David H. Ward

Distribution and stability of eelgrass beds at Izembek Lagoon, Alaska

Spatial change in eelgrass meadows, Zostera marina L., was assessed between 1978 and 1987 and between 1987 and 1995 at Izembek Lagoon, Alaska. Change in total extent was evaluated through a map to map comparison of data interpreted from a 1978 Landsat multi-spectral scanner image and 1987 black and white aerial photographs. A ground survey in 1995 was used to assess spatial change from 1987. Eelgrass beds were the predominant vegetation type in the lagoon, comprising 44–47% (15 000–16 000 ha) of the total area in 1978 and 1987. Izembek Lagoon contains the largest bed of seagrass along the Pacific Coast of North America and largest known single stand of eelgrass in the world. There was a high degree of overlap in the spatial distribution of eelgrass among years of change detection. The overall net change was a 6% gain between 1978 and 1987 and a < 1% gain between 1987 and 1995. The lack of significant change in eelgrass cover suggests that eelgrass meadows in Izembek Lagoon have been stable during the 17-year period of our study.

Seasonal and annual survival of adult Pacific brant

Declining mid-winter counts of Pacific brant (Branta bernicla nigricans) and reduced numbers of nesting birds on their main breeding grounds prompted us to assess factors that may be limiting recovery of this population. We estimated seasonal and annual survival rates of adult brant in 1986-93 from resightings of leg-banded birds. Brant were banded at a major colony on the Yukon-Kuskokwim Delta, Alaska (Y-K Delta) in 1986-92, and resighted there in 1987-93 as well as at major fall and spring migration and wintering areas in 1990-93. Seasonal survival was the same for males and females. Mean monthly survival rate was lowest (P ≤ 0.05) in late spring migration (15 Apr-1 Jun), the period of greatest subsistence harvest on the breeding grounds, and highest in winter (1 Jan-1 Mar), the period of greatest sport harvest. Annual survival rate did not vary among years (F = 0.51; 5, 718 df; P = 0.91) and averaged 0.840 (SE = 0.031) from 1986 to 1993. Subsistence harvest has contributed to low population levels of Pacific brant.

Population genetic structure of annual and perennial populations of Zostera marina L. along the Pacific coast of Baja California and the Gulf of California

The Baja California peninsula represents a biogeographical boundary contributing to regional differentiation among populations of marine animals. We investigated the genetic characteristics of perennial and annual populations of the marine angiosperm, Zostera marina, along the Pacific coast of Baja California and in the Gulf of California, respectively. Populations of Z. marina from five coastal lagoons along the Pacific coast and four sites in the Gulf of California were studied using nine microsatellite loci. Analyses of variance revealed significant interregional differentiation, but no subregional differentiation. Significant spatial differentiation, assessed using θST values, was observed among all populations within the two regions. Z. marina populations along the Pacific coast are separated by more than 220 km and had the greatest θST (0.13–0.28) values, suggesting restricted gene flow. In contrast, lower but still significant genetic differentiation was observed among populations within the Gulf of California (θST = 0.04–0.18), even though populations are separated by more than 250 km. This suggests higher levels of gene flow among Gulf of California populations relative to Pacific coast populations. Direction of gene flow was predominantly southward among Pacific coast populations, whereas no dominant polarity in the Gulf of California populations was observed. The test for isolation by distance (IBD) showed a significant correlation between genetic and geographical distances in Gulf of California populations, but not in Pacific coast populations, perhaps because of shifts in currents during El Niño Southern Oscillation (ENSO) events along the Pacific coast.

Effects of harness-attached transmitters on premigration and reproduction of brant

Radio transmitters are an important tool in waterfowl ecology studies, but little is known about their effects on free-ranging geese. We attached transmitters to female brant (Branta bernicla nigricans) to investigate migration schedules at a fall staging area, return rates to nesting grounds, and nesting rates of returning females in subsequent breeding seasons. Radio-tagged females (n = 62) carried either 35-g back-mounted transmitters attached with ribbon harnesses, or 26 or 32-g back-mounted transmitters affixed with plastic-coated wire harnesses (Dwyer 1972). Arrival and departure schedules at Izembek Lagoon, Alaska, did not differ (P > 0.05) between radio-tagged females and the entire population in 1987-89. Color-banded females with transmitters returned to the breeding colony in subsequent nesting seasons (1988-92) at a lower (P ≤ 0.003) rate (≤4%) than color-banded females without transmitters (57-83%). The 1 returning color-banded female with a transmitter did not breed, while an average 90% of the returning color-banded females without transmitters nested in subsequent breeding seasons (P = 0.005). Back-mounted, harness-attached transmitters may bias data in studies of waterfowl behavior, productivity, and survival

Fidelity and breeding probability related to population density and individual quality in black brent geese Branta bernicla nigricans

1. Patterns of temporary emigration (associated with non-breeding) are important components of variation in individual quality. Permanent emigration from the natal area has important implications for both individual fitness and local population dynamics. 2. We estimated both permanent and temporary emigration of black brent geese (Branta bernicla nigricans Lawrence) from the Tutakoke River colony, using observations of marked brent geese on breeding and wintering areas, and recoveries of ringed individuals by hunters. We used the likelihood developed by Lindberg, Kendall, Hines & Anderson 2001 (Combining band recovery data and Pollocks robust design to model temporary and permanent emigration. Biometrics, 57, 273-281) to assess hypotheses and estimate parameters. 3. Temporary emigration (the converse of breeding) varied among age classes up to age 5, and differed between individuals that bred in the previous years vs. those that did not. Consistent with the hypothesis of variation in individual quality, individuals with a higher probability of breeding in one year also had a higher probability of breeding the next year. 4. Natal fidelity of females ranged from 0.70 +/- 0.07-0.96 +/- 0.18 and averaged 0.83. In contrast to Lindberg et al. (1998), we did not detect a relationship between fidelity and local population density. Natal fidelity was negatively correlated with first-year survival, suggesting that competition among individuals of the same age for breeding territories influenced dispersal. Once females nested at the Tutakoke River, colony breeding fidelity was 1.0. 5. Our analyses show substantial variation in individual quality associated with fitness, which other analyses suggest is strongly influenced by early environment. Our analyses also suggest substantial interchange among breeding colonies of brent geese, as first shown by Lindberg et al. (1998).

Response of fall-staging brant and Canada geese to aircraft overflights in southwestern Alaska

Because much of the information concerning disturbance of waterfowl by aircraft is anecdotal we examined behavioral responses of Pacific brant (Brauta beruicla nigricans) and Canada geese (B. canadensis taverneri) to experimental overflights during fall stagmg at Izembek Lagoon, Alaska. These data were used to develop predictive models of brant and Canada goose response to aircraft altitude, type, noise, and lateral distance from flocks. Overall, 75% of brant flocks and 9% of Canada goose flocks flew in response to overflights. Mean flight and alert responses of both species were greater for rotary-wing than for fixed-wing aircraft and for high-noise than for low-noise aircraft. Increased lateral distance between an aircraft and a flock was the most consistent predictive parameter associated with lower probability of a response by geese. Altitude was a less reliable predictor because of interaction effects with aircraft type and noise. Although mean respouse of brant and Canada geese nerally was inversely proportional to aireraft altitude, reatest respouse occurred at intermediate (305-760 m) altitudes. At Izembek Lagoon and other areas where there are large concentrations of waterfowl, managers should consider lateral distance from the birds as the primary criterion for establishing local flight restrictions, especially for helicopters.

Testing for handling bias in survival estimation for black brant

We used an ultrastructure approach in program SURVIV to test for, and remove, bias in survival estimates for the year following mass banding of female black brant (Branta bernicla nigricans). We used relative banding-drive size as the independent variable to control for handling effects in our ultrastructure models, which took the form: S = S 0 (l - αD), where α was handling effect and D was the ratio of banding-drive size to the largest banding drive. Brant were divided into 3 classes: goslings, initial captures, and recaptures, based on their state at the time of banding, because we anticipated the potential for heterogeneity in model parameters among classes of brant. Among models examined, for which a was not constrained, a model with α constant across classes of brant and years, constant survival rates among years for initially captured brant but year-specific survival rates for goslings and recaptures, and year- and class-specific detection probabilities had the lowest Akaike Information Criterion (AIC). Handling effect, a, was -0.47 ± 0.13 SE, -0.14 ± 0.057, and -0.12 ± 0.049 for goslings, initially released adults, and recaptured adults. Gosling annual survival in the first year ranged from 0.738 ± 0.072 for the 1986 cohort to 0.260 ± 0.025 for the 1991 cohort. Inclusion of winter observations; increased estimates of first-year survival rates by an average of 30%, suggesting that permanent emigration had an important influence on apparent survival, especially for later cohorts. We estimated annual survival for initially captured brant as 0.782 ± 0.013, while that for recaptures varied from 0.726 ± 0.034 to 0.900 ± 0.062, Our analyses failed to detect a negative effect of handling on survival of brant, which is consistent with an hypothesis of substantial inherent heterogeneity in post-fledging survival rates, such that individuals most likely to die as a result of handling also have lower inherent survival probabilities.

Effects of el nino on distribution and reproductive performance of black brant

Climate in low-latitude wintering areas may influence temperate and high-latitude breeding populations of birds, but demonstrations of such relationships have been rare because of difficulties in linking wintering with breeding populations. We used long-term aerial surveys in Mexican wintering areas and breeding areas in Alaska, USA, to assess numbers of Black Brant (Branta bernicla nigricans; hereafter brant) on their principal wintering and breeding area in El Niño and non-El Niño years. We used Pollocks robust design to directly estimate probability of breeding and apparent annual survival of individually marked brant at the Tutakoke River (TR) colony, Alaska, in each year between 1988 and 2001. Fewer brant wintered in Mexico during every El Niño event since 1965. Fewer brant were observed on the principal breeding area following each El Niño since surveys began in 1985. Probability of breeding was negatively related to January sea surface temperature along the subtropical coast of North America during the preceding winter. Between 23% (five-year-olds or older) and 30% (three-year-olds) fewer brant nested in 1998 following the strong El Niño event in the winter of 1997-1998 than in non-El Niño years. This finding is consistent with life history theory, which predicts that longer-lived species preserve adult survival at the expense of reproduction. Oceanographic conditions off Baja California, apparently by their effect on Zostera marina (eelgrass), strongly influence winter distribution of brant geese and their reproduction (but not survival), which in turn affects ecosystem dynamics in Alaska.

THE SIZE SELECTION OF CLAMS BY AFRICAN BLACK OYSTERCATCHERS AND KELP GULLS

The selection of prey by two species of shore-dwelling birds was compared with the predictions of a simple optimal diet model. The preferences of African Black Oystercatchers (Haematopus moquini) and Kelp Gulls (Larus dominicanus) for different size classes of the sand-burrowing bivalve Donax serra were examined. Gulls took only large clams, regardless of the availability of the different size classes. This preference is ascribed to the high energy expenditure required for gulls to open clam shells. Oystercatch- ers, which open bivalve shells relatively easily, took both small and large clams, even when large clams were abundant. The selection of small clams when large clams were abundant appears to have resulted from the lack of discrimination of clam sizes by oystercatchers and the temporal segregation of availability of small and large clams.

Carryover effects associated with winter location affect fitness, social status, and population dynamics in a long-distance migrant.

We used observations of individually marked female black brant geese (Branta bernicla nigricans; brant) at three wintering lagoons on the Pacific coast of Baja California—Laguna San Ignacio (LSI), Laguna Ojo de Liebre (LOL), and Bahía San Quintín (BSQ)—and the Tutakoke River breeding colony in Alaska to assess hypotheses about carryover effects on breeding and distribution of individuals among wintering areas. We estimated transition probabilities from wintering locations to breeding and nonbreeding by using multistratum robust-design capture-mark-recapture models. We also examined the effect of breeding on migration to wintering areas to assess the hypothesis that individuals in family groups occupied higher-quality wintering locations. We used 4,538 unique female brant in our analysis of the relationship between winter location and breeding probability. All competitive models of breeding probability contained additive effects of wintering location and the 1997–1998 El Niño–Southern Oscillation (ENSO) event on probability of breeding. Probability of breeding in non-ENSO years was , , and for females wintering at BSQ, LOL, and LSI, respectively. After the 1997–1998 ENSO event, breeding probability was between 2% (BSQ) and 38% (LOL) lower than in other years. Individuals that bred had the highest probability of migrating the next fall to the wintering area producing the highest probability of breeding.