270 American Birds, Winter 1986 Carohna were found dunng this study, pocos•nswtth dense shrubs had the Summary and dtscusston but severalwere not actuallyrecorded most birds but some, seeminglyiden- tical, siteswere unoccupied. dunngcensus periods; they were limited Pocosinsand associatedplant com- to peripheralcommunities, or atypical Black-throated Green Warbler: These munities occupy an extensiveportion roes not censused. Black-and-white warblers were abundant in Atlantic of the southeastern Coastal Plain. Re- Warbler, Louisiana Waterthrush,and White-cedar forests, but in all other cent concernover developmentor po- American Redstart all seemed to be mature forest communities they were tential exploitation of these areas has hm•ted to advanced forest stagesthat sporadicallydistributed. However, Pot- raisedquestions concerning the impor- cannot be consideredas pocosinasso- ter (pers. comm.) found them common tance of their preservation.This study, crated communities. in tall Pond Pine pocosins in Dare the first attempt to systematicallyeval- Worm-eating Warbler and Ovenbird: County. uate the avifauna of pocosins,indicates Both warblers were regularly, though Pocosinsprobably were important as that while the avifauna composition •s not abundantly, encountered in ad- precolonial habitat refugia. The subcli- interesting, there may be insufficient vancedstage successional communities. mactic communitiesand complex zo- justification for protection of these Thesebirds seemed to persistsouth and nation of pocosinsprovided habitats for habitats on ornithological grounds eastof their documentedbreeding range earlyand intermediatesuccessional bird alone. Index speciesare lacking and m bay and cedarforests, but are absent, speciesthen that would not otherwise most associatesare ubiquitous. Except or nearly so, from other CoastalPlain have occurredregularly in most Coastal for marginal populations of Red-cock- areas.Kentucky Warblersexhibited a Plain habitats.It appearsthat they may adedWoodpeckers, they do not include s•mfiardistributional pattern, but they have provided a reservestock of birds speciesof national concern. This does were also common in high shrub po- that populated short-lived natural not imply that these communities cosmsthroughout the North Carolina openingsin other southeasternplant shouldnot be preservedand managed Coastal Plain. communities. Today pocosinsand re- for their own sake,or for the refugethey SwainsoWs Warbler: Generally re- lated communities are not critical to may provideto unique flora. garded a warbler of southernswamps most mid-successionalstage bird species The habitat diversity found in the and canebreaks, Swainsoh's were found becausegrazing, mowing, lumbering, stagesofpocosin successionmaintained as regular,though not common inha- and other development produce and by storms,flooding, and fire, collectively b•tatsof cedar and bay forestcommu- maintain a wide array of early and in- allows considerable faunal diversity mt•es. They were found in every bay termediatesuccessional stages over ex- over broad areas.At specificsites, how- forest visited in Bladen County. tensiveareas of the CoastalPlain. Thus, ever, diversityis seldomhigh. In most for bird speciesalready associated with cases,the uniformintermediate pocosin Prairie Warbler: Although it was pre- pocosinsand therebywidely distributed stagesof community developmentsup- dicted that the Prairie Warbler would acrossthe CoastalPlain, local expansion port a low diversity but a high density be an important component of shrub into suitable disturbed habitats is not of breedingbirds. Advanced stagesof communities, the abundance of this surprising.In otherwords, species that community development, which are speciesin Atlantic White-cedar com- may have been common only in po- more structurallydiverse, may support munitiesand near absencein bay forest cosinsin the past owing to ecological a higherdiversity, but a modestdensity. •sd•fficult to explain.Nolan (1978) and restrictions,are now able to exploit a Fire, flooding,and limited seasonalfood Terwilligerand Rose(1984) notedthat wide array of disturbed community resourcessingly and collectivelydictate D•smal Swamp cedar stands were types.Except for maritime shrubthick- diversity and density of birds in ways amongthe few placeswhere this warbler ets on barrier islands, there were few that to date have not been measured. has been reported in closed habitats. precolonial natural communities other In the past, pocosin communities Terwilligerand Rose(1984) providean than pocosins that could have sup- may have been more important to the explanationfor this occurrencethat in- ported speciescurrently regarded as CoastalPlain avifaunathan they are to- volvesPleistocene evolutionary behav- ubiquitous on the Coastal Plain. day. Since historical times land man- ioral history of the species.In view of The true pocosin fauna generally agementpractices have openedup vast the relativelyrapid growth of Atlantic consistsof specieswith wide ecological areasfor early successionalspecies that White Cedar from shrub level to tree tolerance with the ability to exploit in the past were probably ecologically height,it seemsjust as likely that suc- small plots of early successionalstages confined to pocosinsand other natu- ceedinggenerations continue to return and areasdisrupted by human activities. rally suppressed serial community to these cedar forests to nest because The more advancedserial stages,no- stages. they are site imprinted. Becausecedar tably bay forestand Atlantic White-ce- From a zoogeographicperspective, a forests are the result of clearing and dar forests,however, support an inter- number of Atlantic Coastal Plain spe- rapidly colonizingcedar seedlings, this esting assemblageof breedingspecies cies presentlyreach either northern, processbecomes all the more feasible. includingseveral birds that, prior to this southern, or easternlimits of distribu- Onans (1971) proposed that site im- study, were consideredlocally rare or tion in pocosinrich areas.The ecolog- pnntingwith successionalchange is one were mostly unknown in the south- ical natureofpocosin habitats, and per- way in which a speciescan exploit new easternCoastal Plain. While interesting, hapsthe ability of theseareas to buffer habitatsand expand distributions. these communities are not really po- temperature extremes, is certainly a Yellow-breasted Chat: Densities of cosins,although it could be arguedthat factor in the geographicdistribution of chatsvaried considerablyin pocosins. it is normal to passthrough a pocosin somesoutheastern species. Tall shrubpocosin and recentlyburned stagebefore a bay forestevolves. In general,the practiceof lumping a

Volume 40, Number 5 1271 variety of communities,ranging from Appendix1. Vegetativespecies arranged in alphabeticalorder by genus. open grasssavannas to mature hard- RedMaple Acerrubrum wood swampforest, as pocosinshas al- Wire Grass Aristida stricta loweda wide arrayof speciesto be con- Giant Cane Arundinaria gigantea sidered as pocosin inhabitats. Such a sedges Carex,sp. systemallows Eastern Meadowlarks and AftanticWhite-cedar Charnaecyparisthyoides Bachman'sSparrows to be inhabitatsof Titi Cyrillaracerniflora the samecommunity in whichone finds Loblolly-bay Gordonialasianthus SweetGallberry Ilex coriacea Prothonotary and Worm-eating war- Inkberry Ilexglabra blers.When only the shrubbog stageof rushes Juncus, sp. successionis regardedas pocosin,the Sheep-Laurel Kalmiacaroliniana fauna is characterizedby a small as- SweetGum Liquidambarstyracifiua semblage of early to intermediate Fetterbush L yonia lucida successional,shrub stage species. SweetBay Magnoliavirginiana Blackgum Nyssasylvatica Misconceptionsand continued mis- Redbay Perseaborbonia quotations in newspaperaccounts, en- ShortleafPine Pinus echinata vironmental impact statements, and LongleafPine Pinuspalustris scholarlypublications make it difficult Pond Pine Pinus serotina to separateemotional concerns from the LoblollyPine Pinustaeda emergingfacts regarding pocosins. Def- Laurel-leavedGreenbrier Smilax laurifolia PondCypress Taxodium(distichurn) ascendans inition and interpretation have caused blueberries Vacciniurn, sp. problemswith inventoryof habitat and Honey-Cup Zenobiapulverulenta projection of habitat loss.The general classificationof pocosinsas a type of wetland,although correct, has led some environmentaliststo assumehigh wild- ment to studycertain areasin the state KOLOGISKI, R. L. 1977. The phytosocaot- hfe values associatedwith pocosinsde- parkssystem. Eloise Potter, Mary Kay ogyof the Green Swamp,North Carohna spitethe previouslack of systematicin- Clark, John B. Funderburg, Jr., and N.C. Agfic. Exp. Sta. Tech. But. No ventories. Statements about pocosin Gilbert Grant (all North Carolina State 250.100 pp. wildlife valuesare further hamperedby LEE, D. S., and E. F. POTTER. 1984 Museum) assistedin portionsof the in- Quantifying breeding birds assocaated the lack of comparableinformation for ventory effort. David K. Clark, Eliza- with pocosins:an exercisein commumty other wetland habitats and historical bethtown,provided assistance in aerial definitionand myth slaying.[Abstract] comparisonsof the avifauna in the surveysand lodgingduring the Bladen 65th Annual Meeting of the Wilson Or- Southeast in general and the North Countyphase of this study.Potter con- nithologicalSociety. Carolina Coastal Plain in particular. In ducted extensive surveys of Dare LYNCH, M. 1982. Breedingbirds of Hall this study I have provided the first at- County and her findings were most Swamp Pocosin, Martin County, N C The Chat 46(4):93-105. tempt to document the breeding bird helpfulin preparingthis manuscriptand MEANLEY, B. 1968. Birds of the Great fauna associatedwith pocosin com- her commentsgreatly improved it. Dismal Swamp. Atlantic Nat. 23:141- munities, and I hope to have started a 142. processthat will allow future researchers --. 1969. Notes on Dismal Swamp bards to define and separatethe fauna of po- Raven 40:47-49. cosins from that of their closely asso- --. 1976. A censusof breedingbirds an the Dismal Swamp, VA. Raven 47 40- ciated but very different communities. 43. --. 1979. An analysis of the birdlife of the Dismal Swamp. pg 261-276 in The LITERATURE CITED Great Dismal Swamp. P. W. Kirk, Jr, ed. University Pressof Virginia., Char- BUELL, M. F., and R. L. CAIN. 1943. The lottesville.427 pp. successionalrole of southern white cedar, McMULLAN, PHILIP S., JR. 1984. Land- Chamaecyparisthyoides, in southeastern clearingtrends on the Albemarle-Pamhco North Carolina. Ecology24:85-93. Peninsula: 300 years of developmentan BURLEIGH, T. D. 1937. Bird life on the a wetlands region. Privately published, North Carolina coast. Auk 54:452-460. McMullan ConsultingDurham, NC. 131 CLARK, M. K., and E. F. POTTER. 1982. PP. Third annual breedingbird foray: Hoke NOLAN, V., JR. 1978. The ecologyand be- County, N.C. The Chat 46(2):29-37. havior of the Prairie Warbler (Dendrotca ACKNOWLEDGMENTS __, D. S. LEE, and J. B. FUNDER- discolor).Omithotogical Monogr. 26 t- BURG, JR. 1985. The Mammal Fauna 595. The United States Fish & Wildlife of Carolinabays, pocosins, and associated ORIANS, G. 197i. Ecologicalaspects of be- Service provided partial support for communities in North Carolina: an havior. Pp. 513-546 in Avian Biology overview.Brirnleyana 1t: 1-38. Vol. I.D. S. Farner and J. R. King, ed, studiesin Dare County under contract HEATH, R. C. 1975. Hydrology of the At- Academic Press,NY. number 14-16-0004-81-056. Bryan bemarle-PamlicoRegion, North Caro- RICHARDSON, CURTIS J., Editor. 1981 Taylor, North Carolina State Parks, lina. Water-ResourceInvestigation 9-75, Pocosinweftands: an integratedanalys•s provided permission and encourage- U.S. GeologicalSurvey. Raleigh 98 pp. of CoastalPlain freshwaterbogs in North

1272 AmericanBirds, Wanter 1986 Appendix 2. Bird speciesarranged in taxonomicorder. Carohna Hutchinson Ross Publishing Co., Stroudsburg.PA. 364 pp. Green-backed Heron Butorides virescens __. 1983. Pocosins:Vanishing wasteland Wood Duck Aix sponsa or valuable wetlands.BioScience 33(10). Black Vulture Coragypsatratus November 1983. Cathartes aura Turkey Vulture SHARITZ, REBECCA R., and J. W. GIB- Red-shouldered Hawk Buteo lineatus BONS. 1982. The ecologyof Southeast- American Kestrel Falco sparverius Northern Bobwhite Colinus virginianus ern shrub bogs(pocosins) and Carolina Mourning Dove Zenaida macroura bays:a communityprofile. U.S. Fishand Yellow-billed Cuckoo Coccyzusminor Wildlife Service,Division of Biological Barred Owl Strix varia Services,Washington, D.C. FWS/OBS- Chuck-will's-widow Caprimulgus carolinensis 82/04. 93 pp. Chimney Swift Chaetura pelagica TERWILLIGER, K. A., and R. K. ROSE Ruby-throatedHummingbird Archilochus colubris 1984. Breedingbirds in cedar standsin Belted Kingfisher Ceryle alcyon the Great Dismal Swamp. Am. Birds Red-headed Woodpecker Melanerpes erythrocephalus 38(1):24-27. Red-bellied Woodpecker Melanerpes carolinus TOOKER, W. W. 1899. The adapted Al- Downy Woodpecker Picoidespubescens gonquin term "Poquosin."Am. Anthro- Hairy Woodpecker Picoides villosus pol., Jan.:162-170. Red-cockaded Woodpecker Picoides borealis WELLS, B. W. 1932. The natural gardens Northern Flicker Colaptesauratus of North Carolina. University of North Pileated Woodpecker Dryocopuspileatus Carolina Press,Chapel Hill. 458 pp. Eastern Wood-Pewee Contopusvirens ß 1946. Vegetation of Holly Shelter AcadianFlycatcher Empidonax virescens Great Crested Flycatcher Myiarchus crinitus WildlifeManagement Area. N.C. Dept Cons. Develop. Bull. 2. 40 pp. EasternKingbird Tyrannustyrannus Purple Martin Prognesubis __ and L. A. WHITFORD. 1976. History Northern'Rough-wingedSwallow Stelgidopteryxserripennis of stream head swamp forests,pocosins, Barn Swallow Hirundo rustica and savannas in the Southeast. Jour Blue Jay Cyanocitta cristata Elisha Mitchell Sci. Soc. 92:148-150. American Crow Corvusbrachyrhynchos WILBUR, HENRY M. 1981. Pocosinfauna Fish Crow Corvusossifragus Pp. 62-68. in Pocosin Wetlands. C. V Carolina Chickadee Parus carolinensis Richardson,ed., Hutchinson Ross Publ Tufted Titmouse Parus bicolor Co., Stroudsburg,PA. 364 pp. White-breasted Nuthatch Sitta carolinensis WOODWELL, GEORGE M. 1956. Phy- Brown-headed Nuthatch Sitta pusilla tosociologyof Coastal Plain wetlands of Carolina Wren Thryothorusludovicianus the Carolinas. Mister's thesis. Duke House Wren Troglodytesaedon University. SedgeWren Cistothorusplatensis Blue-gray Gnatcatcher Polioptila caerulea --North Carolina State Museum, Eastern Bluebird Sialia sialis P.O. Box 27647, Raleigh, Wood Thrush Hylocichla mustelina North Carolina' 27611 Gray Catbird Dumatella carolinensis Brown Thrasher Toxostomarufum European Starling Sturnus vulgaris White-eyedVireo Vireo griseus Yellow-throated Vireo Vireofiavifrons Red-eyedVireo Vireo olivaceus Northern Parula Parula americana Black-throated Green Warbler Dendroica virens Yellow-throated Warbler Dendroica dominica Pine Warbler Dendroica pinus Prairie Warbler Dendroica discolor Black-and-white Warbler Mniotilta varia American Redstart Setophagaruticilla ProthonotaryWarbler Protonotaria citrea Worm-eating Warbler Helmitheros vermivora Swainson's Warbler Limnothlypis swainsonii Ovenbird Seiurus aurocapillus Louisiana Waterthrush Seiurus motacilla Kentucky Warbler Oporornisformosus Common Yellowthroat Geothylpistrichas Hooded Warbler Wilsonia citrina Yellow-breasted Chat Icteria virens Summer Tanager Piranga rubra Northern Cardinal Cardinalis cardinalis Indigo Bunting Passerinacyanea Rufous-sided Towhee Pipdo erythrophthalmus Bachman's Sparrow Aimophila aestivalis Red-winged Blackbird Agelaiusphoeniceus EasternMeadowlark Sturnella magna Common Grackle Quiscalusquiscula Brown-headed Cowbird Molothrus ater Orchard Oriole Icterus spurius

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