Colony Attendance of Least Auklets at St. Paul Island, Alaska: Implications for Population Monitoring
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TheCondor94:93-100 0 The Cooper Ornithological Society 1992 COLONY ATTENDANCE OF LEAST AUIUETS AT ST. PAUL ISLAND, ALASKA: IMPLICATIONS FOR POPULATION MONITORING ’ IAN L. JONES~ Department of Biology, Queens’ University,Kingston, Ontario K7L 3N6, Canada Abstract. Colony attendance by Least Auklets (Aethia pusilla) was monitored during three breeding seasonsat St. Paul Island, Alaska. Maximum counts of birds attending one 150-m2 study plot varied significantly from year to year, with a nearly two-fold difference between highest and lowest years. Maximum numbers on the surface at any one time amounted to about half of the local breeding population. Attendance was high in the year with greatestproportion of adults breeding, low in the year with lowest proportion of adults breeding, and the proportion of adults breeding differed significantly among years. Adult attendance both early and late in breeding seasondiffered significantly among years, but there was no evidence that varying attendance related to changesin the overall adult pop- ulation. Counts during incubation and chick-rearingstages were affected by sub-adults(two- year-olds), which differed significantly in attendance from year to year and sometimes representedup to half the birds on the colony surface.Year-to-year changesin surfacecounts probably related to strength of this sub-adult cohort and to varying attendancebehavior of adults and sub-adultsthat correlated with food availabilitv. These data suavestthat. taken alone, surface counts at Least Auklet colonies must be interpreted cautio&ly in assessing population changes.Suggestions for improvement of countingtechniques and an alternative approach to population monitoring are discussed. Key words: Alcidae; Aethia pusilla; census;colony attendance; Least Auklet; population monitoring;reproduction; seabird; survival. INTRODUCTION result from extreme within-day, daily and sea- The Least Auklet (Aethia pusilla) is a plankton- sonal variability of colony attendance. feeding seabird endemic to the Bering Sea and Attempts to monitor auklet populations have adjacent North Pacific waters and is the most involved counts of birds active on the surfaceof abundant breeding seabird of the region (Sowls rocky slopesof breeding colonies (Bedard 1969, et al. 1978). Least and other auklets represent an Searing 1977, Byrd et al. 1983, Piatt et al. 1990a). important component of the local marine eco- These counts are normally timed to coincide with system, which is thought to be in a dynamic state daily peak periods of abundance and replicate due to natural and man-made disturbances counts are normally conducted during the period (Springer and Roseneau 1985). However, auklet of breeding seasonwhen day-to-day variation in populations have proved to be difficult to census, attendance is lowest. The mean of these counts population estimates for most colonies are rough is presumed to be useful for population moni- estimates, and as yet there is no acceptedmethod toring. For example, changesin counts of auklets of monitoring population changes (Sowls et al. at Kongkok Bay, St. Lawrence Island, in 1964 1978, Piatt et al. 1990a). Problems with counting (Bedard 1969) 1976 (Searing 1977) and 1987 auklets arise becausemost breeding sitesare con- (Piatt et al. 1990a) have been interpreted as pos- cealed in inaccessible crevices in talus that may sible evidence for a massive population increase be several meters deep; therefore, populations (auklet counts increased two-fold over the peri- must be estimated indirectly from the number od). However, there are few data available on of birds on the colony surface.Further difficulties inter-annual variability in colony attendance. A key question is whether changesin surfacecounts from one year to the next, or over longer periods, reflect real changesin auklet populations. I Received 29 April 1991. Accepted 9 September Here I present information on colony atten- 1991. 2Present address:Department of Zoology, Univer- dance of Least Auklets obtained as part ofa study sity of Cambridge, Downing St., Cambridge CB2 3EJ, of behavioral ecology of the speciesin the Prib- United Kingdom. ilof Islands, Alaska. Previous studies have in- I931 94 IAN L. JONES volved counting large numbers of auklets on the daytime activity period to estimate the daily ac- surfaceat many plots. Here I report results of an tivity peak. Although I never directly compared intensive investigation of colony attendance be- the results of these two counting methods, my havior at a single study plot, utilizing a popu- counting technique is likely to compare favora- lation of color-marked auklets. The objectives of bly with auklet counts made in previous studies this study were (1) to document inter-annual because it also estimates peak numbers (John variation in colony attendance, (2) to identify Piatt, pers. comm.). In this study, sub-adults (two factors that influence attendance and ultimately year old birds) were distinguished by their brown (3) to assessthe value of surfacecounts for mon- foreheads with restricted nuptial plumes, worn itoring auklet population changes. flight feathers and spotted throats (Btdard and Sealy 1984, Jones and Montgomerie, in press METHODS further details below). Colony attendance was monitored at a colony of To monitor activity of individual auklets at more than 10,000 Least Auklets near Tolstoi the study plot, a color-marked population of a Point, St. Paul Island, Pribilof Islands, Alaska minimum of about 200 regularly attending birds (57”08’N, 170”17’W) during May to August of was maintained. With help from field assistants 1987, 1988 and 1989. At this site, auklets nest I captured and color-marked 248 auklets (219 among sparsely vegetated beach boulders and adults, 29 sub-adults) in 1987, 369 (306 adults, talus, and in adjacent cliffcrevices. I made counts 63 sub-adults) in 1988, and 145 (all adults) in of auklets visible on the talus (“surface counts”) 1989. To minimize disturbance, banding was re- at one densely occupied 10 m x 15 m study plot stricted to the pre-laying period, banding took on a talus slope about 5 m above sea level. This place every fourth day, and birds were processed site appeared to be representative of auklet hab- and releasedas quickly as possible after capture. itat at St. Paul and had similar densities of auk- Capture and handling of thesebirds did not affect lets on the surface to those reported in previous their reproductive performance or likelihood of studies (e.g., Piatt et al. 1990a). Auklet counts returning in following years (Jones, unpubl. were made within the boundaries of one study manuscript). Each auklet was given a numbered plot between 25 May-l August 1987, 15 May- USFWS stainless steel band and three Darvik@ 9 August 1988, and 8 May-12 August 1989 (Fig. plastic color bands (see Jones and Montgomerie 1) during a four-hour monitoring period timed 199 1, in press, for further details). to include the daily peak of auklet attendance Chick growth and fledging successwere not (11:00-l 8:00 hr Alaska Daylight Saving Time). monitored directly because most Least Auklets Surface counts were made regularly during the nested in inaccessible crevices at Tolstoi, and pre-laying and laying periods,and daily after June because disturbance of nesting crevices reduces 27 in 1987, June 26 in 1988 and June 25 in 1989 reproductive success(Roby and Brink 1986a, until auklet surface activity ceasedat the end of Piatt et al. 1990b). Consequently, breeding per- the breeding season(Fig. 1). The plot boundaries formance and phenology were evaluated by ob- remained the same throughout the study. The serving chick-provisioning behavior of color- monitoring period was shifted later in the day as marked birds. Least Auklets have a clutch size the breeding season progressed,to parallel the of one, and food is delivered to the chick until shift in peak colony attendance. All birds within it fledges 26-31 days after hatching (Roby and plot boundaries were counted at least every 30 Brink 1986a). Starting on the day when the first min and then as frequently as necessaryat time auklet was seen delivering food, watches at the of peak attendance to estimate each day’s highest plot were extended to seven hours daily, encom- number of birds attending at one time. This in- passing the daytime and evening peaks of food volved repeated counts every few minutes about delivery (see Jones and Montgomerie, 1991; in the time of peak numbers on the surface. I used press). Marked auklets that delivered food to single highest counts of adults and sub-adults chicks on at least two occasions were classified made each day to estimate day-to-day variability as active breeders, those never seen delivering in attendance of each age group. BCdard (1969) food were classified as non-breeders. Use of the Searing (1977) and Piatt et al. (1990a) all used term “active breeder” implies successat least to the average of second, third, and fourth highest hatching. Hatching dates were estimated from regular counts of all birds made throughout the date of first appearance of each marked breeding LEAST AUKLET COLONY ATTENDANCE 95 incubation chickrearing . 4 1988 May 1 June 1 July1 August 1 Date FIGURE 1. Patternsof daily variation in colonyattendance of LeastAuklets at Tolstoi Point, St. Paul Island, 1987-1989. Daily adult countsmay be determinedby subtractingthe sub-adultcount from the count of all birds. auklet with a food load and fledging successwas the study plot and throughoutthe colony. In 1988, assumed if a marked bird was observed deliv- attendance was highest during laying and pre- ering food for at least 25 consecutive days. The laying periods, while in 1989 attendance peaked date of the peak of hatching for each year of this late in the incubation period (Fig. 1). Day-to-day study was inferred from the distribution of es- fluctuation in attendance was high in 1989 com- timated hatch dates (n = 184 in 1987, n = 131 pared to 1988 (Fig.