California Gulls Research Articles

Revisiting the Past: Historical Status of Breeding Ring-billed and California Gulls in the Klamath Basin, Oregon

Three summaries of the historical and recent status of breeding Ring-billed (Larus delawarensis) and California (L. californicus) gulls in Oregon mention just two colony sites in the Klamath Basin—Upper Klamath Lake and Spring Lake—and lack numerical data for individual colonies. Nevertheless, nesting of gulls at Lower Klamath Lake before its water source was cut off in 1917 is well documented. Egg-set records, published literature, and archived field notes attest to nesting of both species before 1954 at two additional sites in the Klamath Basin, the Link and Klamath rivers. The largest known colonies up to that time were at Lower Klamath Lake (~2000 pairs, perhaps many more) and along the Klamath River below the town of Klamath Falls (~2000 pairs). Gulls no longer nest at these five sites, but, since at least 2000, they have nested intermittently at Swan Lake and Gerber Reservoir. Though Gabrielson and Jewett (1940) considered the California Gull more abundant in southeastern Oregon than the Ring-billed, egg-collectors’ data from 1926 to 1953 suggest the opposite, as in northeastern California. The California Gull may have been the more numerous of the two in the Klamath Basin when nesting and foraging habitat was dominated by wetlands, but the drying of Lower Klamath Lake and the great expansion of irrigated agriculture starting in the 1920s may have favored the Ring-billed. Trends in the Klamath Basin’s gull populations are unclear, but much nesting habitat has been lost. Documentation of the historical record of nesting gulls and other waterbirds is critical, as conservation efforts typically aim at restoring populations to some historical baseline, which can gradually shift lower with further declines and successive generations accepting a new status quo.

Predator and duck behaviours at depredated nests in wetlands of Great Salt Lake, Utah

Abstract Nest depredation is one of the greatest threats posed to ground-nesting ducks. We employed cameras to monitor 164 duck nests (71 cinnamon teal, Spatula cyanoptera, 44 gadwall, Mareca strepera, 38 mallard, Anas platyrhynchos, and 11 nests of unknown species) in the wetlands surrounding Great Salt Lake, from 2015–2021. Of the 164 nests, 21% were successful, 73% were depredated and 7% were abandoned. We observed predators at 99 of the 119 depredated nests; predators at 20 nests went undetected. Raccoons (Procyon lotor, depredated nests) and striped skunks (Mephitis mephitis, ) were the most common of the 99 nest predators recorded. Other predators that depredated nests included long-tailed weasels (Mustela frenata), northern harriers (Circus hudsonius), California gulls (Larus californicus), Sandhill cranes (Antigone canadensis), common ravens (Corvus corax), coyotes (Canis latrans) and red foxes (Vulpes vulpes). Neither the number of eggs removed per depredation event nor the number of eggs remaining varied by predator species. Depredated nests were easier for predators to find than undisturbed, incubated nests, resulting in 68% of depredated nests being visited by multiple predators. All hens detected the approach of a predator and flushed before the predator reached the nest; no hens attempted to defend their nest or attack the predator. Only 21% of hens returned to their depredated nest, and those that did remained off their nest an average of 33 h and 23 h after their nest was depredated by a raccoon or skunk, respectively. Seventeen percent of hens resumed incubation of their depredated nest, but only 1 nest to which a hen returned successfully hatched an egg. Depredation events of raccoons and skunks were not distributed randomly during the 24-hour day, but rather occurred most often during the night and nautical twilight, and rarely during the day. Depredation events of avian predators occurred during the day, rarely during twilight, and none during the night. Depredation events during the night were more likely when the wind was calm but temperature, humidity, and actual moon illumination had no impact. Depredation events by skunks and raccoons occurred more often during the 1st and 4th phases of the moon (new moon) than in the 2nd or 3rd phase.

Estimating Cause‐Specific Mortality and Survival of Juvenile Fall Chinook Salmon: An Investigation of Avian Predation across Large Spatial Scales

AbstractTo investigate the effects of piscivorous colonial waterbird predation on juvenile salmonids (smolts) and to determine what proportion of all sources of smolt mortality was due to predation, we analyzed a data set describing fall Chinook Salmon Oncorhynchus tshawytscha that were marked with PIT tags in the Columbia River, USA. We used a state‐space Bayesian model that incorporated live detections of tagged fish and recoveries of tags from dead fish to jointly estimate predation and survival over multiple river reaches and years (2008–2019). By incorporating recoveries of tags on bird colonies, the model was also able to increase the precision and spatial extent of survival estimates compared with a more traditional capture–recapture model. Results indicated that the avian predation probability (proportion of available fish consumed) varied depending on the predator species, colony location, and the fish's rearing type. Estimates of cumulative predation probability ranged annually from 0.066 (95% credible interval = 0.049–0.089) to 0.283 (0.210–0.419), and predation accounted for 7.3% (4.3–11.9%) to 29.1% (19.7–40.2%) of all sources of smolt mortality during out‐migration to Bonneville Dam. Probabilities of predation on wild smolts were significantly higher than those for hatchery smolts in most river reaches and years. Predation probabilities were the highest for colonies of American white pelicans Pelecanus erythrorhynchos and double‐crested cormorants Nannopterum auritum, which were located near where wild smolts rear and congregate. Estimates of predation by Caspian terns Hydroprogne caspia, California gulls Larus californicus, and ring‐billed gulls L. delawarensis were often <0.03 of available smolts annually. Collectively, results indicated that the cumulative effects of predation on Chinook Salmon smolts were substantial in some but not all river reaches and years. Future models that consider biological and environmental factors influencing smolt susceptibility to avian predation may provide a more holistic understanding of the degree to which predation limits fish survival.

Predation probabilities and functional responses: How piscivorous waterbirds respond to pulses in fish abundance

How predators respond to changes in prey abundance (i.e., functional responses) is foundational to consumer–resource interactions, predator–prey dynamics, and the stability of predator–prey systems. Predation by piscivorous waterbirds on out-migrating juvenile steelhead trout (Oncorhynchus mykiss) is considered a factor affecting the recovery of multiple Endangered Species Act-listed steelhead populations in the Columbia River basin. Waterbird functional responses, however, may vary by predator species and location, with important implications to predator management strategies. We used a 13-year dataset on waterbird abundance across seven breeding colonies (three Caspian tern [Hydroprogne caspia], two double-crested cormorant [Nannopterum auritum], and two California and ring-billed gull [Larus californicus and Larus delawarensis] colonies) and steelhead tag-recovery data (>645,000 tagged and >32,000 recovered steelhead) to quantify weekly predation probabilities and functional responses across waterbird species, colonies, and years. Weekly predation probabilities were highly variable, ranging from 0.01 to 0.30 at tern colonies, 0.01 to 0.20 at cormorant colonies, and 0.03 to 0.13 at gull colonies. Per capita predation probabilities were an order of magnitude higher at inland tern and cormorant colonies relative to estuary colonies of the same species. Terns displayed Type II functional responses across colonies and years, where predation probabilities peaked at low steelhead abundances and declined as steelhead abundance increased (i.e., predator swamping). Cormorants nesting at the large estuary colony (several thousand birds) displayed a Type III functional response, but cormorants nesting at the smaller inland colony (several hundred birds) displayed a Type II response. Consumption probabilities of steelhead by gulls remained consistent across a large range of steelhead availability, suggesting a Type I or a Type III functional response, but a lack of colony abundance data prevented quantifying functional responses. The level of tern predation combined with Type II functional responses indicate possible population-level impacts that could destabilize small or declining prey populations. Conversely, the apparent Type III functional responses of gulls and estuary nesting cormorants are indicative of prey switching behaviors targeted at periods of high steelhead abundance. Our results illustrate the complexity of predator–prey interactions and the importance of quantifying predator- and location-specific functional responses when predicting the efficacy of management strategies to enhance prey populations.

Cumulative effects of piscivorous colonial waterbirds on juvenile salmonids: A multi predator-prey species evaluation.

We investigated the cumulative effects of predation by piscivorous colonial waterbirds on the survival of multiple salmonid (Oncorhynchus spp.) populations listed under the U.S. Endangered Species Act (ESA) and determined what proportion of all sources of fish mortality (1 –survival) were due to birds in the Columbia River basin, USA. Anadromous juvenile salmonids (smolts) were exposed to predation by Caspian terns (Hydroprogne caspia), double-crested cormorants (Nannopterum auritum), California gulls (Larus californicus), and ring-billed gulls (L. delawarensis), birds known to consume both live and dead fish. Avian consumption and survival probabilities (proportion of available fish consumed or alive) were estimated for steelhead trout (O. mykiss), yearling Chinook salmon (O. tshawytscha), sub-yearling Chinook salmon, and sockeye salmon (O. nerka) during out-migration from the lower Snake River to the Pacific Ocean during an 11-year study period (2008–2018). Results indicated that probabilities of avian consumption varied greatly across salmonid populations, bird species, colony location, river reach, and year. Cumulative consumption probabilities (consumption by birds from all colonies combined) were consistently the highest for steelhead, with annual estimates ranging from 0.22 (95% credible interval = 0.20–0.26) to 0.51 (0.43–0.60) of available smolts. The cumulative effects of avian consumption were significantly lower for yearling and sub-yearling Chinook salmon, with consumption probabilities ranging annually from 0.04 (0.02–0.07) to 0.10 (0.07–0.15) and from 0.06 (0.3–0.09) to 0.15 (0.10–0.23), respectively. Avian consumption probabilities for sockeye salmon smolts was generally higher than for Chinook salmon smolts, but lower than for steelhead smolts, ranging annually from 0.08 (0.03–0.22) to 0.25 (0.14–0.44). Although annual consumption probabilities for birds from certain colonies were more than 0.20 of available smolts, probabilities from other colonies were less than 0.01 of available smolts, indicating that not all colonies of birds posed a substantial risk to smolt mortality. Consumption probabilities were lowest for small colonies and for colonies located a considerable distance from the Snake and Columbia rivers. Total mortality attributed to avian consumption was relatively small for Chinook salmon (less than 10%) but was the single greatest source of mortality for steelhead (greater than 50%) in all years evaluated. Results suggest that the potential benefits to salmonid populations of managing birds to reduce smolt mortality would vary widely depending on the salmonid population, the species of bird, and the size and location of the breeding colony.

Large Scale Aerial Multi-Robot Coverage Path Planning

Autonomous survey and aerial photogrammetry applications require solving a path planning problem that ensures sensor coverage over a specified area. In this work, we provide a multi-robot path planning method that can obtain this coverage over an arbitrary area of interest. We extend our previous method, path optimization for population counting with overhead robotic networks (POPCORN), by a divide-and-conquer scheme, split and link tiles (SALT), which drastically decreases the time needed for route planning. These POPCORN instances can be computed in parallel and combined with SALT in a scalable manner to produce coverage paths over very large areas of interest. To demonstrate this algorithm’s capabilities, we implemented our planning algorithm with a team of drones to conduct multiple photographic aerial wildlife surveys of the Cape Crozier Adélie penguin rookery on Ross Island, Antarctica, one of the largest Adélie penguin colonies in the world. The colony, which contains over 300,000 nesting pairs and spans over 2 km, was surveyed in about 3 hours. In contrast, previous human-piloted single-drone surveys of the same colony required over 2 days to complete. We also have deployed our survey system at several islets at Mono Lake, California, to survey a California gull colony as well as at a 2000-acre ranch in Marin, California. We provide this survey path planning tool as an open-source software package named wadl.

A Gallus gallus Model for Determining Infectivity of Zoonotic Campylobacter.

To better understand public health implications of waterfowl as reservoirs for zoonotic sources of Campylobacter in recreational waters, we developed a Gallus gallus (chick) model of infection to assess the pathogenicity of environmental isolates of Campylobacter. This method involved exposure of 1-day-old chicks through ingestion of water, the natural route of infection. Viable Campylobacter from laboratory-infected animals were monitored by using a modified non-invasive sampling of fresh chick excreta followed by a passive polycarbonate-filter migration culture assay. The method was used to evaluate the infectivities of three laboratory strains of Campylobacter spp. (Campylobacter coli, Campylobacter jejuni, and Campylobacter lari), three clinical isolates of C. jejuni, and four environmental Campylobacter spp. isolated from California gulls (Larus californicus). The results revealed that chicks were successfully infected with all laboratory and clinical isolates of Campylobacter spp. through ingestion of Campylobacter-spiked water, with infection rates ranging from <10 to >90% in a dose-dependent manner. More importantly, exposure of chicks with Campylobacter spp. isolated from Gallus gallus excreta also resulted in successful establishment of infection (≤90%). Each monitored Campylobacter spp. contained ≥7.5 × 104 CFU⋅g–1 of feces 7 days post-exposure. These results suggest that a G. gallus model can be used to assess infectivity of Campylobacter isolates, including gull and human clinical isolates. Use of an avian animal model can be applied to assess the importance of birds, such as the G. gallus, as potential contributors of waterborne-associated outbreaks of campylobacteriosis.

Cumulative Effects of Avian Predation on Upper Columbia River Steelhead

AbstractTo investigate the cumulative effects of colonial waterbird predation on fish mortality and to determine what proportion of all sources of fish mortality (1 − survival) was due to bird predation, we conducted a mark–recapture–recovery study with upper Columbia River steelhead Oncorhynchus mykiss that were PIT‐tagged and released (N = 78,409) at Rock Island Dam on the Columbia River, USA. We used a state–space Bayesian model that incorporated live detections and dead recoveries of tagged fish to jointly estimate predation and survival probabilities during smolt out‐migration to the Pacific Ocean over an 11‐year study period. Estimated cumulative (all colonies combined) avian predation probabilities ranged from 0.31 (95% credible interval [CRI] = 0.27–0.38) to 0.53 (95% CRI = 0.42–0.64) annually, indicating that avian predation was a substantial source of mortality. Of the predator species evaluated, predation by Caspian terns Hydroprogne caspia was often the highest, with predation probabilities ranging from 0.11 (95% CRI = 0.09–0.14) to 0.38 (95% CRI = 0.29–0.47). Probabilities of predation by double‐crested cormorants Phalacrocorax auritus and mixed colonies of California gulls Larus californicus and ring‐billed gulls L. delawarensis were generally lower than the probabilities for terns but were also substantial, with upwards of 0.04 (95% CRI = 0.03–0.07; cormorants) and 0.31 (95% CRI = 0.25–0.39; gulls) of steelhead consumed. Comparisons of total smolt mortality with mortality due to avian predation indicated that avian predation accounted for 42% (95% CRI = 30–56%) to 70% (95% CRI = 53–87%) of total mortality, suggesting that more steelhead were consumed by avian predators than died from all other mortality sources combined. Results indicate that avian predation, although not the original cause of steelhead declines in the basin, is now a factor limiting the survival of upper Columbia River steelhead. Using the analytical framework developed in this study, future studies can consider the cumulative impact of multiple mortality sources across large spatial and temporal scales to more fully understand the extent to which they limit fish survival.

Whole-Genome Sequencing of Four Campylobacter Strains Isolated from Gull Excreta Collected from Hobie Beach (Oxnard, CA, USA).

Campylobacter spp. are commensal organisms in avian species and are one of the leading causes of bacterial foodborne human diarrheal disease worldwide. We report the draft genome sequences of Campylobacter volucris, C. lari, and C. jejuni strains isolated from California gull (Larus californicus) excreta collected from a California beach.

Common Terns Sterna hirundo incubating Common Garden Snail shells Helix aspersa on Rockabill Island

Many ground-nesting bird species have been observed incubating foreign objects in their nests (Mellink 2002). Pine cones, golf balls, guano, hermit crabs, mammalian bones and many more objects have all been recorded in bird nests (Knight &amp; Erickson 1977; Mellink 2002; Langlois et al. 2012). A study by Conover (1985) showed that 10% of Ring-billed Gull Larus delawarensis and 6% of California Gull L. californicus nests contained a foreign object. These foreign objects were more similar in size and shape to the gull eggs than to randomly selected pebbles. Little Terns Sterna albifrons have been observed incubating egg-like pebbles for several weeks on two separate occasions at a breeding colony in Kilcoole, Co. Wicklow, Ireland (Farrelly 1990). Several hypotheses have been put forward to explain this behaviour. Coulter (1980) suggested that foreign objects may act as an incubation stimulus. Many tern and gull species, unlike other birds, have three brood patches not one (Gochfeld 1976; Reid 1987). Coulter (1980) conducted incubation studies of several species of gulls suggesting that birds incubating three eggs sat for longer periods of time, resettled less frequently, have a shorter incubation period and a higher hatching success than birds on either fewer eggs or more. He speculated that gulls are highly adapted for three-egg clutches whereby the foreign objects provide an important stimulus for incubation behaviour when there are less than three eggs (Beer 1961; Beer 1965; Baerends et al. 1970; Coulter 1980). Alternatively, the mistaken-food hypothesis suggests that a predatory bird brings a foreign object back to the nest, regurgitates the object and then proceeds to incubate it, mistaking it for one of its own eggs (Sugden 1947; Twomey 1948). A third hypothesis is that the foreign object is mistaken by the bird for one of its own eggs and rolled into the nest, in this scenario the foreign object is likely to have originated from close to the nest (Conover 1985).

California Gull (Larus californicus) Space Use and Timing of Movements in Relation to Landfills and Breeding Colonies

Expanding gull (Laridae) populations throughout the world have been attributed to the availability of anthropogenic food subsidies. The influence of landfills on California Gull (Larus californicus) space use and the timing of their movements was evaluated in San Francisco Bay, California, USA. Using radio telemetry, 108 California Gulls were tracked, > 7,000 locations were recorded, and > 1 million detections were obtained at automated logger systems placed at the two main landfills and three major breeding colonies. Population home range (31-35 km2) and core use areas (2-3 km2) overlapped landfills and colonies, and expanded after breeding. California Gull attendance at landfills (1.6-19.0 km from colonies) increased throughout breeding and post-breeding, whereas attendance at colonies was low during pre-breeding (20%-40% per day), increased during breeding (60%-80% per day), and declined into and during post-breeding (< 20% per day). California Gull attendance at landfills was greatest when garbage was delivered from 06:00 hr in the morning until 18:00 hr at night. In contrast, California Gull attendance at colonies during breeding was greater at night from 20:00 hr to 05:00 hr (50%-70% per hr) than during the day from 06:00 hr to 18:00 hr (30%-40% per hr). Landfills played a predominant role in California Gull space use and the timing of their movements in this highly urbanized estuary.

Spatial trends in a biomagnifying contaminant: Application of amino acid compound-specific stable nitrogen isotope analysis to the interpretation of bird mercury levels.

Levels of biomagnifying contaminants are greatest in high-trophic level biota (e.g., predatory birds such as gulls). Gull eggs have been used to assess contaminant spatial patterns and sources, but such assessments must consider how organism trophic position may influence spatial inferences. Stable nitrogen isotopes (δ15 N) in bulk tissue are routinely used in this context. However, bulk δ15 N values are only useful if spatial differences in baseline δ15 N values are considered. Amino acid compound-specific stable nitrogen isotope analysis can generate estimates of baseline δ15 N values and trophic position from the same sample. In the present study, eggs (n = 428) of California (Larus californicus), herring (Larus argentatus smithsonianus), and ring-billed (Larus delawarensis) gulls were used to assess spatial patterns in mercury (Hg) availability in 12 western Canadian lakes located over 14 degrees of latitude, with amino acid compound-specific stable isotope analysis adjustment of egg Hg levels for trophic position. Mean trophic position-adjusted egg Hg levels (micrograms per gram, dry wt) were greatest at sites in receiving waters of the Athabasca River (X¯ = 0.70) compared to southern (X¯ = 0.39) and northern (X¯ = 0.50) regions. Research is required to investigate factors (e.g., local Hg released as a result of human activities, processes influencing Hg methylation) which may be responsible for greater Hg availability in the lower Athabasca River basin. However, it is clear that amino acid compound-specific stable isotope analysis is a valuable tool for assessing contaminant spatial patterns. Environ Toxicol Chem 2018;37:1466-1475. © 2018 SETAC.

Volatile Methylsiloxanes and Organophosphate Esters in the Eggs of European Starlings (Sturnus vulgaris) and Congeneric Gull Species from Locations across Canada.

Volatile methylsiloxanes (VMSs) and organophosphate esters (OPEs) are two suites of chemicals that are of environmental concern as organic contaminants, but little is known about the exposure of wildlife to these contaminants, particularly in birds, in terrestrial and aquatic ecosystems. The present study investigates the spatial distributions of nine cyclic and linear VMSs and 17 OPEs in the eggs of European starlings (Sturnus vulgaris) and three congeneric gull species (i.e., herring gull (Larus argentatus), glaucous-winged gull (L. glaucescens), and California gull (L. californicus)) from nesting sites across Canada. ∑VMS concentrations for all bird eggs were dominated by decamethylcyclopentasiloxane (D5), dodecamethylcyclohexasiloxane (D6), and octamethylcyclotetrasiloxane (D4). With European starlings, birds breeding adjacent to landfill sites had eggs containing significantly greater ∑VMS concentrations (median: 178 ng g-1 wet weight (ww)) compared with those from the urban industrial (20 ng g-1 ww) and rural sites (1.3 ng g-1 ww), indicating that the landfills are important sources of VMSs to Canadian terrestrial environments. In gull eggs, the median ∑VMS concentrations were up to 254 ng g-1 ww and suggested greater detection frequencies and levels of VMSs in aquatic- versus terrestrial-feeding birds in Canada. In contrast, the detection frequency of OPEs in all European starling and gull eggs was lower than 16%. This suggested that low dietary exposure or rapid metabolism of accumulated OPEs occurs in aquatic feeding birds and may warrant further investigation for the elucidation of the reasons for these differences.

Avian diets in a saline ecosystem: Great Salt Lake, Utah, USA

Saline lakes provide a prey-rich, predator-free environment for birds to utilize during migration and stopover periods. The Great Salt Lake (GSL), Utah is the largest salt lake in North America and is utilized by millions of migratory birds. It also is host to multiple commercial endeavors. Proposed expansion of commercial use of the GSL would result in increased impounded area and water extraction for mineral production, which may increase the GSL's salinity and negatively impact invertebrate abundance. I review previous literature and synthesize diets of avian species utilizing the GSL to determine the importance of each invertebrate species, including brine shrimp (Artemia franciscana) and brine flies (Ephydra spp.), and clarify the anthropogenic impacts on food sources and avian populations. Species considered are eared grebes (Podiceps nigricollis), northern shovelers (Anas clypeata), green- winged teals (Anas crecca), common goldeneyes (Bucephala clangula), American avocets (Recurvirostra americana), black-necked stilts (Himantopus mexicanus), Wilson's phalaropes (Phalaropus tricolor), red-necked phalaropes (Phalaropus lobatus), and California gulls (Larus californicus). Brine shrimp and brine fly adults are consumed by all species considered. Alterations in prey abundance due to increased salinity may alter the ability of the GSL to support large avian populations.

Mercury contamination and stable isotopes reveal variability in foraging ecology of generalist California gulls

Environmental contaminants are a concern for animal health, but contaminant exposure can also be used as a tracer of foraging ecology. In particular, mercury (Hg) concentrations are highly variable among aquatic and terrestrial food webs as a result of habitat- and site-specific biogeochemical processes that produce the bioaccumulative form, methylmercury (MeHg). We used stable isotopes and total Hg (THg) concentrations of a generalist consumer, the California gull (Larus californicus), to examine foraging ecology and illustrate the utility of using Hg contamination as an ecological tracer under certain conditions. We identified four main foraging clusters of gulls during pre-breeding and breeding, using a traditional approach based on light stable isotopes. The foraging cluster with the highest δ15N and δ34S values in gulls (cluster 4) had mean blood THg concentrations 614% (pre-breeding) and 250% (breeding) higher than gulls with the lowest isotope values (cluster 1). Using a traditional approach of stable-isotope mixing models, we showed that breeding birds with a higher proportion of garbage in their diet (cluster 2: 63–82% garbage) corresponded to lower THg concentrations and lower δ15N and δ34S values. In contrast, gull clusters with higher THg concentrations, which were more enriched in 15N and 34S isotopes, consumed a higher proportion of more natural, estuarine prey. δ34S values, which change markedly across the terrestrial to marine habitat gradient, were positively correlated with blood THg concentrations in gulls. The linkage we observed between stable isotopes and THg concentrations suggests that Hg contamination can be used as an additional tool for understanding animal foraging across coastal habitat gradients.

Nest survival is influenced by parental behaviour and heterospecifics in a mixed‐species colony

Studies of avian nest success often focus on examining influences of variation in environmental and seasonal factors. However, in‐depth evaluations can also incorporate variation in individual incubation behaviour to further advance our understanding of avian reproductive ecology. We examined these relationships in colonially nesting Black‐crowned Night‐Herons Nycticorax nycticorax using intensive video‐monitoring methods to quantify incubation behaviours. We modelled nest survival as a function of both extrinsic factors and incubation behaviours over a 3‐year period (2010–12) on Alcatraz Island, USA. Model‐averaged parameter estimates indicated that nest survival increased as a function of greater incubation constancy (% of time spent incubating eggs within a 24‐h period), and average daily precipitation throughout the nesting stage. Common Ravens Corvus corax are the only known nest predator of Night‐Herons on Alcatraz Island, as on many other coastal Pacific islands. We also investigated the effects of heterospecific nesting of California Gulls Larus californicus and Western Gulls Larus occidentalis in a mixed‐species colony with Night‐Herons, based on nesting proximity data collected over a 2‐year period (2011–12). This second analysis indicated that, in addition to incubation behaviours, nesting heterospecifics are an important factor for explaining variation in Night‐Heron nest survival. However, contrary to our original expectation, we found that Night‐Herons experienced increased nest survival with increasing distance from gull colony boundaries. These results may apply to other areas with multiple colonial nesting species and similar predator communities and climatic patterns.

The effects of heterospecifics and climatic conditions on incubation behavior within a mixed‐species colony

Parental incubation behavior largely influences nest survival, a critical demographic process in avian population dynamics, and behaviors vary across species with different life history breeding strategies. Although research has identified nest survival advantages of mixing colonies, behavioral mechanisms that might explain these effects is largely lacking. We examined parental incubation behavior using video‐monitoring techniques on Alcatraz Island, California, of black‐crowned night‐heron Nycticorax nycticorax (hereinafter, night‐heron) in a mixed‐species colony with California gulls Larus californicus and western gulls L. occidentalis. We first quantified general nesting behaviors (i.e. incubation constancy, and nest attendance), and a suite of specific nesting behaviors (i.e. inactivity, vigilance, preening, and nest maintenance) with respect to six different daily time periods. We employed linear mixed effects models to investigate environmental and temporal factors as sources of variation in incubation constancy and nest attendance using 211 nest days across three nesting seasons (2010–2012). We found incubation constancy (percent of time on the eggs) and nest attendance (percent of time at the nest) were lower for nests that were located &lt; 3 m from one or more gull nest, which indirectly supports the predator protection hypothesis, whereby heterospecifics provide protection allowing more time for foraging and other self‐maintenance activities. To our knowledge, this is the first empirical evidence of the influence of one nesting species on the incubation behavior of another. We also identified distinct differences between incubation constancy and nest attentiveness, indicating that these biparental incubating species do not share similar energetic constraints as those that are observed for uniparental species. Additionally, we found that variation in incubation behavior was a function of temperature and precipitation, where the strength of these effects was dependent on the time of day. Overall, these findings strengthen our understanding of incubation behavior and nest ecology of a colonial‐nesting species.

Female-Female Pairs and Other Unusual Reproductive Associations in Ring-Billed and California Gulls

FEMALE-FEMALE PAIRS AND OTHER UNUSUAL REPRODUCTIVE ASSOCIATIONS IN RING-BILLED AND CALIFORNIA GULLS MICHAEL R. CONOVER, 1 •3 DON E. MILLER, I AND GEORGE L. HUNT, JR. 2 'Department of Zoology, Washington State University, Pullman, Washington 99164 USA, and 2 Department of Ecology and Evolutionary Biology, University of California, Irvine, California 92717 USA ABSTRACT.-Most Ring-billed Gulls (Larus delawarensis) and California Gulls (L. californicus) form monogamous, heterosexual pairs during the breeding season. We report here the discovery of low frequencies (1-2%) of female-female pairings in Ring-billed and California gulls in three colonies in eastern Washington. The nests of these female-female pairs usually contained 5-6 eggs rather than the more normal 2-3. Between 60-70% of the eggs in the nests of the female-female pairs were fertile. We also found evidence of brood parasitism, which appears to be one cause of 4-egg clutches, and one case of polygyny in the Ring-billed Gulls. This polygynous association built a single communal nest that was incubated by three females and one male and apparently contained the eggs of at least two females. Received 13 November 1978, accepted 21 November GULLS characteristically form monogamous, heterosexual pair-bonds and lay 2- or 3-egg clutches (Paludan 1951, Tinbergen 1953). In the Western Gull (Larus oc- cidentalis), however, up to 14% of the nests from colonies in the California Channel Islands contain 4-6 eggs, most of which are infertile (Hunt and Hunt 1977). These supernormal clutches (4-6 eggs) apparently are laid by two females that share in- cubation and chick-rearing duties. Males are seldom observed at these nests. Su- pernormal clutches also occur in colonies of the Ring-billed Gull (L. delawarensis) and the California Gull (L. californicus), two species differing from the Western Gull in several aspects of their breeding biology. Whereas the Western Gull is non- migratory and spends much of the year on widely dispersed territories on the marine islands where it breeds, the Ring-billed and California gulls are migratory, stay in the nesting grounds only during the breeding season, and breed in dense colonies located on islands of inland lakes and rivers. METHODS On 12-13 May 1978 we surveyed three mixed-species colonies of Ring-billed and California gulls (Sprague Lake, Banks Lake and Potholes Reservoir colonies) in eastern Washington to determine the frequency of supernormal clutches, egg fertility, and female-female pairs. These colonies are described by Conover et al. (in press). Egg fertility was determined by opening eggs in the field and inspecting them for developing embryos or for lacunae in the blastodisc, which indicates infertility (Hunt and Hunt 1977). To determine the sex of birds associated with various sized clutches, adults of both species of gull were noosed on their nests (Miller 1974) at the Sprague Lake colony and sexed by laparotomy (Hunt and Hunt 1977). Sexed birds were banded, dye-marked or wing-tagged, and released. The nests occupied by these gulls were subsequently observed at repeated intervals over the next month to determine whether they were brooded and attended solely by the marked birds. RESULTS AND DISCUSSION We found that 3.3% of all Ring-billed Gull clutches examined contained more than 3 eggs (Table 1). Fertility in 5- and 6-egg clutches was less than that found in Present address: Department of Biology, Ball State University, Muncie, Indiana 47306 USA. The Auk 96: 6-9. January 1979

Unintended consequences of management actions in salt pond restoration: cascading effects in trophic interactions.

Salt evaporation ponds have played an important role as habitat for migratory waterbirds across the world, however, efforts to restore and manage these habitats to maximize their conservation value has proven to be challenging. For example, salinity reduction has been a goal for restoring and managing former salt evaporation ponds to support waterbirds in the South Bay Salt Pond Restoration Project in San Francisco Bay, California, USA. Here, we describe a case study of unexpected consequences of a low-dissolved oxygen (DO) event on trophic interactions in a salt pond system following management actions to reduce salinity concentrations. We document the ramifications of an anoxic event in water quality including salinity, DO, and temperature, and in the response of the biota including prey fish biomass, numerical response by California Gulls (Larus californicus), and chick survival of Forster's Tern (Sterna forsteri). Management actions intended to protect receiving waters resulted in decreased DO concentrations that collapsed to zero for ≥ 4 consecutive days, resulting in an extensive fish kill. DO depletion likely resulted from an algal bloom that arose following transition of the pond system from high to low salinity as respiration and decomposition outpaced photosynthetic production. We measured a ≥ 6-fold increase in biomass of fish dropped on the levee by foraging avian predators compared with weeks prior to and following the low-DO event. California Gulls rapidly responded to the availability of aerobically-stressed and vulnerable fish and increased in abundance by two orders of magnitude. Mark-recapture analysis of 254 Forster's Tern chicks indicated that their survival declined substantially following the increase in gull abundance. Thus, management actions to reduce salinity concentrations resulted in cascading effects in trophic interactions that serves as a cautionary tale illustrating the importance of understanding the interaction of water quality and trophic structure when managing restoration of salt ponds.

Quantifying Avian Predation on Fish Populations: Integrating Predator‐Specific Deposition Probabilities in Tag Recovery Studies

AbstractAccurate assessment of specific mortality factors is vital to prioritize recovery actions for threatened and endangered species. For decades, tag recovery methods have been used to estimate fish mortality due to avian predation. Predation probabilities derived from fish tag recoveries on piscivorous waterbird colonies typically reflect minimum estimates of predation due to an unknown and unaccounted‐for fraction of tags that are consumed but not deposited on‐colony (i.e., deposition probability). We applied an integrated tag recovery modeling approach in a Bayesian context to estimate predation probabilities that accounted for predator‐specific tag detection and deposition probabilities in a multiple‐predator system. Studies of PIT tag deposition were conducted across three bird species nesting at seven different colonies in the Columbia River basin, USA. Tag deposition probabilities differed significantly among predator species (Caspian terns Hydroprogne caspia: deposition probability = 0.71, 95% credible interval [CRI] = 0.51–0.89; double‐crested cormorants Phalacrocorax auritus: 0.51, 95% CRI = 0.34–0.70; California gulls Larus californicus: 0.15, 95% CRI = 0.11–0.21) but showed little variation across trials within a species or across years. Data from a 6‐year study (2008–2013) of PIT‐tagged juvenile Snake River steelhead Oncorhynchus mykiss (listed as threatened under the Endangered Species Act) indicated that colony‐specific predation probabilities ranged from less than 0.01 to 0.17 and varied by predator species, colony location, and year. Integrating the predator‐specific deposition probabilities increased the predation probabilities by a factor of approximately 1.4 for Caspian terns, 2.0 for double‐crested cormorants, and 6.7 for California gulls compared with traditional minimum predation rate methods, which do not account for deposition probabilities. Results supported previous findings on the high predation impacts from strictly piscivorous waterbirds nesting in the Columbia River estuary (i.e., terns and cormorants), but our findings also revealed greater impacts of a generalist predator species (i.e., California gulls) than were previously documented. Approaches used in this study allow for direct comparisons among multiple fish mortality factors and considerably improve the reliability of tag recovery models for estimating predation probabilities in multiple‐predator systems.Received August 1, 2014; accepted November 11, 2014