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THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 124. No. 2. June 2012 
hours patrolling their home range, stopping on 
successive perches and watching the surrounding 
areas, and flying out to inspect distant sparrows or 
meeting females. Males apparently practice a 
form °l scramble-competition polygyny, hut the 
implications of this behavior need to be further 
explored. Only one other promiscuous passerine, 
the Aquatic Warbler (Acrocephulus paludicola) 
(Dyrcz 1989: Birkhead 1993; Schulze-Hagen 
et al. 1995, 1999), is also known to perform 
scramble-competition polygyny, but differs in 
some details from the behavior we report for 
Saltmarsh Sparrows. One difference is the ab¬ 
sence of meet-and-pounce behavior in Aquatic 
Warblers. A key element in the Saltmarsh 
Sparrow sexual system is that most males behave 
as if they do not know the locations of most nests. 
Two forms of evidence support this conclusion’ 
the rare appearance of males at active nests of 
females, and the frequent practice of meet-and- 
pounce interactions in all stages of nesting. The 
alter behavior suggests that males often do not 
know the fertility status of females in their home 
ranges Females are quite stealthy around their 
nests during early stages of the nest cycle and 
chase males that approach the nest site. Male 
patrolling and meet-and-pounce interactions in 
conjunct,on with female sexual solicitation, must 
be understood in the context of individual marine 
ta ct,cs (Trivers 1972, Johnson and Burley 1997) 
sexual* h N* ,ta , m * cs °f Mating.- Male and female 
sexual behavior ,n Saltmarsh Sparrows may 
involve aspects of male coercion (forced mating? 
female control, and female choice. We found no 
common!? 0 ' d ° minance P^Vgyny, which 
commonly occurs m other promiscuous avian 
i 
iwcvjss Sr and ° ring i977 - Bradbu ^ 
species. Overall the U " expressed 1,1 the 
ized by apparent n l" 8 syste,n is character- 
females^ nS? foLT m,SCU - y by malcs 
female control of coerced bch ° vior ’ and 
oerced mounting attempts. We 
found no evidence that male Saltmarsh Spanovw 
followed and guarded females in New York, nor 
did Shriver et al. (2007) in Maine. Consequently, 
we must examine the possibility of forced mating 
behavior by males and female control of this 
behavior, and consider whether female choice 
could have a role in the mating system exhibited 
by Saltmarsh Sparrows. 
It males do not know the locations of nests of 
most females, or do not check them regularly, as 
we believe, some males may pursue a mixed 
sexual strategy of accepting solicited matings 
initiated by females and practicing forced mating 1 
with females when they find them at other times 
(McKinney et al. 1984). We do not know how 
widespread this second behavior may be among 
males, but we witnessed it daily during any 
extended period of observations in two marshes. 
Our interpretation of mounting behavior as 
forced mating is provisional because we provide 
no evidence that insemination actually occurs 
without female cooperation. However, based on 
behavior and morphology, we argue that such 
matings are possible, and perhaps likely. The 
aggressive tactics (pecking, wing-beating, grap¬ 
pling, and tumbling) that occurred between a male 
and female during pounces in Saltmarsh Sparrows 
were similar to actions used by males fighting 
over territories (Andrew 1956, Post and Greenlaw 
1975). The coercion tactics of males resemble 
those ol socially monogamous species that also 
practice forced mating; it occurs as well -n 
extrapair matings (McKinney et al. 1984. Birk- 
head ct al. 1985. Edinger 1988. Burley et al. 
1994). Phis behavior is well-known among 
dabbling ducks (McKinney et aJ 1983, Sorenson 
1994). 
Female resistance has been viewed as evidence 
ol forced mating (Birkhead et al. 1989 ), yet recent 
studies indicate that females initiate and control 
extrapair matings (Wagner 1991. Stutchburv and 
Neudorf 1998). The ability of female Saltmarsh 
Sparrows to either discourage some males I rum 
attempting to mount, or to thwart their mounting 
efforts, are elements of female control, but not of 
female choice. Some males, perhaps those more 
vigorous and persistent, are able to overcome the 
resistance of females. Some observers (e.g- 
ohwer 1978) are convinced that successful 
copulations in birds require the cooperation of the 
female but Morris (1957) reported that in Bronze 
Mannikins (Lonchura cucullata ), cloacal contact 
> males when females were uncooperative 
