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Territory Establishment, Size, and Tenacity by Male Red-Winged Blackbirds

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Territory Establishment, Size, and Tenacity by Male Red-Winged Blackbirds TERRITORY ESTABLISHMENT, SIZE, AND TENACITY BY MALE RED-WINGED BLACKBIRDS JAROSLAVPICMAN Departmentof Biology,University of Ottawa,30 SomersetE., Ottawa,Ontario KIN 6N5, Canada ABSTRACT.--Mostmale Red-wingedBlackbirds (Agelaius phoeniceus) established their first territoriesby replacingterritory holdersthat disappeared.These territories were significantly larger than those establishedeither through subdivisionof the vacatedterritory between two or three replacementmales, or through insertion of a new territory through displacement of one or more resident males. Almost all returning male Red-wings settled on their former territoriesin consecutiveyears. Their territoriesremained similar in sizeand locationbetween years. Strong territory tenacity apparently is favored by lower costs of territory defense becauseof reduced aggressionbetween familiar neighborsand site dominanceeffects. The lower costsof territory defensepresumably also allowed the site-tenaciousmales to establish larger territories in consecutive years. Strong territory tenacity by returning males could explain much of the settlementpattern of new males.As a result, the spatialarrangement of male territories in the study marsh (i.e. the location of territorial boundaries)was similar in consecutiveyears. Traditionalism in the male territorial systemshould influence Red-wing mating patterns.Received 27 January1986, accepted 5 January1987. MANY birds return to the same territories in between years. This, along with strong female consecutiveyears and frequently breed with site tenacity, could lead to a high degree of previousmates (e.g. Richdale1957; Coulson and "traditionalism" in both the distribution and White 1958, 1960; Krebs 1971; Richdale and number of male territories, and the distribution Warham 1973; Picman 1981; B•dard and La- of females in a given marsh. Pointe 1984; Cuthbert 1985; Gratto et al. 1985). Territory tenacityshould also be favored be- Territory tenacity and mate fidelity may in- causeof strongintrasexual competition for lim- crease the fitness of returning individuals ited breeding space(Rohwer 1982). This is be- through acceleratedpair formation (Lehrman causefamiliarity reducesthe level of aggression and Wortis 1967, Morris and Ericson 1971, Er- between neighbors(Yasukawa et al. 1982) and, icson 1973), increasedforaging efficiency,and thus, presumably lowers the costsof territory improved escape from predators (e.g. Hinde establishmentby returning males. Strong site 1956, Tinbergen 1957). Hence, site tenacity and tenacity by returning males could, in turn, de- mate fidelity play an important role in arian termine the location and size of territories es- reproductive strategies. tablished by new males. In addition, territory Territorytenacity could affect significantly the tenacityby returning malescould influence the mating pattern of polygynous speciessuch as size of their territories. With respectto benefits the Red-winged Blackbird (Agelaiusphoeniceus; associatedwith territory tenacity, the less te- Searcy 1979, Picman 1981). Female Red-wings naciousindividuals should experiencehigher return to their previous territories in consecu- rates of territorial interactions, and these could tive years,regardless of whether their previous negativelyinfluence the sizeof their territories. mate returned (Picman 1981). As a result, new The lower costsof territorydefense derived from malesmay "inherit" the haremsof the previous the high degreeof territory tenacityin return- territory holders(Picman 1981).Previous stud- ing males, however, should allow an increase ies on Red-wings suggestthat males also gen- in territory size in consecutiveyears if neigh- erally return to their territories in subsequent boring territoriesare vacated.Because breeding years(Nero 1956,Case and Hewitt 1963, Searcy space is often limited in marsh-nesting Red- 1979). Becauseoverwinter return rates of male wings (e.g. Orians 1961, Hurly and Robertson Red-wingsare relatively high (over 50%;Fank- 1985),acquiring a larger territory may increase hauser 1967, Picman unpubl. data), the distri- the probabilityof acquiringa largerharem (Pic- bution of Red-wing territoriesshould be similar man 1980b). 405 The Auk 104: 405-412. July 1987 406 JAROSLAVPICMAN [Auk, Vol. 104 Returningto the sameterritory shouldbe dis- tion of new territories, and the degree of territory advantageousfor malesthat experiencedlow tenacity by returning males, were examined in two reproductive success.To evaluate the role of ways. I measured the absolute overlap (m2) between previousbreeding experience,we need to de- the territoriesof a given male in successiveyears and the year-to-year change in size of the territories of termine if territory size, harem size, and fledg- returning males. To obtain an index of change in ing successof malesaffect territorial shifts be- territory location, I divided the overlapping area of tween years.This informationshould indicate the new and previous territories by the size of the the relative importance of intrasexual compe- new territory. This index estimatedthe proportion of tition in determining the pattern of territory a new territory locatedwithin the previous territory establishmentby male Red-wings.Thus, if low of a given male, and ranged from 0 for a nonover- reproductivesuccess did not influencethe de- lapping territory to 1 for a new territory locatedwith- greeof territoryshifts, competition among males in the previous territory. An index of changein ter- for breedingspace presumably played the major ritory size was estimatedas follows:territory size in role in favoring the high degree of territory year 2 - territory size in year 1 + 6,500 (this constant was added to transform negative into positive values, tenacityby males. and correspondsto the highest negative value of ter- I report the resultsof a 5-yr study on male ritory changedue to territory sizereduction from year Red-wingedBlackbird territoriality. My objec- I to 2). An Apple GraphicsTablet was usedto analyze tives were to examine the pattern of territory spatialdata on territory size and the amount of over- establishmentby new males;the degreeof ter- lap betweennew and previousterritories of returning ritory tenacity in males that returned to the and replacementmales. Methods of studying repro- marsh in consecutiveyears; the effect of terri- ductive and mating successof males are described tory size,mating success,and reproductivesuc- elsewhere (Picman 1980a, 1981). The critical level for cesson shifts of male territories between years; rejectinga null hypothesiswas P < 0.05. One-tailed testswere usedwhenever directional predictions could and the effect of territory tenacity on the size be made. of male territories. RESULTS METHODS Patternof territoryestablishment by newmales.-- The study was conductedbetween 1976 and 1980 in an extensive, brackish-water marsh at George C. A total of 95 territories defended by 61 males Reifel MigratoryBird Sanctuary,on WesthamIsland, (3 present for 4 yr and 1 throughout the 5-yr Delta, British Columbia. The higher grounds of this study period) was observed.In 1977-1980,41 marsh are coveredby a relatively homogeneouscat- new males established first territories in the tail (Typhalatifolia) vegetation. Sedges (Carex sp.) study area (Table 1). Most new males(66%) re- and bulrushes (Scirpussp.) dominate the lower areas placedterritory holders that disappearedfrom near the seashore.Only two passerinespecies, the the studymarsh (n = 26) or that abandonedtheir Red-wingedBlackbird and the Marsh Wren (Cistotho- old territory and establisheda new territory ruspalustris), commonly breed in this marsh. elsewhere in the marsh (n = 1). Approximately Early in April 1976 I establisheda grid systemby 78% of territories establishedby replacement dividing the study area (between 15 and 25 ha in differentyears) into 20 x 20-m quadrats.This made overlappedby at least 60% with the original it possibleto map the locationof singingperches of territories (Fig. 1). Somereplacement males ex- male Red-wings and to quantify their interactions panded the territoriesof their predecessorsby with intruders. Eachyear unbanded maleswere cap- including areas located outside the breeding tured using a decoy trap (Picman 1979) and color- areain a previousyear. This reduced the degree banded. Territories of individual males were deter- of overlap between the original and new ter- mined eachyear in April and early May by recording ritories. Despitethese changes, the original and the locationof singing perches,flight paths,and in- new territories were similar in size (Table 2). teractionswith conspecificintruders on mapsof the As a result, the size of new territories estab- studyarea. Territory sizes were calculatedby the min- lished by replacementmales correlated signif- imum convexpolygon method. Territory establishmentby new males (i.e. males icantly with the original territory size (r = 0.60, df = 17, P < 0.001). that appearedin the marsh for the first time) was studied by examining their recruitment into the Eight adult malesestablished their first ter- breeding population. The effect of replacementof ritoriesby dividing three vacatedterritories be- malesthat disappearedby singlemales on the loca- tween 2 (n = 1) or 3 (n = 2) replacement indi- July 1987] Territorialityin Male Red-wingedBlackbirds 407 TABLE1. Recruitmentof male Red-winged Blackbirdsinto the breeding population, 1977-1980. Number of new males settling in: 1977- Mode of territory establishment 1977 1978 1979 1980 1980 Replacementof a male
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