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Habitat Barriers to Movement of Understory Birds in Fragmented 944 ShortCommunications and Commentaries [Auk, Vol. 113 docarpusNational Park in southern Ecuador(ca. 100 COLLAR, N. J., L. P. GONZAGA, N. KRABBE, A. km north of Angashcola),where recent recordsof S. MADROI•O-NIETO, L. G. NARANJO, T. A. PARKER, cinctusderive (Bloch et al. 1991), and the Reservadel AND D.C. WEGE. 1992. Threatened birds of the Alto Quindio Acaime/Los Nevados National Park/ Americas: The ICBP/IUCN Red Data Book. In- Ucumarl RegionalPark complexin Colombia.Other ternational Council for Bird Preservation, Cam- potentiallyviable populationsof S.cinctus in Ecuador bridge, United Kingdom. probably occurwithin SangayNational Park (Chim- FJELDS•.,J., ANDN. KRABBE.1990. Birdsof the high borazo/Morona-Santiago) and the Cayambe/Coca Andes. Zoological Museum, University of Co- EcologicalReserve (R. S. Ridgely pets. comm.).Strin- penhagen, Denmark. gent protectionof theseareas should be stronglysup- LOISELLE,B. A., AND J. G. BLAKE. 1991. Temporal ported. variation in birds and fruits along an elevational Acknowledgments.--Sponsorsof the Amaluza ex- gradient in CostaRica. Ecology72:180-193. peditions are listed in Williams and Tobias (1994), O'NEILL, J. P., AND T. S. SCHULENBERG.1979. Notes and to these we are indebted. We also thank: Paul on the Masked Saltator, Saltator cinctus, in Peru. Atkinson, ThomasBrooks, Robert Clay, RobertRidge- Auk 96:610-613. ly, and Paul Salaman for reviewing the manuscript RENJIFO,L. M. 1991. Discovery of the Masked Sal- and providing ideas;Rodrigo Tapia-Castro (along with tator in Colombia,with noteson its ecologyand severalmembers of Arcoiris) and David Espinosafor behavior. Wilson Bulletin 103:685-690. assistancein the field; and Nancy Hilgert de Bena- RIDGELY, R. S., AND G. TUDOR. 1989. The birds of vides (with the staff of CECIA) for hospitality and South America, vol. 1. University of TexasPress, help during our visits to Ecuador. Austin. SENAR,J. C., A. BORRAS,T. CABRERA,AND J. CABRERA. LITERATURE CITED 1993. Testing for the relationship between co- niferous crop stability and Common Crossbill BLOCH, H., M. K. POULSEN,C. RAHBEK,AND J. F. residence.Journal of Field Ornithology 64:464- RASMUSSFaq.1991. A surveyof the montanefor- 469. est avifauna of the Loja province, southern Ec- WILLIAMS,R. S. R., AND J. A. TOBIAS. 1994. The con- uador. BirdLife International (Study Report No. servation of southern Ecuador's threatened avi- 49), Cambridge,United Kingdom. fauna. BirdLife International (Study Report No. COLLAR,N. J., M. J CROSSY,AND A. J. S?ATTERSFIELD. 60), Cambridge,United Kingdom. 1994. Birds to watch 2: The world check-list of threatened birds. BirdLife International, Cam- Received20 November1995, accepted29 February1996. Associate Editor: R. L. Hutto bridge, United Kingdom. The Auk 113(4):944-949, 1996 Habitat Barriersto Movement of Understory Birds in Fragmented South-TemperateRainforest KATHRYN E. SIEVING,1'3 MARY F. WILLSON,2 AND TONI L. DE SANTO2 •Departmentof WildlifeEcology and Conservation, University of Florida, 303 Newins-ZieglerHall, Gainesville,Florida 32611, USA; and 2PacificNorthwest Research Station, Forestry Sciences Laboratory, 2770 SherwoodLane, Suite 2A, Juneau,Alaska 99801, USA The temperaterainforests in SouthAmerica are re- and fragmented by agricultural expansion.Habitat strictedto southernChile and adjacentsouthwestern fragmentation has detrimental effectson forest bird Argentina(Vuilleumier 1985).Home to a numberof populations(Wilcove et al. 1984, Lovejoy et al. 1986, endemic bird species,some that are listed as endan- Hagan and Johnston1992, Donovan et al. 1995, Rob- gered or threatened(Glade 1988,Collar et al. 1992), inson et al. 1995). Among forest bird species,terres- this forestbiome is rapidly beingharvested for timber trial or understoryinsectivores repeatedly have been identifiedas highly sensitiveto forestfragmentation, particularly in the tropics (Leck 1979, Willis 1979, E-mail: [email protected] Karr 1982,Lovejoy et al. 1986,Sieving and Karr 1997) October1996] ShortCommunications andCommentaries 945 but also in temperateregions (Recher and Serventy All five speciesare permanentresidents, strongly ter- 1991, Willson et al. 1994). ritorial (pers. obs.), and feed primarily on inverte- One potential causeof bird populationdeclines in bratesgleaned from leaf litter andlow foliage(John- fragmentednatural habitatsis isolationby expansion son 1965,pers. obs.). of altered human habitats,which may restrict dis- Experimentalmethod.--We used taped song play- persalmovements between fragments (Opdam et. al. backs (Falls 1981, McGregor 1992a) to determine 1985, Dunning et al. 1995). As a preliminary step to whetherbirds were willing to crossnon-forested hab- assessingavian use of forestedcorridors between itats betweenforest patches where testanimals held fragments(Simberloff et al. 1992),we examinedthree territories. Territorial songsof each specieswere re- habitats that may serve as movement barriers in a corded from commercial and field recordings and temperaterainforest landscape in southernChile. Us- playedfrom speakersin a standardizedmanner (de- ing songplayback experiments (McGregor 1992a), we scribedbelow) to individuals holding territoriesnear quantifiedthe willingnessof five understoryspecies edgesof forestfragments. Our playbackssimulated a to enter previouslycleared areas between forest frag- potentialterritorial invasion and elicited agonistic ap- ments. proachesby targetbirds (Falls 1981, Becker 1982, Col- Studysite and species.--Isla Grande de Chilo• (41ø55'S, gan1989). During preliminary trials in theforest, all 73ø35'W)is a largecontinental island less than 10km five speciesresponded strongly to playbacksin and offshore; it has been isolated from mainland Chile for neartheir territories,making close approaches to the approximately I0,000 years (Willson and Armesto speakerwhile displayingaggressive behaviors. 1996). Agriculture is the dominant land-use pattern Playbackexperiments were conducted during daily in the northern, eastern, and southern portions of singing/activitypeaks for all species(0600-1000 and Chilo& Farmersof Chilo$use animal power to clear 1600-1830h, local time) during two breedingseasons their land. Thus, the processof pasture creation is (January,and Novemberto Decemberof 1994).All slow and the landscapeis still diverse; fields with five specieswere engagedin nestingactivities during intermediatevegetative cover are interspersedamong thesetimes (Johnson 1965, Sieving et al. unpubl.data), completely cleared fields and forest fragments.For- althoughactual stage of thenesting cycle varied among estedareas are maintainedby farmersfor wood ex- individuals.We usedthe followingprotocol. A 6-watt traction and winter livestockshelter. Our study sites portablespeaker (Sony SRS-58) attached to a hand- were located adjacent to forest patches,ranging in held tapeplayer by a 10-mcord was set 15 to 20 m size from less than one to several hundred hectares, from the forestedge. Up to four observers,including on private farms in northeasternChilo& In our ex- the tapeplayer operator,were stationed20 to 40 m periment, potential barrier habitatswere divided into apart on either side of the speaker.Thus, approxi- three categories:(1) open pasture,with short (<5 cm mately60 m of forestedge could be observedon either tall) grassoffering no cover to forestbirds; (2) scat- side of the speaker.After all observerswere in place tered cover, where stumps,sparse shrubs, fern clus- for 2 min,the target species' song sequence was played ters, piles of cut branches,and a few regenerating from the speaker.Songs were played for varying saplingswere interspersedwith patchesof grass;and amountsof time dependingupon the behaviorof (3) dense cover, comprisedof older clearingswith subjects.A responsewas defined as a bird moving continuouscover of bamboo(Chusquea sp.), ferns, sap- (while seenor heardsinging by any observer)to the lings, and b_hash piles. forestedge and in the directionof the speakerafter We selectedfive understoryspecies (four rhino- the playbackbegan. If no subjectresponded by ap- cryptidsand onefurnariid) that persistin fragmented proachingthe forestedge and speakerfrom the forest forestsof Chilo• (Willson et al. 1994).All are endemic interior after 5-10 min, the playbackwas terminated to the south-temperaterainforest of southernChile and not used.Once a subjecthad responded, however, and Argentina,but their geographicranges vary in the tape was playeduntil the subjecteither left the size. The smallestof the rhinocryptids is the Magel- forestedge and movedback to forestinterior, clearly lanic Tapaculo(Scytalopus m. magellanicus;ca. 11 g), refusingto enter the testhabitat after at least 10 min which also has the widest geographicdistribution of activeresponse (singing and/or pacingand perch- (Sibleyand Monroe 1990,Ridgely and Tudor 1994). ing at the edge),or until it enteredthe test habitat The Ochre-flankedTapaculo (Eugralla paradoxa; ca. 25 and approachedthe speaker.The latter behaviorwas g) and the ChucaoTapaculo (Scelorchilus rubecula; ca. recordedas a barrier entry that indicated the respon- 40 g) have very small geographicranges. The Black- dent was willing to enter the test habitat.Our ex- throatedHuet-huet (Pteroptochostarnii) is the largest perimentaldesign is describedby McGregor(1992b) of the four rhinocryptids(ca. 150 g) and has an in- as the single-measureapproach to quantifyingplay- termediategeographic range. Des Murs' Wiretail (Syl- back responses. viorthorhynchusdesmursii; ca. 10 g), the one furnariid, We were conspicuouswhen
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