Impact of Interspecific Aggression and Herbivory by Mute Swans on Native
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744 ShortCommunications andCommentaries [Auk,Vol. 111 ervation of avian blood and tissue samplesfor predominantlymonogamous bird: Genetic evi- DNA analysis.Can. J. Zool. 69:82-90. dence. Anim. Behav. 35:877-886. SHERMAN,P. W. ANDM. L. MORTON.1988. Extra-pair WESTNEAT,D.F. 1990. Geneticparentage in the In- fertilizationsin mountain White-crownedSpar- digo Bunting: A study using DNA fingerprint- rows. Behav. Ecol. Sociobiol. 22:413-420 ing. Behav. Ecol. Sociobiol. 27:67-76. SH•N, H.-S., T. A. BARGIELLO,B. T. C•a•ac, F. R. JACKSON, WESTNEAT,D. F., P. W. SH•RMAN, AND M. L. MORTON. AND M. W. YOUNG. 1985. An unusual coding 1990. The ecology and evolution of extra-pair sequencefrom a Drosophilaclock gene is con- copulations in birds. Curt. Ornithol. 7:331-369. served in vertebrates. Nature 317:445-448. WETTON, J. H., R. E. CARTER,D. T. PARICIN,AND D. SMITH, H. G., R. MONTGOMERIE,T. POLDMAA, B. N. WALTERS.1987. Demographicstudy of a wild WHITE,AND P. T. BOAG. 1991. DNA fingerprint- HouseSparrow population by DNA fingerprint- ing reveals relation between tail ornaments and ing. Nature 327:147-149. cuckoldry in Barn Swallows, Hirundorustica. Be- hav. Ecol. 2:90-98. Received30 June1993, accepted 17 November1993. WESTNEAT,D. F. 1987. Extra-pair fertilizations in a The Auk 111(3):744-748, 1994 Impact of InterspecificAggression and Herbivory by Mute Swanson Native Waterfowl and Aquatic Vegetation in New England MICHAEL R. CONOVER • AND GARY S. KANIA 2 •BerrymanInstitute and Department of Fisheriesand Wildlife, Utah State University, Logan,Utah 84322, USA; and 2Depart:aentof WildlifeManagement, National Rifle Association, 11250 Waples Mills Rd., Fairfax,Virginia 22030, USA One of the greatestthreats to American avifauna is whether theseconcerns are real or whether swanpop- the establishmentof free-rangingexotic avian pop- ulationsshould be controlled.At present,Mute Swans ulations (Temple 1992). Despite evidenceof substan- are protectedin some statesand unprotectedin oth- tial harm causedby exotic birds, most exotic species ers;in still others,government employees attempt to are so poorly studiedthat many of their alleged en- control swan populationsby shaking eggs and re- vironmental impacts remain largely undocumented moving adults (Allin et al. 1987). In this study, we (Temple 1992). One such exotic speciesis the Mute examined interspecificaggression by free-ranging Swan (Cygnusolor), a Eurasianspecies that has been adult Mute Swanswith breeding territories in fresh- introduced severaltimes into North America, begin- water ponds and the impact of their herbivory on ning in the late 1800s(Allin et al. 1987).By the 1970s, aquatic vegetation. free-ranging populations existed in Michigan, Min- Methods.--Territorial pairs of free-ranging Mute nesota,Wisconsin, Wyoming, British Columbia, On- Swanswere observedat 15 freshwaterponds (2 to 30 tario, and in Atlantic coastalstates from Maryland to ha) in New Haven County,Connecticut from 1982to Massachusetts(Allin 1981). In the Atlantic Flyway, 1987.These ponds represented all freshwatersites in free-rangingpopulations increased from 200 birds in the study area known to have nestingpairs of free- 1955 to 5,300 in 1987 (Allin et al. 1987). ranging Mute Swans in 1982. Observationswere lim- Some biologistsare concernedthat the increasing ited to freshwater sites because these were the main populationof free-rangingMute Swansmay havean areas where Mute Swans came into contact with na- adverse impact on native waterfowl, owing to the tive waterfowl. swan'saggressive nature (Reese1975, Williams 1989). Data on the impactof interspecificaggression were Swanssometimes attack other waterfowl, causingin- collectedon both membersof a swan pair simulta- jury or death (Stone and Marsters1970, Willey and neously;data for males and femaleswere analyzed Halla 1972,Allin et al. 1987).Furthermore, aggressive separately.Sex of pair memberswas determinedby swansmay displaceother waterfowl (Willey and Hal- the larger body size and larger fleshy knob on the la 1972). An additional concern is that the foraging forehead of males (Bellrose 1980). Pairs were observed behavior of swansmay adverselyaffect aquaticplant year-round for 30-rain periods, randomly selected biomass,reducing the food available for other wa- amongdaylight hours.Observations were madefrom terfowl. Currently, data are insufficient to judge shore,usually from a car to minimizedisturbance of July 1994] ShortCommunications and Commentaries 745 the birds.During an observationperiod, we notedall T^BLE1. Number of aggressiveinteractions per hour aggressivebehaviors by swans,the other speciesin- (œñ St) initiatedby free-rangingmale (n = 15)and volved, and the outcomes.Agonistic behaviors by female (n = 15) Mute Swans against swans and swansincluded threats (raising wings above back), other waterfowl at 15 freshwater sites in Connect- icut. chases,and physical attacks. All sites were searched weekly from a boat or shorefor nestingwaterfowl. Samplesizes for all statisticaltests were the number Speciesbeing attacked Male Female pa of subjectsunder observation.Paired t-testswere used to ascertainif there were sexualdifferences in ag- Mute Swan 0.62 ñ 0.21 0.31 ñ 0.11 0.03 gressivebehavior. To test whether seasonalvariation Mallard 0.10 ñ 0.03 0.07 ñ 0.01 0.07 occurredin swanaggressiveness, data on eachswan's American Black Duck 0.03 ñ 0.01 0.01 ñ 0.001 0.10 behaviorduring March to May, Juneto August,and Canada Goose 0.38 ñ 0.29 0.01 ñ 0.01 0.11 Septemberto Februarywere analyzedusing a two- Human 0.25 ñ 0.07 0.16 ñ 0.06 0.08 wayANOVA (seasonsvs. subjects). Linear regressions Total interspecific were usedto compareinterspecific aggression rates aggression 0.80 ñ 0.30 0.31 ñ 0.09 0.04 of individual swansto their intraspecificaggression Paired one-tailed t-test comparingmales and female swans. rates. To determinewhether waterfowlwere avoiding areas near swans, we prepared a map of each site site was consideredas being composedof five sec- noting water depths.At the beginning of eachob- tions. Eachtime a site was sampled,all of the above- servation period, we noted a swan's location and ran- groundvegetation was harvestedwithin a 0.33 x 0.33 domlyselected from the mapanother point of similar m sampling frame placed within each section. Sub- depth.At 5-rainintervals, we thenvisually estimated sequent sampling was taken elsewhere in each sec- the distancefrom the nearestindividual of everywa- tion so that no area was harvested more than once. terfowl speciespresent to both the swan and the ran- Samplingwas conductedin late May or early June domlyselected point. For eachindividual swan,these when the exclosureswere established(year 1, spring), datafrom all observationperiods were then combined three months later (year 1, fall), and 15 months later to yield a single mean distancevalue between it and (year 2, fall). Vegetationwas segregatedby species, each waterfowl species.A similar mean value was driedfor morethan 24 h at 110øC,and weighed.Paired obtainedfor the pairedrandom points. Hence, sample t-testswere usedto comparethe biomassof grazed sizes consisted of the number of individual swans and ungrazedsites. One grazedand one ungrazed under study. A paired t-test was then used to test samplewere obtainedfrom eachpond with eachsam- whethermean distances between a waterfowlspecies ple consistingof 10 sampleframes collected from the and a swan differed from the distance between that samepond (five sampleframes per exclosurex two speciesand a random point. exclosuresper pond). Hence, for thesestatistical tests, The experimenton the impactof swanherbivory the samplesize equalled the number of ponds under was conductedat 12 of the freshwaterponds used in study. the other part of this study.Exclosures were erected Results.--During405.0 h of observationon 15 males in late May or early June at two ponds in 1983, six and 398.5h on 15females, we observed870 aggressive ponds in 1984, two in 1985, one in 1986, and one in interactionsby free-ranging Mute Swans;410 were 1987.At each pond, we locatedtwo sites (3 x 3 m directedagainst other swansand 460 againstother each)between 20 and 50 m apart that were similar in species.Swans were aggressivetowards humans 174 water depth (between0.5 and 1.0 m) and appeared times (usuallyinvolving a threat displaydirected at to have similar vegetation. One of these sites was peopletrying to feed them). They were alsoaggres- randomly selectedto serve as an open site where sive towardsGreat Blue Herons (Ardeaherodias) three swanscould graze (hereafterreferred to as grazed times, gulls (Larusspp.) three times, scaups(Aythya site)and markedwith a woodenpost. The other site spp.) twice, Gadwalls (Anasstrepera) once, domestic served as the ungrazed site and a 3 x 3 m swan ducksseven times, and domesticgeese 21 times.On exclosure was centered on it. The exclosure was built all other occasions,interspecific aggressive behavior by attaching a 0.3 to 0.5 m wide chicken-wire fence was directedat Mallards (Anasplatyrhynchos), Amer- to eight posts.The fence was built so that its bottom icanBlack Ducks (A. rubripes),and CanadaGeese (Branta edge was level with the water surfaceso that it would canadensis;Table 1). excludeswans but not other aquaticherbivores such There were no significantdifferences among years as fish, turtles,diving ducks,and muskrats.Another in the aggressivebehavior of swans.Hence, data were pair of grazedand exclosuresites was set up on the summarizedfor eachindividual subjectacross years oppositeside of the pond. Data from thesetwo pairs