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Copulation and Mate Guarding in the Northern Fulmar

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Copulation and Mate Guarding in the Northern Fulmar COPULATION AND MATE GUARDING IN THE NORTHERN FULMAR SCOTT A. HATCH Museumof VertebrateZoology, University of California,Berkeley, California 94720 USA and AlaskaFish and WildlifeResearch Center, U.S. Fish and WildlifeService, 1011 East Tudor Road, Anchorage,Alaska 99503 USA• ABSTRACT.--Istudied the timing and frequency of copulation in mated pairs and the occurrenceof extra-paircopulation (EPC) among Northern Fulmars (Fulmarus glacialis) for 2 yr. Copulationpeaked 24 days before laying, a few daysbefore females departed on a prelaying exodus of about 3 weeks. I estimated that females were inseminated at least 34 times each season.A total of 44 EPC attemptswas seen,9 (20%)of which apparentlyresulted in insem- ination.Five successful EPCs were solicitated by femalesvisiting neighboring males. Multiple copulationsduring a singlemounting were rare within pairsbut occurredin nearly half of the successfulEPCs. Both sexesvisited neighborsduring the prelayingperiod, and males employed a specialbehavioral display to gain acceptanceby unattended females.Males investedtime in nest-siteattendance during the prelaying period to guard their matesand pursueEPC. However, the occurrenceof EPC in fulmars waslargely a matter of female choice. Received29 September1986, accepted 16 February1987. THEoccurrence and significanceof extra-pair Bjorkland and Westman 1983; Buitron 1983; copulation (EPC) in monogamousbirds has Birkhead et al. 1985). generated much interest and discussion(Glad- I attemptedto documentthe occurrenceand stone 1979; Oring 1982; Ford 1983; McKinney behavioral contextof extra-pair copulationand et al. 1983, 1984). Becausethe males of monog- mate guarding in a colonial seabird,the North- amous species typically make a large invest- ern Fulmar (Fulmarus glacialis). Fulmars are ment in the careof eggsand young, the costof among the longest-lived birds known, and fi- being cuckolded is high, as are the benefits to delity to the same mate and nest site between the successfulcuckolder. Males are expectedto years is high (Macdonald 1977, Ollason and pursueopportunities for copulationoutside the Dunnet 1978,Hatch 1985). Only one egg is laid pair bond (Trivers 1972) and to reduce as far as per clutch, and re-laying in the same season possiblethe uncertainty of paternity for the after the loss of a clutch is unknown. The sexes young they help raise. The incidence of EPC share about equally in incubation and chick- may in general be higher in colonial species rearing duties (Hatch 1985). Thus, fulmars ex- than in solitary nesters,and the threat of cuck- hibit a highly conservativesocial system. Some oldry is postulatedto be one of the principal features of the breeding biology make this disadvantagesof colonial breeding (Alexander species(and perhaps other Procellariiformes) 1974,Hoogland and Sherman 1976). particularly interesting from the standpoint of Mate guarding may be defined asany behav- copulation behavior and potential sperm com- ior by a mated male whose principal function petition. At Pacificcolonies, fulmars arrive syn- is to reduce the likelihood of encounters be- chronouslysome 6-8 weeksbefore the first eggs tween his mate and other malesduring the time are laid. There is thus a considerableprelaying when fertilization of her eggsis possible.The period for social interaction. Foraging occurs importanceof mate guarding in the monoga- over large oceanareas in boutsof severaldays, mousmale's reproductive strategy has been rec- and it is difficult for males and females to co- ognizedin a variety of species(e.g. Beecher and ordinate perfectly their attendancepatterns at Beecher 1979; Birkhead 1979, 1982; Power and the nest site. Females are receptive to copula- Doner 1980; Power et al. 1981; Werschku11982a; tion over the whole prelaying period,and sperm are storedin specialglands in the utero-vaginal (UV) region of the oviduct (Hatch 1983).Sperm Present address. remain viable in the UV glands for several 450 The Auk 104: 450-461. July 1987 July 1987] Copulationand Mate Guardingin Fulmars 451 weeks, as frequently there is no contact be- at a nest site (the host). Visitors usually arrived on tween the male and female during the "pre- the wing, but closeneighbors also moved between laying exodus" (Warham 1964), a time of con- siteson the ground. tinuousforaging immediately before egg laying In May 1980I madenotes opportunistically on ex- (lasting up to 38 days in fulmars; Hatch 1985). tra-pair visits and completed21.3 h of dedicatedob- The combination of intermittent attendance servationson one plot of about 130 nest sites (plot Q). Observationson plot Q in 1980 encompassedthe patterns,sperm storage, and a prolonged recep- whole plot, without regard to particular focal pairs. tive period in the female renders the male ful- For 3-6 h on eachof 5 daysI watchedfor interactions mar particularly susceptibleto being cuckold- amongknown individualsand recordedon audiotape ed. the characteristicsof all visits detected. In addition, I studied the timing and frequencyof copu- whenever I detecteda visit involving known breed- lation in mated pairs, both in relation to cal- ing birds during my daily rounds of other plots, I endar date and relative to the laying dates of stopped to observethe encounter to its conclusion, individual females. Social relations outside the and recorded information on duration, behavior of pair bond, including EPC,were documentedin the sexes,and the visitor'sidentity. All visitsinvolv- a seriesof extendedwatches on a group of ing breedingbirds of oppositesex were placedin one of four categoriesfor the purposeof presentation.In known individuals during 2 yr. Finally, I ex- type ! a breeding male visited an unattendedbreed- amined patternsof nest-siteattendance during ing female,in type 2 a breedingmale visiteda breed- the prelayingperiod during 6 yr for evidence ing pair, in type 3 a breeding female visited a lone of mate guarding. breedingmale, and in type 4 a breedingfemale visited a breeding pair. A standard copulation count was conducted each METHODS afternoonor evening (3-26 May). All occurrencesof mountingin 1 h were noted, including both success- The study was conductedon the Semidi Islands, ful and unsuccessfulcopulations (see below), and the Alaska (56øN, 156øW),where an estimated440,000 ful- number of pairs on the plot was recordedat the be- marsbreed (Hatch and Hatch 1983).The mainstudy ginning and end of the hour. areaon ChowletIsland contained 6-8 plotsthat were In 1981 all behavioral observations were confined usedto monitor colonyattendance from 1976 to 1981. to plot Q. Two observersworked alone or simulta- The plotscomprised 500-700 nest sites. Monitoring neously,each observing a sampleof 24-31 nest sites began 4-8 weeks before egg laying and continued for 1-9 h daily from 17 April to 26 May, for a total throughthe late chick stage in mostyears. Daily counts of 176 h. The samesample of nest siteswas observed of single birds and pairs on the plots were made be- throughout the season,but not all sitesunder obser- tween 0900 and 1600. I also monitored several hundred vation at any one time were occupied.I sampled oc- sitesindividually, noting the attendanceof adultsand cupied sitesfor 4,173 site-hours,including 1,655site- the presenceof eggsor young eachday. Bothpro- hoursfor singlesand 2,518site-hours for pairs.Again, ceduresprovided information on nest-siteattendance the visit involving known breeding birds was the usedin the presentanalysis (see also Hatch 1985). focalunit of behavior,and the large majorityof oc- All birds usedin the study of socialinteractions currencesprobably was detected. Identity and breed- (1980-1981)were sexedby their position in copula- ing statuswere determinedas necessaryfor birds that tion, and their breedingstatus was determined by site interactedwith focal pairs. Information recorded on attachmentand egg laying. Fulmarsoccurred in a audiotapefor eachvisit includedtime, identity and wide range of color phases,and most birds also had generalbehavior of the participants,duration of the distinguishingblack marks on the culmen. Thus, a visit, occurrenceof EPC attemptsor other physical combinationof plumage differencesand bill mark- contact,and presenceor absencein the colonyof the ings provided a reliable systemfor individual iden- visitor's mate. All occurrencesof copulation in focal tification. Initially, I sketchedeach bird's bill mark- pairs during behavior watches were recorded, and ings on a templatedrawing of the head to provide a standard1-h copulationcounts were conducteddaily permanentrecord of individualidentity. After mem- in 1981. orizing the layout of nestsites and individual markers Successfulcopulations were easily distinguished (an easytask becausesite-holding fulmars were rel- from unsuccessfulattempts because the appositionof atively sedentaryon land), all observationsof behav- the birds' cloacaewas a laboredand conspicuouspro- ior were ascribedto individualsaccording to sexand cess,as was the final thrustingby the male (lasting site number. several seconds) once cloacal contact was achieved. The basic unit of behavior ! recorded was the visit, Copulation sequencesthat included cloacal contact defined as the directed approachof one bird (the and thrusting most likely resulted in insemination.I visitor)to within 0.5 m of anotherindividual or pair refer to thoseas "completedcopulations." I use the 452 SCOTTA. HATCH [Auk, Vol. 104 RESULTS 1500 Copulations(no,/pr,/h) • 0.23 0.33 0.42 Success rate - 0,63 0,69 0.64 1980 Sample size (pr,-h) - 594 438 195 Behaviorduring
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