Comparison and Assessment of Aerial and Ground Estimates of Waterbird Colonies

Abstract: Aerial surveys are often used to quantify sizes of waterbird colonies; however, these surveys would benefit from a better understanding of associated biases. We compared estimates of breeding pairs of waterbirds, in colonies across southern Louisiana, USA, made from the ground, fixed-wing aircraft, and a helicopter. We used a marked-subsample method for ground-counting colonies to obtain estimates of error and visibility bias. We made comparisons over 2 sampling periods: 1) surveys conducted on the same colonies using all 3 methods during 3–11 May 2005 and 2) an expanded fixed-wing and ground-survey comparison conducted over 4 periods (May and Jun, 2004–2005). Estimates from fixed-wing aircraft were approximately 65% higher than those from ground counts for overall estimated number of breeding pairs and for both dark and white-plumaged species. The coefficient of determination between estimates based on ground and fixed-wing aircraft was ≤0.40 for most species, and based on the assumption that estimates from the ground were closer to the true count, fixed-wing aerial surveys appeared to overestimate numbers of nesting birds of some species; this bias often increased with the size of the colony. Unlike estimates from fixed-wing aircraft, numbers of nesting pairs made from ground and helicopter surveys were very similar for all species we observed. Ground counts by one observer resulted in underestimated number of breeding pairs by 20% on average. The marked-subsample method provided an estimate of the number of missed nests as well as an estimate of precision. These estimates represent a major advantage of marked-subsample ground counts over aerial methods; however, ground counts are difficult in large or remote colonies. Helicopter surveys and ground counts provide less biased, more precise estimates of breeding pairs than do surveys made from fixed-wing aircraft. We recommend managers employ ground counts using double observers for surveying waterbird colonies when feasible. Fixed-wing aerial surveys may be suitable to determine colony activity and composition of common waterbird species. The most appropriate combination of survey approaches will be based on the need for precise and unbiased estimates, balanced with financial and logistical constraints. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):697–706; 2008)     

Recruitment failure in Florida Keys <Emphasis Type="Italic">Acropora palmata,</Emphasis> a threatened Caribbean coral

Recovery of Acropora palmata from its currently imperiled status depends on recruitment, a process which is poorly documented in existing Caribbean coral population studies. A. palmata is thought to be well adapted to proliferate through the recruitment of fragments resulting from physical disturbances, such as moderate intensity hurricanes. This study monitored fifteen 150 m² fixed study plots on the upper Florida Keys fore-reef for asexual and sexual recruitment from 2004 to 2007. Between July and October 2005, 4 hurricanes passed by the Florida Keys, producing wind speeds on the reef tract of 23 to 33 m s⁻¹. Surveys following the hurricanes documented an average loss of 52% estimated live tissue area within the study plots. The percentage of “branching” colonies in the population decreased from 67% to 42% while “remnant” colonies (isolated patches of tissue on standing skeleton) increased from 11% to 27%. Although some detached branches remained as loose fragments, more than 70% of the 380 fragments observed in the study plots were dead or rapidly losing tissue 3 weeks after Hurricane Dennis. Over the course of the study, only 27 fragments became attached to the substrate to form successful asexual recruits. Meanwhile, of the 18 new, small encrusting colonies that were observed in the study, only 2 were not attributable to asexual origin (i.e., remnant tissue from colonies or fragments previously observed) and are therefore possible sexual recruits. In summary, the 2005 hurricane season resulted in substantial loss of A. palmata from the upper Florida Keys fore-reef from a combination of physical removal and subsequent disease-like tissue mortality, and yielded few recruits of either sexual or asexual origin. Furthermore, the asexual and sexual fecundity of the remaining population is compromised for the near future due to the lack of branches (i.e., “asexual fecundity”) and overall loss of live tissue.     

EDITORIAL

Understanding breeding site preferences is important for management of colonial waterbirds. The number of active colonial waterbird nests at a series of four small constructed wetlands in Cape Town was counted monthly from 1999 to 2008. In total 491 pairs belonging to 11 waterbird species were involved. Between 1997 and 2004 a number of different artificial structures were used to attract colonial waterbirds to breed at one of the constructed wetlands. The changes in use of the different artificial structures, and of natural reedbeds, for nesting revealed the site preferences of different waterbird species. Structures that simulated flooded dead trees were most successful in attracting breeding waterbirds. Reedbeds were used for nesting when the number of such artificial structures was reduced. When the reedbeds were removed some birds returned to the artificial structures, whereas others moved to a more distant wetland. Some species were pioneers that provided public information about safety from predation that apparently encouraged birds in other species to settle. Safety from potential mammalian predators was clearly the prime consideration in breeding locality selection. Adverse winds and human disturbance were less important factors.     

Individual dispersal among colonies of Little Egrets Egretta garzetta

Colonial waterbirds are unusual in that competition for nest-sites or mates may occur at a scale of a few metres, whereas thousands of birds may overlap in their foraging range at a larger scale. Dispersal has been evaluated for only a few such species, and its adaptive significance remains unclear. We studied Little Egret dispersal among all the colonies within the Camargue, southern France. The overall probability of dispersal between successive years was 0.45. The probability of dispersal was unaffected by a bird's age, or by any density-dependent effect of colony size. Juveniles dispersed at distances that would be expected if colony selection were random, while adults tended to remain within 10 km of their previous colony. We found no obvious environmental 'trigger' for an individual to disperse. Although our evidence is inconclusive, the short dispersal distances of adults are not consistent with foraging conditions as the primary trigger for dispersal. Little Egrets generally forage within 8 km of their colony, so birds dispersing less than 10 km would gain little advantage in response to unfavourable foraging conditions. Our data, with 75% of dispersing birds coming from decreasing colonies and 72% joining increasing colonies, suggest that individual dispersal depended on colony dynamics as a whole, i.e. (1) a social component of dispersal at the individual level, or (2) a simultaneous colony response to unfavourable environmental conditions or (3) both. Further investigation at a higher social level may be necessary to understand dispersal of this colonial nesting species.     

THE PROCESS OF COLONY FORMATION AMONG HERRING GULLS LARUS ARGENTATUS NESTING IN NEW JERSEY

We examined the pattern of colony occupation and egg-laying in five colonies of Herring Gulls nesting in New Jersey, U.S.A. Colonies formed from epicentres located in sparse bushes. The number of epicentres related to the number of birds nesting in the colonies. Colonies of over 250 pairs had more than one epicentre, whereas those with under 250 pairs had only one epicentre. Epicentres were not always in the geographical centre of the colonies. New territory-hunting pairs filled in the epicentre areas, and then nested outside these areas. The egg-laying pattern followed the settling pattern, but was more synchronous than the settling pattern. There was greater synchrony of egg-laying within sub-areas of the colonies than in the colonies as a whole. Further, synchrony correlated with the number of nests in sub-areas.     

A STUDY OF THE CINNAMON-BREASTED ROCK BUNTING Fringillaria tahapisi tehepisi

Brown, C. J. &; Piper, S. E. 1988. Status of Cape Vultures in the Natal Drakensberg and their cliff site selection. Ostrich 59:126-136.

Ground, aerial and questionnaire surveys on the status of the Cape Vulture Gyps coprotheres in the Little and High Drakensberg mountain ranges of Natal were carried out from 1981–1983. The area supported at least 1325 Cape Vultures, 60% on the High Drakensberg in 17 nesting colonies (mean of 31 birds per colony) and 38 roosts (mean of 7 birds). In the Little Drakensberg 17 sites were found, six of which were confirmed nesting colonies (mean of 53 birds) and seven were roosts (mean of 16 birds). A minimum of 215 nests was recorded while the actual number of breeding pairs was probably about 325. In the Natal Drakensberg 84% of nests and 77% of roosting birds occupied ledges that faced between east and south, on the lee side of the generally westerly and northwesterly “Bergwind”. Nesting ledges in the High Drakensberg were on average about 100 m lower than roosting ledges. The northern Drakensberg supported the largest numbers of Cape Vultures (19 birds per 10 km of cliffs), while the smallest numbers were recorded in the south (8 birds per 10 km of cliffs). In the south smallstock farming predominates and the availability of carrion is higher than in the north where mainly largestock are farmed. The use of poisons (mainly strychnine) for killing mammalian predators on farms is considered to be the main reason for fewer birds occurring in the south, and is the main threat to Cape Vultures in the Natal Drakensberg.     

Breeding displays of the African Spoonbill Platalea alba

Breeding colonies of the African Spoonbill Platalea alba were studied in Kenya and South Africa during 1980. In Kenya nesting was in trees and in South Africa in reedbeds; the nesting area in both colonies was shared with other species of waterbirds.
During the breeding season, adult African Spoonbills acquire a more immaculate plumage and brighter soft-part coloration. Sexes are distinguishable by the larger size and more robust proportions of males.
Males are more sedentary and aggressive during pair formation, and females are more mobile and submissive. The general pattern of pair formation is described, with a variety of ritualized courtship displays acting to bring and keep the sexes together.
Comfort movements, vocalizations, and 14 distinct ritualized social displays are described and illustrated for the African Spoonbill. Some similarities and possible homologies with displays of other ciconiiform birds are mentioned. This paper represents the first part of a continuing study of the comparative ethology of the five species of spoonbills of the world.     

Climate- and resource-driven long-term changes in dormice populations negatively affect hole-nesting songbirds

Global climate change has been shown to cause variable shifts in phenology in a variety of animals and unexpected outcomes across food chains are to be found. Here we examined how rising annual spring temperatures affected the interactions between seed masting, cavity nesting birds and dormice using long-term data from Eastern Czech Republic. We have shown that climate change was associated with unequal shifts in the phenology of two cavity-breeding groups: dormice and birds. Rising spring temperatures have progressively advanced the termination of hibernation for the edible dormouse Glis glis , a common bird predator, leading to an increasing overlap in the use of nesting boxes between dormice and birds. In contrast, only the collared flycatcher Ficedula albicollis , of the four cavity-nesting bird species, advanced its breeding dates in response to rising temperatures. At the same time, favourable weather conditions, coupled with good seed masting years, have been associated with a substantial rise in dormice numbers. Concurrent with the increasing dormice abundance, the number of bird nests destroyed significantly increased in three out of four bird species. We showed that while there was a significant change in the date that the dormice emerged from hibernation during the course of the study, it did not significantly contribute to predation levels when controlling for their abundance and timing of breeding in birds. We found that the increasing dormice abundance was the main factor causing high brood losses in birds, while the timing of breeding in birds had a variable effect between bird species. This study illustrates how changes in climate might affect organisms at various trophic levels with often unexpected outcomes. Limited evidence from other study organisms suggests that species most at risk are those at different trophic levels that do not shift at the same rate or in the same direction as their food resources, predators or competitors.     

A Large Accumulation of Avian Eggs from the Late Cretaceous of Patagonia (Argentina) Reveals a Novel Nesting Strategy in Mesozoic Birds

We report the first evidence for a nesting colony of Mesozoic birds on Gondwana: a fossil accumulation in Late Cretaceous rocks mapped and collected from within the campus of the National University of Comahue, Neuquén City, Patagonia (Argentina). Here, Cretaceous ornithothoracine birds, almost certainly Enanthiornithes, nested in an arid, shallow basinal environment among sand dunes close to an ephemeral water-course. We mapped and collected 65 complete, near-complete, and broken eggs across an area of more than 55 m². These eggs were laid either singly, or occasionally in pairs, onto a sandy substrate. All eggs were found apparently in, or close to, their original nest site; they all occur within the same bedding plane and may represent the product of a single nesting season or a short series of nesting attempts. Although there is no evidence for nesting structures, all but one of the Comahue eggs were half-buried upright in the sand with their pointed end downwards, a position that would have exposed the pole containing the air cell and precluded egg turning. This egg position is not seen in living birds, with the exception of the basal galliform megapodes who place their eggs within mounds of vegetation or burrows. This accumulation reveals a novel nesting behaviour in Mesozoic Aves that was perhaps shared with the non-avian and phylogenetically more basal troodontid theropods.     

Constrained by available raptor hosts and islands: density-dependent reproductive success in red-breasted geese

In this paper we aim to explain the distribution of red-breasted geese Branta ruficollis over different nesting habitats. To be safe from land predators red-breasted goose colonies were restricted to i) islands on rivers, ii) cliffs with peregrine falcons Falco peregrinus , and iii) the close proximity of snowy owl Nyctea scandiaca and rough-legged buzzard Buteo lagopus nests. Among years nest site availability varied by fluctuations in numbers of owls and buzzards in association with cycles in lemming abundance, but the total number of goose nests found in the study area did not vary. The distribution of geese, in combination with data on reproductive success, suggested a despotic mechanism: at cliffs, goose numbers were constant among years with an invariably high reproductive success, whereas large fluctuations in numbers on islands coincided with opposite trends in success. Apparently, geese nesting with owls or buzzards moved to the few islands present in the study area during years when these birds of prey were absent. Consequently, in such years the average density of geese on islands was more than twice as high as at cliff colonies (5.4 and 2.3 pairs per ha of foraging habitat, respectively). Colony size at cliffs may have been restricted by territorial behaviour of the geese, though there is evidence that, additionally, the host falcons also limited the number of nesting geese. Apparently rare in closely related species, we observed a negative density-dependent effect on reproductive success during the nest phase, and attribute this to limited food resources, reinforced by the high frequency of territorial interactions. This leads to the conclusion that, in addition to predation pressure, nesting density is an important agent in the link between lemming cycles and goose breeding success.     

Testosterone and group size in cliff swallows: testing the "challenge hypothesis" in a colonial bird

The "challenge hypothesis" states that increases in testosterone levels of male animals during the breeding season are directly related to the extent of intrasexual competition for resources or mates that they experience. Although often tested in territorial species, the challenge hypothesis has not been evaluated for colonial animals that live in groups of different sizes and that thus experience different intensities of intrasexual competition. We measured circulating testosterone levels of male and female cliff swallows (Petrochelidon pyrrhonota) in southwestern Nebraska, where these birds nest in colonies of widely different sizes. Males had significantly higher testosterone levels than females, as expected. For males especially, there was a seasonal rise in testosterone levels early in the nesting cycle, corresponding to the period when birds were establishing nest ownership and egg laying, and then a fall as they switched to parental duties. Testosterone levels varied significantly with colony size; for both sexes, birds in larger colonies had higher levels of testosterone than those in smaller colonies when controlling for date. Age and body mass were not related to testosterone levels. Higher levels of testosterone for birds of both sexes in larger colonies probably reflect greater competition for matings, often extra pair, in the more social nesting situations. The results support the predictions of the challenge hypothesis.     

Panmixia on a continental scale in a widely distributed colonial waterbird

Many highly mobile species, such as migratory birds, can move and disperse over long distances, yet exhibit high levels of population genetic structuring. Although movement capabilities may enable dispersal, gene flow may be restricted by behavioural constraints such as philopatry. In the present study, we examined patterns of genetic differentiation across the range of a highly mobile, colonial waterbird. American white pelicans (Pelecanus erythrorhynchos) breed across continental North America and are currently experiencing a range expansion, especially on the eastern range limit. To assess patterns of genetic structuring, we sampled 333 individuals from 19 colonies across their North American range. The use of ten variable microsatellite markers revealed high levels of allelic richness with no population differentiation. Both Bayesian and frequentist approaches to examining genetic structuring revealed a single panmictic population. We found no evidence of genetic structuring across the Continental Divide or between migratory and non‐migratory colonies. The lack of any genetic structure across the range indicates that, unlike other waterbirds with similar life‐history characteristics, extensive gene flow and presumably low philopatry appear to preclude genetic differentiation. The lack of population genetic structure in American white pelicans provides an example of range‐wide panmixia, a rare phenomenon in any terrestrial species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 583–592.     

Nest spacing in relation to settlement time in colonial cliff swallows

How colonial animals space their nests in relation to conspecifics may provide clues as to whether coloniality provides net benefits or occurs only because breeding sites are limited. We examined how nearest-neighbour distance varied in relation to settlement time in the highly colonial cliff swallow, Petrochelidon pyrrhonota, comparing observed nearest-neighbour distances to those expected if birds spread out to maximize nest spacing. Cliff swallows generally settled closer to each other than required by the available substrate, and clustered their nests closer in large colonies than in small ones. The first settlers at a colony site spaced themselves further apart than later arrivals but did not maximize nearest-neighbour distances. The first arrivals maintained greater nest spacing throughout the season than did birds that arrived later. Colony size and amount of nesting substrate had no effect on initial settlement distances of the first arrivals, but eventual nearest-neighbour distances declined with colony size. First arrivals may gain less from nesting with conspecifics and thus are less likely to cluster their nests than later arrivals, which may often be young or na?ve birds that gain more from the social benefits of colonial nesting. The results are consistent with the presumed social advantages cliff swallows receive from coloniality and do not support the hypothesis that colonies result from nesting site limitation. Copyright 2000 The Association for the Study of Animal Behaviour.     

Using flight initiation distance to evaluate responses of colonial‐nesting Great Egrets to the approach of an unmanned aerial vehicle

Remote‐controlled, unmanned aerial vehicles (UAVs) can be used to collect information in difficult‐to‐access places while potentially minimizing human disturbance. These devices have been widely used in a variety of ecological and behavioral studies in recent years, but additional study is needed to assess the magnitude of disturbance they cause to birds. We examined the responses of Great Egrets (Ardea alba) to a UAV in a breeding colony in Louisiana in 2016 where isolated patches of common reed (Phragmites australis) were used as nest sites by multiple breeding pairs. We examined the flush responses and flight initiation distances (FIDs) of nesting adults to the direct vertical approach of a UAV. Incubating adults were more likely to flush from nests and flushed at greater distances when approached by a UAV than adults with nestlings, suggesting that adult assessment of risk was based on the greater reproductive value of nestlings. We observed fewer flush responses and calculated lower set‐back distances using a UAV to approach nesting Great Egrets (~50 m) than set‐back distances calculated using traditional methods of approach (e.g., walking or boating; 87–251 m). We found that FIDs were shorter when more adults were present in nesting patches, suggesting that the perception of predation risk may be based in part on the reactions of other birds. Our results suggest that UAVs may be a useful alternative for monitoring colonial‐nesting waterbirds. However, our analyses were based exclusively on behavioral observations. Additional studies of the physiological responses of birds to the approach of UAVs are needed to better understand the stress responses of birds to these devices.     

THE BIOLOGY OF THE WAVED ALBATROSS DIOMEDEA IRRORATA OF HOOD ISLAND, GALAPAGOS

As a nesting species, the Waved Albatross Diomedea irrorata is restricted to Hood Island in the Galapagos archipelago where 12,000 pairs bred in 1971. Outside the islands the species occurs over the northern parts of the Humboldt Current. Two colonies were studied in detail (1970–1971). At the start of a season, males returned first to the colonies and defended a small territory. Copulation occurred without any elaborate ceremony and the female spent little time on land before laying. There was no fixed nest-site, even within a season, and birds moved their eggs considerable distances. This resulted in heavy egg losses. Younger birds bred later than older birds and laid longer but narrower eggs. The average incubation spell varied from four to five days at the extremes of the incubation period to 19 days in the middle. The average incubation and fledging periods were 60 and 167 days respectively. Pairs which lost an egg sometimes adopted the abandoned egg of another bird and successfully reared the chick. Most pairs nested in both seasons. Nesting success was extremely variable, both between years and between colonies. Between 1961 and 1971 at Punta Suarez, virtually no young were reared in four seasons. Even in 1970–71, where nesting success was good, some groups of birds deserted their eggs en masse whereas in neighbouring areas up to 80% of the pairs reared young. The main foods of the young were squid and fish. Birds did not moult wing and tail feathers at the breeding colonies, and about 50% retained some primaries for more than one season, suggesting that successful pairs had difficulty in fitting in a complete moult between breeding attempts. Old feathers were normally found among the inner primaries and at the next moult were preferentially replaced, though adjacent newer feathers were sometimes retained for another season. Some birds bred in their fourth years, but most not until a year or two older. Immatures were present at the colonies late in the breeding cycle, the youngest returning latest and remaining until the last young fledged. Survival of adults and young averaged at least 95% and 93% per annum over many years. Adults and young ringed in 1961 survived equally well. The significance of the timing of the return of immatures and of the large-scale desertion of eggs, apparently not due to food shortage or disturbance, is discussed.     

Estimating the abundance and hatching success of breeding Curlew Numenius arquata using survey data

Counts of individual Curlew and estimates of breeding pairs from ‘field-by-field’ surveys were highly correlated with the numbers of nesting pairs, as determined by intensive studies on 27 sites located on four grassland-dominated study areas. The mean count of individual Curlew over three standard survey visits to each site was used to estimate numbers of nesting pairs. This estimate exceeded the number of nesting pairs on all study areas (as assessed by intensive studies) by 12% but was more accurate than the maximum number of pairs estimated from the three survey visits on each site (the previous convention for estimating breeding pairs). Estimates of the number of pairs hatching chicks were assessed by recording alarm-calling Curlew during late survey visits. The maximum number of alarm-calling pairs was used to estimate the number of pairs hatching chicks, overestimating this by 54%. Three of the study sites were adjacent to extensive moorland which produced overestimates during surveys because moorland nesting birds fed and led broods onto these sites. Omitting these sites from consideration reduced the degree of overestimation to 1% for the number of pairs and 7% for the number of pairs hatching chicks.     

The current state of populations of the common tern Sterna hirundo (Linnaeus, 1758) and the Kamchatka tern S. camtschatica (Pallas, 1811) in northern Sakhalin

This paper presents data on the numbers of nesting common and Kamchatka tern populations on the northeastern coast of Sakhalin Island over a more than 20-year period. Distribution maps of the main tern colonies (Chaika, Lyarvo, and Wrangel islands, and an unnamed island situated in Chaivo Bay) are given. Taking into account the current data concerning southern Sakhalin colonies, the total number of terns in Sakhalin Island is about 9000 pairs for the Kamchatka tern and up to 16000 pairs for the common tern. The main biotic and abiotic factors that determine the population dynamics and distribution of nesting terns in northern Sakhalin Island are enumerated.     

Multi-scale habitat models for reintroduced bird populations: a case study of South Island saddlebacks on Motuara Island

Understanding resource selection by animals is important when considering habitat suitability at proposed release sites within threatened species recovery programmes. Multi-scale investigatory approaches are increasingly encouraged, as the patchy distribution of suitable habitats in fragmented landscapes often determines species presence and survival. Habitat models applied to a threatened New Zealand forest passerine, the South Island saddleback (Philesturnus carunculatus carunculatus), reintroduced to Ulva Island (Stewart Island) found that at landscape scale breeding pairs? preferences for sites near the coast were driven by micro-scale vegetation structure. We tested these results by examining models of breeding site selection by a reintroduced saddleback population on Motuara Island (Marlborough Sounds) at two scales: (1) micro-scale, for habitat characteristics that may drive breeding site selection, and (2) landscape scale, for variations in micro-scale habitat characteristics that may influence site colonisation in breeding pairs. Results indicated that birds on Motuara Island responded similarly to those on Ulva Island, i.e. birds primarily settled at the margins of coastal scrub and forest and later cohorts moved into larger stands of coastal forest where they established breeding territories. Plant species composition was also important in providing breeding saddleback pairs with adequate food supply and nesting support. However, Motuara Island birds differed in their partitioning of habitat use: preferred habitats were used for nesting while birds were foraging outside territorial boundaries or in shared sites. These differences may be explained because Motuara has a more homogeneous distribution of microscale habitats throughout the landscape and a highly bird-populated environment. These results show that resource distribution and abundance across the landscape needs to be accounted for in the modelling of density?bird?habitat relationships. In the search for future release sites, food (invertebrates and fruiting tree species) should be abundant close to available nesting sites, or evenly spread and available throughout the landscape.     

Central–periphery gradient of individual quality within a colony of Black‐headed Gulls

Central nesting sites within avian colonies are often more profitable in terms of fitness, as they can offer better protection against predators compared with colony edges. Thus, central sites are expected to attract high‐quality individuals, which should produce a clear central–periphery gradient in the phenotypic quality of nesting individuals and reproductive output within colonies. The aim of this study was to assess spatial patterns of individual quality within a colony of a common larid, the Black‐headed Gull Chroicocephalus ridibundus. We found that laying was advanced in the central zone of the colony when compared with the periphery, and central pairs tended to lay larger eggs and had higher hatching success than peripheral pairs. Centrally nesting individuals had higher blood haemoglobin concentrations, size‐corrected body mass and heterozygosity (males only), indicating their better condition and higher genetic quality. In contrast, central females had lower plasma albumin concentration, which could possibly be attributed to their greater investment in egg production. Finally, pairs breeding in the central zone of the colony had an elevated heterophil/lymphocyte ratio, suggesting that social stress can be recognized as an important cost of nesting in dense aggregations. The results provide empirical support for central–periphery gradients in the individual quality of colonially nesting Black‐headed Gulls, indicating that spatial gradients in pair quality should be carefully taken into account when conducting research on colonial birds.     

Nest and breeding population abundance of Least Terns: assessing bias and variation in survey timing and methods

Methods commonly used to estimate the number of nests and size of the breeding population at colonies of Least Terns (Sternula antillarum) and other waterbirds include walk‐through counts of nests (ground‐nest counts) and counts of incubating adults from the colony perimeter (incubating‐adult counts). The bias and variance of different methods and the comparability of repeated surveys versus once‐annual censuses are poorly understood. Our objectives were to assess (1) the potential bias and variation of the more rapid incubating‐adult counts compared to the time‐intensive, and presumably more accurate, ground‐nest counts, and (2) how accurately a once‐annual census captured peak nesting abundance. We studied nine Least Tern colonies at Cape Lookout National Seashore (CALO), North Carolina, from April to August 2010–2012. We analyzed observer and survey method agreement with concordance correlation coefficients (ρ_c). We deployed time‐lapse cameras at 156 nests and used repeated‐measures logistic regression to determine if the proportion of time spent incubating varied with colony, time of day, or time of season. We found substantial agreement in abundance estimates of Least Tern nests and incubating adults between observers and survey methods, and among different times of day and seasons (all comparisons ρ_c > 0.97). Least Terns incubated eggs 94% of the time on average during daylight hours, irrespective of colony, nesting stage, or month. Although the nesting peak at CALO occurred during the recommended census period for Least Terns, abundance estimates for surveys conducted at different times during that period varied by as much as 39%. We recommend conducting incubating‐adult counts to estimate nest and breeding population abundance of Least Terns or other waterbirds when vegetation or dunes do not obstruct views of nesting colonies. In addition, given the variation in abundance estimates for surveys conducted at different times during the recommended survey period, incubating‐adult counts should be performed at least twice during the census period, with the maximum count reported as peak nest abundance.