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Age-Specific Survival and Philopatry in Three Species of European Ducks: a Long-Term Study ’

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Age-Specific Survival and Philopatry in Three Species of European Ducks: a Long-Term Study ’ The Condor98:61-74 0 The CooperOrnithological Society 1996 AGE-SPECIFIC SURVIVAL AND PHILOPATRY IN THREE SPECIES OF EUROPEAN DUCKS: A LONG-TERM STUDY ’ PETERBLIJMS,~ AIVARS MEDNIS,~LMARS BAUGA Instituteof Biology,Latvian Academy of Sciences,Micra 3, LV-2169, Salaspils,Latvia JAMES D. NICHOLS AND JAMES E. HINES PatuxentEnvironmental Science Center, National Biological Service, Laurel, MD 20708 Abstract. Capture-recaptureand band recovery modelswere usedto estimateage-specific survival probabilities for female Northern Shovelers (Anasclypeata), Common Pochards (Aythyaferina), and Tufted Ducks (Aythya.fuZigula)at Engure Marsh, Latvia, in 1964-1993. We banded more than 65,100 day-old ducklings of both sexesand captured 10,211 incu- bating females (3,713 new bandings and 6,498 recaptures).We developed a set of 3-age capture-recapturemodels to estimate annual survival rates for female ducklings,yearlings (SY), and adults (ASY) using programs SURGE and SURVIV and selectedparsimonious models usina a method develoned bv Akaike (1973). Survival rates of SY and ASY females were highest-forTufted Ducks intermediate for Common Pochards,and lowest for Northern Shovelers. Survival rates of SY and ASY females varied in parallel for shovelers and po- chards. We believe that much of the difference in survival estimates between SY and ASY birds was caused by mortality rather than permanent emigration. Estimates of day-old duckling survival, reflecting both mortality and permanent emigration, were 0.12 for shov- eler, 0.06 for pochard, and 0.03 for Tufted Duck. For all species,duckling survival varied over years, but the pattern of variation was not similar to that of the other age classes. Estimatesof survival usingband recovery data for SY + ASY female pochardsand Tufted Ducks were similar to the capture-recaptureestimates, suggestingthat surviving females returned to the breeding marsh with probabilities approaching 1. Key words: age-speciJicfemale survival;Anatidae; Anas clypeata;Aythya ferina; Aythya fuligula; breedingphilopatry;permanent emigration; Latvia. INTRODUCTION Common Goldeneye, Bucephala clangula (Dow Capture-recaptureand band recovery models are and Fredga 1984) or island nesting Common Ei- becoming increasingly important for estimation der, Somateria mollissima (Coulson 1984), all of of survival rates of birds, including waterfowl. which are relatively easy to capture. Traditionally, band recovery models have been Most evidence on breeding philopatry in fe- used extensively to estimate survival rates of male waterfowl comes from estimates of return ducksand geese(reviewed by Johnsonet al. 1992), rate, however, this statistic incorporates three but capture-recapturemodels have received little probabilities and can be only used to draw ten- attention, primarily becauseof the lack of long- tative inferences about homing (Anderson et al. term studieswith marked birds. There have been 1992, Johnsonet al. 1992). If both band recovery few long-term capture-recapture studies of and capture-recapture survival estimates can be breeding ducks that estimated survival proba- obtained for breeding females banded in a par- bilities basedon modem statisticalmethods. Most ticular location then it is possible to directly es- ofthese have focusedon box-nesting speciessuch timate unconfounded homing probability (see as Wood Duck, Aix sponsa (Hepp et al. 1987, Methods). Becauseof the lack of adequate data Dugger 1991, Hepp and Kennamer 1993) and no suchestimates are currently available (but see Hepp et al. 1987). In this paper we use 27 years of band recovery and 18 years of capture-recapture data to esti- mate survival and breeding probabilities of fe- LReceived 31 March 1995. Accepted 9 November male ground-nesting and over-water nesting 1995. * Presentaddress: Gaylord Memorial Laboratory,The ducks on a single study area in Latvia. Aging of School of Natural Resources,University of Missouri- breeding females and banding of day-old duck- Columbia, Puxico, MO 63960, USA. lings with special oval bands (Blums et al. 1994), 1611 62 PETER BLUMS ET AL. permitted inferencesthat have not been possible tions, many artificial elevated islands were con- using previous methods (i.e., we estimated sur- structed on the flooded sections of two natural vival probabilities for females of three age class- islands during 198 1-1983 (Blums and Mednis es). To determine whether females are philopa- 199 1). Beginning in 1984,82 islands totaling 14.3 tric, we compared survival estimates from band ha in area were available for nesting within the recovery and capture-recapturemodels, and also previous island territory. We believe that frag- evaluated band recovery locations of females mentation of large islands increasedthe carrying captured on the nests. capacity of island breeding habitats despite the substantial decreasein total surface area. This METHODS was confirmed by the highest ever number of STUDY AREA AND BREEDING duck nests recorded on all islands in the early POPULATIONS OF DUCKS 1990s (Blums et al. 1993). A long-term capture-recapturestudy of Northern In addition to the natural and artificial islands, Shoveler (Anus clypeuta, hereafter shoveler), nest searcheswere expanded to three isolated Common Pochard (Aythyu ferina, hereafter po- areas of persistent emergent marsh in 1972, to- chard), and Tufted Duck (Aythyu fuligulu) was taling approximately 111 ha of reed-beds and conducted from 1958 to 1993 on Engure Marsh, cattail stands, excluding open water. Thus, per- Latvia, Eastern Europe. The 35-km2 Engure manent sampling areas included natural and ar- Marsh is a shallow, permanently-flooded pal- tificial islands, 1958-1993, and emergent marsh- ustrine marsh (Cowardin et al. 1979) on the east es, 1972-1992. coast of the Baltic Sea (57”15’N, 23”07’E). The During the last 20 years, the marsh supported marsh has gradually changed from an open to a about 2,000 breeding pairs of ducks, with about hemi-marsh (Weller and Spatcher 1965) domi- 60% consisting of pochards, Tufted Ducks, and nated by tall, robust hydrophytes such as Com- shovelers.The averagenumbers of breeding pairs mon Reed (Phrugmitesuustralis) and cattail (Ty- on the entire marsh during 1977-1993 were as phu spp.). Human activities are prohibited on follows (Blums et al. 1993): Common Pochard islands and areas of emergent vegetation during 900 (range 560-1640), Tufted Duck 280 (160- the breeding season but most of the marsh is 360), and Northern Shoveler 33 (range 19-59). open to waterfowl hunting during early August Of thesenumbers 99% of shovelers,42% of Tuft- through early November. Different management ed Ducks, and 23% of pochards nested within activities (such as construction of artificial is- permanent sampling areas. Shovelers nested al- lands, vegetation and predator control, and at- most exclusively on the islands within perma- traction of gulls and terns) were conducted on all nent sampling areas, thus the entire breeding sampling areas throughout the study that pre- population was monitored each year during rou- vented the decline of carrying capacity of breed- tine nest searches.The breeding populations of ing habitats (Mihelsons et al. 1976, Blums and Tufted Ducks and shovelers were fairly stable Viksne 1990, Blums and Mednis 1991). Preda- throughout the study period however, the num- tors were systematically controlled for two to ber of shovelers increased substantially during three months beginning with the break-up of ice, the last five years (Blums et al. 1993). Pochards and an average of 11 (range 1-17) American increased during the last 16 years. Minks (Musteluvison), 83 (44-l 44) Marsh Har- riers (Circus aeruginosus),21 (O-63) Ravens FIELD METHODS (Corvuscorux), 17 (6-34) Hooded Crows (Corvus We conducted two to three complete searchesfor corone),14 (2-25) Herring Gulls (Larusurgen- duck nestson the permanent sampling areasfrom tutus),etc., were removed or relocated out of the mid-May to mid-June. All breeding habitats marsh during eachbreeding season(records from within permanent sampling areas were system- 1978 through 1993). atically searchedto locate nests by walking par- Permanent sampling areas included five nat- allel transects.We adjusted the distance between ural islands with a total surfacearea at low water transectsfrom 1.5 to 3.0 m in relation to vege- of approximately 20 ha, from 1958 to 198 1. Large tation density and height. Additional intensive portions of theseislands were flooded during high effortswere made each breeding seasonto locate water conditions and were not always suitable new nests by flushing females and watching lone for nesting. To maintain stable breeding condi- individuals of both sexes.We believe the effec- SURVIVAL AND PHILOPATRY OF FEMALE DUCKS 63 tivenessof nest searcheswas high becausecertain ings or a narrow brownish ring at the inner part sampling areas were assigned to the biologists of iris. who worked on the same areas for lo-30 con- Newly hatched ducklings were captured by secutive field seasonsand knew potential nest hand at nests and banded with plasticine-filled sites. That knowledge was especially important oval aluminum bands (Blums et al. 1994) rather for finding nestsin reed-beds.Experimental burns than with the web tagscommonly usedby Amer- on islands, after the last nests hatched, verified ican biologists(Grice and Rogers 1965, Haramis that more than 95% of all nests were found each and Nice 1980). Band loss for ducklings was es- year. We captured incubating females on nests timated to be extremely low, ~0.5% (Blums et during
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