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Pair Formation in Relation to Climate: Mallard, Eurasian Wigeon and Eurasian Teal Wintering in Iceland

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Pair Formation in Relation to Climate: Mallard, Eurasian Wigeon and Eurasian Teal Wintering in Iceland Pair formation and climate 55 Pair formation in relation to climate: Mallard, Eurasian Wigeon and Eurasian Teal wintering in Iceland. J. E. Jonsson' & A. Gardarsson Institute of Biology, University of Iceland, Grensasvegur 12, 108 Reykjavik, Iceland Email: arnthorldhi.is 'Present address: School of Forestry, Wildlife and Fisheries, Louisiana State University, Baton Rouge, LA 70803, U.S.A. Email: jjonss 10lsu.edu The pairing chronologies of Mallard Anas platyrhynchos, Eurasian Wigeon A. penelope and Eurasian Teal A. crecca wintering in Southwest Iceland were studied over the winters of 1997-98 and 1998-99. Mallard paired first of the three species, with over 60% of all females paired by November. Wigeon and Teal had started pairing in early October, and the proportion of paired females increased gradually through winter until spring. In Mallard, the percentage of paired females was 60-70% during midwinter, the last females became paired in March and April. All three species paired later in Iceland than reported in more southern European and North American areas. In all winter months, especially November, December and January, the proportion of paired Mallard females at each location showed a positive relationship with mean monthly temperature. Fewer data are available on other dabbling duck species, but their pairing chronology seems related to climatic conditions. Key Words: Anas platyrhynchos, Anas penelope, Anas crecca, Anatidae, pairing chronology, winter climate. ©W ildfowl & Wetlands Trust W ildfowl 12001] 52: 55-68 56 Pair formation and climate Mate defense is energetically costly, 1988). The sex ratio of northern hemi­ because courtship, breeding and sphere ducks is usually male-biased repelling rivals cost time and energy (Bellrose 1980; McKinney 1986), which (Afton & Sayler 1982; Wishart 1983). At means that some males will be left high latitudes, limited food availability unpaired every season, even though all and rough weather in winter may cause the females become paired. This leads starvation and poor physiological con­ to sexual selection among males, and dition (Hepp 1989). As well as being at females can probably afford to be risk from starvation, birds face the dan­ selective in their search for a mate ger of hypothermia in cold and windy (Rohwer & Anderson 1988). The conditions. At air temperatures below females choose males by age, plumage the thermoneutral zone, birds need to quality, body mass and the intensity of conserve energy by limiting energeti­ their courtship displays (Wishart 1983; cally costly behaviour, such as Bossema & Roemers 1985; Hepp 1986; courtship displays (Newton 1998). Holmberg ef al. 1989; Weidmann 1990; The wintering range of many north­ Heitmeyer 1995). ern dabbling ducks is between 30°N In contrast to the life-long pair and 60°N. Individuals of widely distrib­ bonds of swans and geese, ducks form uted species, such as Mallard Anas pair bonds each year (Williams & platyrhynchos, may face very variable McKinney 1998). Ducks form pairs on tem peratures during a year. the wintering grounds and a pair will Behavioural decisions of ducks winter­ remain together until the female starts ing at the edges of the range should to incubate. Then the male deserts the differ from those of individuals winter­ female. In the autumn, the sexes meet ing in central areas. Among species, again on the wintering grounds, and the ability to tolerate cold weather observations of marked individuals depends partly on body size, as larger have shown that pairs from the previ­ species benefit from a relatively slower ous season may reunite if both metabolic rate, reduced surface-to-vol- members return to same wintering ume ratio, and thicker plumage locality (Savard 1985; Mjelstad & compared to smaller species (Blem Scetersdal 1990; Robertson et at. 1998; 1990; Dawson & O'Connor 1996). Williams & McKinney 1998). The timing Perhaps even more important is the of pair formation differs between higher ability of larger species to store species and populations (Weller 1965; endogenous reserves (Skutch 1962; Hepp & Hair 1983; Rohwer & Anderson Afton 1980; Afton & Paulus 1992). 1988). The formation and maintenance Ducks, swans and geese form pairs of a pair bond is costly to the male, on the wintering grounds and migrate since he must actively defend the to the breeding grounds in pairs female from rival males and predators (McKinney 1986; Rohwer & Andersson (Afton & Sayler 1982; Wishart 1983). On Pair formation and climate 57 the other hand, pairing is beneficial to has been described in detail by the females because paired females Egilsson (1985). The climate is subarc­ are able to forage more efficiently than tic and oceanic. January is the coldest single females (Paulus 1983). month with a mean monthly tempera­ Mallard Anas platyrhynchos, Wigeon ture of -0.5°C and an average number Anas penelope and Teal Anas crecca are of 21 frost days. Along the coast there common breeding birds in Iceland. The are important estuarine and marine Mallard population apparently does not habitats used by waterfowl and shore­ leave Iceland during winter, but the birds. Several lakes, ponds and majority of Wigeon and Teal migrate to streams are open through the winter. western Europe, mostly the British Isles (Gardarsson 1975). However, Observations some Wigeon and Teal overwinter in south Iceland. In Iceland, all three Mallard, Wigeon and Teal were species are at the northern edge of counted twice per month from both their breeding and wintering November to February, and in April. range (Cramp & Simmons 1978). During autumn and spring migrations, The aim of this study was to additional counts were made to ensure describe and compare the pairing adequate sample sizes: four counts chronology of Mallard, Wigeon and Teal were made in September, and three in in Iceland. Wintering in Iceland, these October and March, respectively. The species are evidently near the lower counts were made in the four hours limit of their temperature tolerance, around low tide. pairing chronologies different from The birds were identified as pairs those reported at southern locations and single males and females. Pairing can be expected. In order to test this status was assigned after Paulus prediction, the results were compared (1983): Birds were considered paired if with published data from North they: (1) Mutually avoided or chased America and Europe. other birds. (2) Exhibited consistent synchrony in activity, especially loco­ motion. (3) Remained within 3 m Methods distance from each other. To reduce risk of error, birds in very dense tempo­ Study area rary flocks, such as ducks being fed by Observations were made on the the public, were excluded from the southern coastline of Faxafloi, south­ analysis. For each set of observations west Iceland, from September 1997 to of pairing status of females, 95% confi­ April 1998 and from August to April dence limits were estimated from the 1999. Reykjavik (64°09'N, 21°57'W) lies Standard Error of Proportion, tJlpq/n], at the centre of the study area. The area where t is Student's t, P=proportion paired females, q=proportion unpaired 58 Pair formation and climate females and n=fotal number of females Mallard paired mostly in October, examined. Wigeon and Teal at a slower rate through the winter (Figure 1). All three Comparison with other areas species began pairing in the autumn, Mallard by early September, Wigeon in The pairing chronology of Mallard mid-September and Teal at the end of has been described for several loca­ September. By late October, 60-70% of tions on both sides of the Atlantic the female Mallard had paired. The Ocean. Three studies have been carried percentage of paired female Mallard out in continental Europe: in Munich, remained at around 60% until early Germany (Bezzel 1959); in Middelburg, March. In late April, at the beginning of The Netherlands (Lebret 1961) and in the nesting season, 75% of all females Helsinki, Finland (Raitasuo 1964). In were paired in 1998 and 90% in 1999. At North America, pair formation of the end of October, the percentage of Mallard has been studied in Ithaca, paired Wigeon females was 29% in New York (Johnsgard 1960b); in the 1997 and 19% in 1998. In March, 70% of Mingo basin, Missouri (Heitmeyer 1987) all female Wigeon had become paired, and on the Atchafalaya River delta, and in April well over 80%. Pairing pro­ Louisiana (Johnson & Rohwer 1998). gressed at a sim ilar rate in Teal as in The relationship between the pro­ Wigeon; by March about 70% of the portion of paired females and mean females had paired and in April about monthly temperatures was tested by 90%. analysis of covariance (ANCOVA, PROC Differences in the progress of pair­ GLM, SAS Institute 1990). The final ing between Mallard and the other two model was determined by backward species were most pronounced, and selection. The variables entered into highly significant in October and the initial model were: month (October- November. For instance, combined March), as a class variable; mean observations from 31 October 1997 and monthly temperature for each location; 23 October 1998 yield %2=219.65 and the month χ temperature interac­ (P<0.001) for Mallard vs. Wigeon, tion. Data residuals were tested for χ 2=22.63 (P<0.001) for Mallard vs. Teal normality by the Shapiro-Wilkes test and χ 2=8.03 (P<0.01 ) for Wigeon vs. (SAS Institute 1990). Teal; and for 14 November 1997 plus 3 November 1998 χ 2=51.92 (P<0.001) for Results Mallard vs. Wigeon, χ 2=20.81 (P<0.001) for Mallard vs. Teal, but χ 2=0.002 (n.s.) Observations in Iceland for Wigeon vs. Teal. Mallard used mainly two habitats: Pairing chronology of all three intertidal areas, either mudflats or sea­ species was sim ilar in 1997-98 and weed beds, or urban fresh waters.
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