494 THE WIl.SON JOURNAL OF ORNITHOLOGY • Vol. 119, No. 3, September2007 The Wilson Journal ofOrnithology 119(3):494^98, 2007 Yellow-breasted Chat and Gray Catbird Productivity in a Fragmented Western Riparian System Tawna C. Morgan, Christine A. Bishop, and Tony D. Williams* — ABSTRACT We studied the effects of habitat some conditions, habitat fragmentation may fragmentation on productivity ofYellow-breastedChat lead to a decline in population numbers (Sher- (Icteria virens) and Gray Catbird (Dumetella caroli- ry and Holmes 1992, 1993). npeanrsiiasn)eicnossyosuttehme.rnNeBsrtiitinsghdCaotelsumwbeirae ilnatearwienstiseorlnatreid- Robinson et al. (1995, 2000) documented a habitat patches than in continuous habitat patches for relationship between lower productivity and both species. We found no direct evidence that habitat increased habitat fragmentation in eastern for- fragmentation decreased productivity in eitherspecies. ests. Western riparian ecosystems are frag- Average fecundity did not significantly differbetween mented by nature because of frequent flood- continuous (2.54 fledglings forGray Catbird; 1.78 and ing, ephemeral water sources, and mountain- 1.67 fledglings for Yellow-breasted Chat in 2002 and ous topography (Hejl 1992, 1994; Ohmart 2003, respectively) and isolated sites (1.33 fledglings forGray Catbird; 1.78 and 0.87 fledglings for Yellow- 1994). The few studies that have examined the breasted Chat in 2002 and 2003, respectively). Terri- effects offragmentation in western landscapes tory size, as measured by mapping perch locations for have generally found a less direct relationship breeding adults, was smallerforYellow-breastedChats between habitat fragmentation and productiv- breeding in the Okanagan Valley (0.25 ha) than for ity (Tewksbury et al. 1998). chats in mid and high-density southern populations. We tested the hypothesis that fragmented However, overall fecundity and nest success were sim- ilar to more southerly populations. The.se results sug- habitat negatively affects reproductive success gest that both .species can persist in a relatively frag- of Yellow-breasted Chat {Icteria virens) and mented ecosystem. Received 30 June 2005. Accepted Gray Catbird {Dumetella carolinensis) in 17 October 2006. western riparian habitats throughout the Okan- agan Valley, British Columbia, Canada. Both species predominantly use wild rose {Rosa Riparian areas provide critical breeding and spp.) as a nesting substrate and forage in close stopover habitat for a large diversity of birds. proximity to one another (Cannings et al. Despite its importance to avian communities, 1987; TCM, pers. obs.). Yellow-breasted Chat there has been considerable loss of riparian and Gray Catbird nests are commonly within habitat throughout North America (Croonquist 5 m ofone another. The Okanagan Valley rep- and Brooks 1993). Thus, there is a need to resents the northwest fringe of the Yellow- understand avian population trends in relation breasted Chat’s breeding range (Eckerle and to habitat alteration. Several studies have doc- Thompson 2001), while Gray Catbird popu- umented a decrease in productivity and an in- lations extend farther north into northern Brit- crease in brood parasitism in association with ish Columbia (Cimprich and Moore 1995). increased habitat fragmentation (Wilcove and We compared reproductive parameters for Robinson 1990, Donovan et al. 1995). Under Yellow-breasted Chat with more southerly populations. ' Centre for Wildlife Ecology, Simon Fraser Uni- METHODS versity, 8888 University Dr., Burnaby, BC V3J 1S6, Canada. Aerial photographs ofriparian habitat in the 2Environment Canada, 5421 Robertson Rd., Delta, Okanagan Valley, British Columbia and Yel- BC V4K 2N2, Canada. low-breasted Chat surveys in 2001 were used ’Current address; Department ofBiology andWild- to categorize four large riparian patches (15- life, Institute of Arctic Biology, University of Alaska, Fairbanks, AK 99775, USA. 70 ha) as continuous and five smaller patches Corresponding author; e-mail: as isolated (<2 ha). Continuous sites were [email protected] tracts of unbroken riparian habitat that con- SHORT COMMUNICATIONS 495 tained >15 ha of unbroken habitat with flow- the mean number of fledglings per female per ing water. Isolated sites were small fragment- season. All females were single brooded. A ed patches of riparian habitat, primarily along second method of calculating fecundity, in roadsides and drainage ditches. Isolated sites which only successful nests were considered, were separated from other riparian habitats by was used for comparison with other studies. >5 km and had ephemeral water sources. All Percent parasitism was calculated as the per- sites were <600 m in elevation and were be- centage of all monitored nests that contained tween the International Boundary (49° 02' N, >1 Brown-headed Cowbird {Molothrus ater) 1 19° 27' W) and the city ofPenticton (49° 30' egg or young at any point during the nesting N, 119° 35' W). All study sites were used by cycle. Gray Catbirds were omitted from this chats in 2002. Three of the isolated sites and portion of the study because they reject cow- one continuous site were unused by Yellow- bird eggs. breasted Chats in 2003. Gray Catbird nests Spot mapping was conducted for 20 Yel- were monitored at the same sites as Yellow- low-breasted Chat territories (5 observations breasted Chats in 2003. All sites were moni- per territory) in 2003. Territory visits were at tored for breeding activity between 13 May least 3 days apart and observations were con- and 15 August 2002 and 2003. ducted for one-half hr during which all Yel- Surveys for breeding Yellow-breasted low-breasted Chat sightings were recorded. Chats and Gray Catbirds were conducted at Observations began 15 min after entering a all sites every 3 weeks to detectbreeding pairs territory. If a focal bird disappeared for more and locate nests. We checked nests using mir- than 5 min, observations were excluded from ror-poles or direct observation every 3 days the analysis. Time of observation, bird activ- during the nesting cycle to examine clutch ity, location, perch height, and any other com- size, cowbird parasitism status, hatch date, ments pertaining to bird behavior or surround- and nest success or failure (Martin et al. ing ecology were recorded for all sightings. UTM 1997). locations were taken after the observa- Nestlings were measured and banded with tion period based on maps and descriptive in- a single Canadian Wildlife Service band and formation recorded during the observation pe- three-band color combination at 6 (Yellow- riod. Territory size was calculated using breasted Chat) or 7 (Gray Catbird) days of Arcview GIS 3.1 area calculator. age. All nestlings were banded between 0700 We used STATISTICA (StatSoft 2000) for and 0930 hrs PDT. Adults were captured using all statistical analysis. One-way analysis of passive and target netting techniques. We variance (Sanders 1995) was used to test for banded adults with a three-band color com- differences in fecundity, clutch size, territory bination and an aluminum Canadian Wildlife size, nest initiation, and hatching and fledging Service band. All fieldwork was in accordance dates between years and patch type. Student’s with Simon Fraser University Animal Care r-test was used to test for differences in nest permit 663B-03, bird banding permit 10365 success between years. There were between CN, and Canadian Wildlife Service permit 59- year differences in nest success rate, and the 03-0426. We used mist nets for up to one-half data are presented and analyzed by year. Re- hr within territories at least 6 m from nests. sults were considered significant at P <0.05 Locally-recorded species-specific male songs and are presented as mean ± SD unless oth- were played for a maximum of 10 min to at- erwise stated. tract territorial males and females. We also RESULTS placed passive nets within territories near known flight paths, but did not play tapes to Twenty-five Yellow-breasted Chat nests capture some birds. were monitored in 2002 while 32 Yellow- Daily nest success probability was estimat- breasted Chat nests and 34 Gray Catbird nests ed using maximum likelihood (Hensler and were monitored in 2003. Initiation of incuba- Nichols 1981, Rotella et al. 2000). A nest or tion began the third week of May and contin- female was considered successful if >1 Yel- ued through 12 July in 2002-2003 for Yellow- low-breasted Chat or Gray Catbird nestling breasted Chats and 4 June to 12 July 2003 for fledged. Female fecundity was calculated as Gray Catbirds. Nest incubation, hatching, and — 5 496 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 119, No. 3, September2007 fledging dates were later in all years at isolat- ed sites for Yellow-breasted Chats and Gray m 00 lo 8.72 11.41 Catbirds (Table 1). There were no significant differences in clutch size (3-6 eggs) or number offledglings Jul Jul between years for Yellow-breasted Chats (Ta- 00 CO oc 'It 23 9 ble 1). Percent parasitism ranged from 0 to ^ E 04 44% for Yellow-breasted Chat nests during 2002-2003. There was no difference in fecun- "XTOTi. 0o—0i 0io0in 00O40' i^oinit 9.74 11.34 dity between parasitized and non-parasitized UJ females (2.26 ± 1.53 and 1.71 ± 1.74 fledg- 0li.n2g8s2,).reFsipfetcyt-isveevleyn,pter=cen1t.0o1f,Ydefllo=w-5b1r,eaPste=d 3 1£) “UC3ji _"35 I033—0 33 9Jul Jun CO 00 CO 28 Chat nests monitored fledged at least one nest- CO 04 ling in 2002-2003. Similarly, there was no csligunticfhicsainzte adinfdfefreecnucneditiny oefitYheerlloywe-abrreawshteend o —OO'; 'i—Int 00 d’00 7.94 10.96 Chat females breeding in isolated sites were compared with females in continuous tracts of habGirtaaty(CTaatbblierd1).clutch size ranged from two £I•Fs CFI (N (N — 2Ju7n 1Ju6n to five and did not differ between isolated (3.67 ± 1.37) and continuous sites (3.76 ± — 0d0 c (OdNN 1.75 1.65 0.72). Female fecundity of Gray Catbirds in ' ri isolated (1.33 ± 1.75) and continuous (2.54 ± 1.65) sites did not differ {t = 0.23, df = 29, - P = 0.814; t = 1.84, df = 32, P = 0.075; '3o^£,5 1.33 2.54 = Table 1). There was no difference in nest suc- li! cess between habitats for Gray Catbird (Table 1). 1.37 0.72 Yellow-breasted Chat nest success rates dif- fered between years and between continuous and isolated sites. In 2002, nest success was iTi higher in continuous habitats than isolated d din dTf (dn 3.67 3.76 habitats while in 2003 there was no difference bhiegthweereninhiasboiltaattesd (siTtaebslein12)0.0N3estthasnuicnceisssolawtaesd Tt O (N 00 0 0 sites in 2002 (Table 1). Mean Yellow-breasted Chat territory size in 2003 was 0.25 ± 0.24 ha {n = 20). No sig- tniwfeiecannthadbiiftfaetretnycpeein(0t.e2r2rit±ory0.s2i8zefeoxrisitseodlabtee-d ^Ov in do 0.916 0.914 sites, 0.29 ± 0.16 for continuous sites; t = -0.56, df = 18, P = 0.584). Territory size O' 00 —sC ^ 6 28 (N was not measured in 2002. ^ ^ DISCUSSION 2003 2003 04 (N (N 0^ We found no evidence that fragmentation or isolation ofriparian habitats in a western land- scape reduced reproductive success or in- Catbird creased brood parasitism of Yellow-breasted Continuous Isolated Chats or Gray Catbirds. Percent parasitism, Gray clutch size, and fecundity did not differ be- SHORT COMMUNICATIONS 497 tween isolated and continuous habitat patches birds can successfully reproduce in a highly for either species. However, delayed incuba- fragmented environment. Settlement patterns tion at isolated sites by both species suggests in this study suggest that, although continuous continuous sites may be preferred habitat, fill- and isolated sites may be comparable in qual- ing up sooner and causing late arrivals to ity, continuous tracts may be preferred breed- breed elsewhere (Fretwell and Lucas 1970). ing locations and have the capability to sup- Differences between years may reflect local port higher breeding densities. Alternatively, environmental fluctuations. More breeding isolated patches may lag behind continuous pairs were detected in 2003 but decreased nest patches in resource availability for reproduc- success in 2002 may be indicative of overall tion. It is also possible that continuous patches decreased habitat quality in that year. may cater to the semi-colonial nature thought Both Gray Catbirds and Yellow-breasted to be a characteristic ofYellow-breastedChats Chats had similar patterns of productivity and (Griscom 1923). nest success between habitats in 2003. When The current level of riparian habitat frag- compared to other Yellow-breasted Chat pop- mentation does not appear to be a limiting fac- ulations breeding at the core of their range tor for either Yellow-breasted Chat or Gray (Thompson and Nolan 1973, Schadd and Catbird populations in the Okanagan Valley, Ritchison 1998, Rickets and Ritchison 2000), British Columbia. Parameters for Yellow- the Okanagan population had similar levels of breasted Chat breeding at the northwest extent productivity. of their range are comparable to more south- Yellow-breasted Chat territory size (0.25 erly populations, suggesting that conditions at ha) in the Okanagan Valley was smaller than the edge of the species’ range do not reduce reported for mid and high-density southern productivity. The smaller territory size ofYel- Yellow-breasted Chat populations (0.5-1.0 ha, low-breasted Chats in the Okanagan Valley Dennis 1958; 1.24 ha, Thompson and Nolan may suggest a higher concentration of breed- 1973). Clutch size in the Okanagan (3.5 eggs/ ing resources. nest) was the same as that of more southerly ACKNOWLEDGMENTS populations (3.5 eggs/nest; Thompson and Nolan 1973). Overall fecundity (2.86 young/ We thank Environment Canada and WWF Endan- successful nest) was within the range of that gered Species Recovery Eund forfunding support. We reported for southeastern populations (3.25 also thank those who offered research expertise and young/successful nest, Schadd and Ritchison assistance in the field forthis project. In particular, we a1n9d98;non2-.p3a-r2a.s9itiyzoeudngn/essutcs,cesWshfiutlehpeaardasiettizaeld. tDKh.iabnbJ.kinEalmhleetrlhype,eddeRcdooinlclaetHcaetldlt,fhieeDsleadvdaisadstias.NtiaWenl.tds,;E.Ma.EnadsAt.RoenBn,eezeKn.MecrL-,. 2000). The percentage of nests that were suc- DeGroot, J. M. N. Smith, and Barry Lancasterprovid- cessful (57%) was midrange when compared ed editorial help. We thank Audra Moir and the Oso- to other studies (22.5%, Thompson and Nolan yoos Indian Band for providing access to their land, 1973; 84.2%, Schadd and Ritchison 1998; and all members of the community who provided in- 45%, Rickets and Ritchison 2000). The con- formationoncurrentandhistoricYellow-breastedChat sistency of fecundity values between the saingdhtRi.ngsA,.iCnanpnairtnigcsu.laWreStaelvseoOtghlaen,kLtahurreeieanRoocnkywmeoluls, Okanagan Valley and southern populations in- reviewers for their suggestions. dicates conditions at the fringe of this species range are not negatively affecting productivi- LITERATURE CITED ty* Cannings, R. A., R. J. Cannings, andS. G. Cannings. Despite the narrow range in latitude for all 1987. Birds of the Okanagan Valley, British Co- sites monitored, later nest initiation and fledg- lumbia. Royal British Columbia Museum, Victo- ing dates were found in isolated riparian ria, Canada. patches. Consequently, we expected birds CiMPRiCH, D. A. AND E R. Moore. 1995. Gray Catbird breeding in this habitat to have reduced nest (Dumetella carolinensis). The birds of North success or fecundity. The absence of a signif- America. Number 167. Croonquist, M. j. and R. R Brooks. 1993. Effects of icant difference in fecundity between isolated habitat disturbances on bird communities inripar- riparian patches and large tracts of habitat in- ian corridors. Journal of Soil and Water Conser- dicates Yellow-breasted Chats and Gray Cat- vation 48:65-70. 498 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 119, No. 3, September2007 Dennis, J. V. 1958. Some aspects ofthebreedingecol- R. Whitehead, and J. Faaborg. 1995. Regional ogy of the Yellow-breasted Chat (Icteria virens). forest fragmentation and the nesting success of Bird Banding 29:169-183. migratory birds. Science 267:1987-1990. Donovan, T. M., F. R. Thompson III, J. Faaborg, and Rotella, j. j., M. L. Taper, and A. J. Hansen. 2000. J. R. Probst. 1995. Reproductive success of mi- Correcting nesting-success estimates for observer gratory birds in habitat sources and sinks. Con- effects: maximum-likelihood estimates of daily servation Biology 9:1380-1395. survival rateswithreducedbias. Auk 117:92-109. Eckerle, K. P. and C. F. Thompson. 2001. Yellow- Sanders, D. H. 1995. Statistics: firstcourse. FifthEdi- breasted Chat {Icteria virens). The birds ofNorth tion. McGraw-Hill Inc, New York, USA. America. Number 575. SCHADD, C. A. AND G. RiTCHisoN. 1998. Provisioning Fretwell, S. D. and H. L. Lucas, Jr. 1970. On ter- of nestlings by male and female Yellow-breasted ritorial behaviorandotherfactorsinfluencinghab- Chats. Wilson Bulletin 110:398-402. itat distribution in birds. I. Theoretical develop- Sherry, T. W. and R. T. Holmes. 1992. Population ment. Acta Biotheoretica 19:16-36. fluctuations in a long-distance migrant: demo- Griscom, L. 1923. Birds ofthe New York City region. graphic evidence for the importance of breeding Handbook Series 9. American MuseumofNatural season events in the American Redstart. Pages History, New York, USA. 431-442 in Ecology and conservation ofneotrop- Hejl, S. j. 1992. The importance oflandscape patterns ical migrant landbirds (J. M. Hagan III and D. S. to bird diversity: a perspective from the northern Johnston, Editors). Smithsonian Institution Press, Rocky Mountains. NorthwestEnvironmentalJour- Washington D.C., USA. nal 8:119-137. Sherry, T. W. and R. T. Holmes. 1993. Are popula- Hejl, S. J. 1994. Human-induced changes in bird pop- tions ofneotropical migrant birds limited in sum- ulations in coniferous forests in western North merorwinter? Implicationsformanagement. Pag- Americaduringthepast 100years. Studiesin Avi- es47-57 in Status and managementofneotropical an Biology 15:232-246. Hensler, G. L. andj. D. Nichols. 1981.The Mayfield mEdiigtroartso)r.y Gbeirndesra(lD.TMe.chFniinccahl aRnedpoP.rtW.RMSt-e2n2ge9l., method for estimating nest success: a model, es- USDA, Forest Service, Fort Collins, Colorado, timators and simulation results. Wilson Bulletin USA. 93:42-53. MartCBiHnBA,ICRTH.DKEA.,p,rCot.P.ocRo.All.PlaeMinnoe,n,taCa.nnadJ.CCWooo.npewJraeaynt,kiivnWes..WMi.l1d9lH9io7f-.e TSteawtkSpsorbfoutgr.rya,2m0J0.m0a.Jn.,uSaSTl.A.JT.TIuHSlesTjalI,,COAaknlfdoarhTo.WmiEan.,dMoUawrSstAi.cno.mp1u9t9e8.r Research Unit, University of Montana, Missoula, Breeding productivity does not decline with in- USA. creasing fragmentation in a western landscape. Ohmart, R. D. 1994. The effects of human-induced Ecology 79:2890-2907. changes on the avifauna of western riparian hab- Thompson, C. F. and V. Nolan. 1973. Population bi- itats. Studies in Avian Biology 15:273-285. ology of the Yellow-breasted Chat {Icteria virens Rickets, M. S. and G. Ritchison. 2000. Nesting suc- /.) in southern Indiana. Ecological Monographs cess ofYellow-breasted Chats: effects ofnest site 43:145-171. and territory vegetation structure. Wilson Bulletin Whitehead, M. A., S. H. Schweitzer, and W. Post. 12:510-516. 2000. Impact ofbrood parasitism on nest survival 1 Robinson, S. K., J. P. Hoover, and J. R. Herkert. parameters and seasonal fecundity ofsix songbird 2000. Cowbird parasitism in a fragmented land- species in southeastern old-field habitat. Condor scape: effect of tract size, habitat and abundance 102:946-950. ofcowbirds and hosts. Pages 280-297 in Ecology WiLCOVE, D. S. AND S. K. Robinson. 1990. Theimpact and management of cowbirds and their hosts (J. of forest fragmentation on bird communities in N. M. Smith, T. L. Cook, S. I. Rothstein, S. K. eastern North America. Pages 319-331 in Bio- Robinson, and S. G. Sealy, Editors). Universityof geography andecologyofforestbirdcommunities Texas Press, Austin, USA. (A. Keast, Editor). SPD Academic Publishing, Robinson, S. K., F. Thompson III, T. M. Donovan, D. The Hague, Netherlands.