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Why Are Nesting Marsh Wrens and Yellow-Headed Blackbirds Spatially Segregated?

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Why Are Nesting Marsh Wrens and Yellow-Headed Blackbirds Spatially Segregated? WHY ARE NESTING MARSH WRENS AND YELLOW-HEADED BLACKBIRDS SPATIALLY SEGREGATED? MARTY L. LEONARDAND JAROSLAVPICMAN Departmentof Biology,University of Ottawa,Ottawa, Ontario KIN 6N5,Canada ABSTR^CT.--Theactivity centers of MarshWrens (Cistothorus palustris) and Yellow-headed Blackbirds(Xanthocephalus xanthocephalus) are spatiallysegregated. This segregationmay oc- curbecause (1) one speciesexcludes the otheror (2) the two speciesprefer different habitats. Thesehypotheses were testedby documentingchanges in the sizeand locationof Marsh Wren territoriesthroughout the breedingseason, and by the removalof conestingYellow- headedBlackbirds. The expansionof Marsh Wren territoriesinto blackbirdbreeding areas after boththe naturaldeparture and the removalof blackbirdsbest supports the explanation basedon the activeexclusion of Marsh Wrensby Yellow-headedBlackbirds. Received 8 Feb- ruary1985, accepted 21 August1985. ECONOMICdefensibility of resources(Brown Red-winged Blackbirds(Nero 1956, Burt 1970, 1964)necessary for reproductionshould favor Runyan 1979, Picman 1980). Yellow-headed the evolution of breeding territories in birds. Blackbirds also nest in marshes, and limited Resourcesare defended most often against con- evidencesuggests that yellow-headsmay force specificsbecause these individuals are gener- compressionof Marsh Wren territories (Orians ally more similar in their requirementsthan and Willson 1964, Verrter 1975). heterospecifics.However, territories also may Spatial segregation between Marsh Wrens be defended against individuals of closely re- and Yellow-headed Blackbirds can be ex- lated speciesand morphologically or ecologi- plained by the following hypotheses.(1) One callysimilar species (e.g. Cody 1969,Reed 1982). speciesexcludes the other from mutually pre- Of particular interest are incidencesof inter- ferred habitat, either directly through behav- specificaggression and territoriality between ioral interactions (interference competition) or markedlydissimilar species (Willson 1967;Ver- indirectly through resourcedepletion (exploi- ner 1975; Picman 1980, 1982). tation competition) (Ricklefs 1979). (2) Each Marsh Wrens (Cistothoruspalustris) puncture speciesprefers a different habitat. These hy- eggsand kill nestlingsof other marsh-nesting pothesesmake the following predictions.First, passerines(Orians and Willson 1964,Burt 1970, if one speciesexcludes the other, then the re- Picman 1977, Bump 1983). Aggressive interac- moval of the dominant speciesshould result in tions have been noted between Marsh Wrens the movement of the subordinate speciesinto and Swamp Sparrows (Melospiza georg•ana; that area. This expansionshould not occur in Willson 1967), Song Sparrows (M. melodia; areasinhabited only by the subordinatespecies. Willson 1967),Red-winged Blackbirds (Agelaius If, however,spatial segregation results from dif- phoeniceus;Orians and Willson 1964, Willson ferencesin habitatpreference, then both species 1967, Picman 1983), and Yellow-headed Black- should remain in the samearea throughout the birds (Xanthocephalusxanthocephalus; Orians and breeding season,regardless of the presenceor Willson 1964, Verner 1975, Bump 1983). Pre- absenceof the other species,and barring any vious evidencesuggests that egg pecking has major seasonalchanges in the habitat. Alter- evolvedprimarily asa meansof reducingcom- natively, if one speciesactively excludesthe petition and not of obtaining food (Picman other, then aggressiveinteractions should be 1984). Marsh Wren nest locationsand feeding observed between them. The absence of ag- areasare often spatiallysegregated from other gressiveinteractions would supportthe expla- co-occurringpasserines (Willson 1967, Verner nation based on indirect exclusion. 1975, Picman 1980). The purposeof our study was to determine The most intensive studies have concentrat- which hypothesisbest explains spatial segre- ed on interactions between Marsh Wrens and gation between Marsh Wrens and Yellow- 135 The Auk 103: 135-140. January 1986 136 LEONARDAND PICMAN [Auk, Vol. 103 TABLE1. Dates of sampling periods for 1983 and 1984 and the maximum number of Yellow-headed Blackbird nests active during those intervals at Site 2. No. (%) of active yellow- Year Period Date head nests 1983 1 22 May to 5 June 197(87%) 2 5-19 June 131 (58%) 3 19 June to 3 July 19 (8.4%) 4 3-15 July 5 (2.2%) 1984 1 24 May to 7 June 194 (88%) 2 7-21 June 180 (82%) 3 21 Juneto 5 July 14 (6.4%) 4 5-17 July 2 (0.9%) headed Blackbirds. Two types of data were collected to test the above predictions:(1) de- scriptive data on interactions between the speciesand on seasonalchanges of wren ter- ritories in one area with and another area with- out yellow-heads;and (2) experimentaldata on the responsesof Marsh Wrens to the removal of nesting yellow-heads. METHODS Studysites.--The descriptive study was conducted from 22 May to 16 July 1983and 24 May to 17 July 1984 in two marsh sites in Delta, Manitoba, Canada. Site 1 wasa relativelydry, homogeneouscattail (Ty- phaspp.) marsh, approximately 6 ha in area,that sup- Fig. 1. Map of Marsh Wren territories at Site 2 for ported 13 Marsh Wren territories in 1984. In addi- Period 2 when yellow-headswere present (dotted tion, 2 pairs of Common Yellowthroats (Geothlypis lines)and Period3 when yellow-headswere absent trichas)and 3 pairs of Red-wingedBlackbirds nested (solidlines), 1984.Shaded areas represent water. The along the drier edgesof this site. Site 2 was 4 ha in boundary of the yellow-head colony is represented areaand consistedof phragmites(Phragmites australis) by a dashedline. Resultswere similarfor 1983. and cattailsalong the edges,and bulrush(Scirpus sp.) and cattail patchessurrounding two central ponds. In 1983 this marsh contained 67 Yellow-headed Territory movements.--Maleyellow-heads in the Blackbird and 6 Marsh Wren territories; in 1984, there Delta area arrive approximatelythree weeks before were 63 yellow-headand 10 Marsh Wren territories. male Marsh Wrens (pets. obs.). Yellow-head males No other passetinesnested in this area. The vegeta- were capturedand markedindividually with colored tion at both studysites became denser as the season leg bands. Territory boundarieswere mapped by progressed(i.e. with the growth of new vegetation). plotting songperches and flight pathsof individual However, water depth and the compositionand dis- maleson a map of the studysite. Each male was ob- tribution of vegetation within each site remained servedfor a 30-min interval twice a week throughout constant.Both siteswere divided into a grid of 20 x the breedingseason. 20-m squares,marked by 2-m-high wooden stakes. Twenty-six of 29 male Marsh Wrens were trapped Water depth was measuredat each stakeusing a 100- and marked. Territory boundarieswere mapped on cm-high stick divided into 5-cm intervals. The ex- a weekly basisthroughout the study period by ob- perimentalstudy was conductedfrom 31 May to 11 serving the movementsof individual males during June 1984 at a third site in Delta. This site was ap- 1-h intervals in 1983 and 30-min intervals in 1984. proximately 1.5 ha in size and composedof a rela- Songperches, courtship nests (Verner 1963),and flight tively homogeneouscattail stand surrounding a small pathswere plotted on mapsof the studysites for each pond.The areacontained 21 yellow-headand 4 Marsh interval.Territory areas were calculatedby the min- Wren territories. imum convex polygon method for four 2-week in- January1986] SpatialSegregation Between Wrens and Blackbirds 137 T^BI•E2. Mean water depth (cm, + SD) measured in Marsh Wren and Yellow-headed Blackbird territories at Site 2 in 1983 and 1984 and mean water depth in Marsh Wren territories at Site I in 1984. Site 2 Site 1 1983 1984 1984 Marsh Wren 23.0 + 10.9 17.0 + 11.6 8.98 + 5.42 n = 28 n = 26 n = 76 Yellow-head 39.2 + 8.9 33.9 + 9.7 n = 35 n = 60 Statistical t = 6.31 t = 7.63 comparison P < 0.001 P < 0.001 tervals. These intervals represent two periods with in 1984). Marsh Wren and yellow-head terri- respectto yellow-heads(Table 1): when yellow-heads tories did not overlap except for one case in were presentin the study area (PeriodsI and 2) and 1983 and one in 1984. Yellow-heads at Site 2 when yellow-headswere essentiallyabsent (Periods defended territories surrounding the central 3 and 4). Although there were no yellow-headsat ponds, whereas Marsh Wrens establishedter- Site 1, territory sizes for the same intervals were re- corded as controls. ritories along the edges of the marsh (Fig. 1). Behavioral interactions.--Behavioral interactions be- The water was significantlyhigher in yellow- head territories than in Marsh Wren territories tween Marsh Wrensand yellow-headswere recorded during time-budgetstudies. In 1983 a maximum of 4 at Site 2 in both 1983 and 1984 (Table 2). Water territorial male wrens/day were observedfor I h each. depth measuredat Site 1 in 1984 was signifi- In 19848 males/daywere observedfor 30 min each. cantly lower than water depth measuredin the The observationperiod for both yearswas between Marsh Wren territories at Site 2 in 1984 (Table 0530 and 0930, correspondingto the birds' mostac- 2; t = -3.59, P < 0.001). tive period (pets.obs.). The duration,type, and num- Territorymovements.--There were no signifi- ber of separateinteractions were recorded for each cant differencesamong the four periods (ran- period. Any of the following activitieswas consid- domized block ANOVA; F = 1.23; df = 3, 26; ered to be an interaction:(1) a chaseinvolving in- dividualsof the two species,(2) a yellow-headflying P > 0.10) in the
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