Incubation in uni-parental Arctic breeding waders

Bruinzeel, L., Tulp, I., Jukema, J. & Stepanova, O. 2000. Incubation in mi-parental arctic breeding waders. Ostrich 71 (1 & 2): 345.

Little Stints Calidris minuta are amongst the smallest breeding waders. This, in combination with the fact that they have a peculiar breeding system, in which each adult takes care of her/his own nest and young, puts high demands during incubation. Due to uni-parental feeding system, feeding time is limited. The accumulation of he1 stores prior to breeding may be crucial to get through the 21-day incubation period. Incubation rhythms, body mass dynamics and energy expenditure of Little Stints was studied at Medusa Bay, Western Taimyr in the summer of 1996. Repeated body mass measurements during incubation were taken by catching the birds on the nest. Concurrently, we measured nest attentiveness, using Tiny Talk dataloggers and temperature sensitive probes, which were placed in the nests. Arthropod availability was measured to estimate food availability. In comparison, body mass dynamics were also studied in Curlew Sandpiper, another uni-parental breeder, but three times bigger. The first (preliminary) results of this study were presented.     

Phenotypic variation in the breeding phenology of the woodlouse Armadillidium vulgare

Summary The breeding phenology of temperate wood-lice is strongly seasonal, the result of physiological constraints and precise environmental cues for reproduction. The adaptive value of such mechanisms is that the release of offspring coincides with favourable conditions for growth and survival (Willows 1984). We recorded the breeding phenology of Armadillidium vulgare (Latreille) on two grassland sites in Great Britain and found between-site and between-year variation in the onset of reproduction, the duration of reproductive activity, the release of offspring, the size of reproductive females and the number of broods per female. Between 82.7 and 97.7% of gravid females sampled were semelparous at 23 months, with the remainder iteroparous, producing a second brood after 35 months. On one site (Weeting Health) improved growth conditions during 1984 allowed some females (19.3% of gravid females sampled in that year) to produce a brood after 11 months. There was also an increase in the number of 3-year-old females found to be gravid. An experimental manipulation of the same habitat confirmed that such changes in life history tactics could be phenotypic responses. The observed phenotypic variation was sufficient to produce a range of life history tactics within a population. Mixtures of life history tactics within a population may be typical of invasive species and populations at the edge of the species range. Our results support the idea that phenotypic plasticity can be an appropriate tactic to maximise fitness in a fluctuating environment (Caswell 1983, 1989).     

Loss of periodicity in breeding success of waders links to changes in lemming cycles in Arctic ecosystems

Lemming population cycles in the Arctic have an important impact on the Arctic food web, indirectly also affecting breeding success in Arctic‐nesting birds through shared predators. Over the last two decades lemming cycles have changed in amplitude and even disappeared in parts of the Arctic. To examine the large scale effect of these recent changes we re‐analysed published data from the East Atlantic Flyway (EAF), where a relationship between lemming cycles and wader breeding success was earlier found, and new data on breeding success of waders in the East Asian–Australasian Flyway (EAAF). We found that 1) any long‐term periodicities in wader breeding success existed only until the year 2000 in the EAAF and until the 1980s in the EAF; 2) studying these patterns at a smaller spatial scale, where the Siberian–Alaskan breeding grounds were divided into five geographical units largely based on landscape features, breeding success of waders from the EAAF was not correlated to an index of predation pressure, but positively correlated to Arctic summer temperatures in some species. We argue that fading out of lemming cycles in some parts of the Arctic is responsible for faltering periodicity in wader breeding success along both flyways. These changed conditions have not yet resulted in any marked changing trends in breeding success across years, and declining numbers of waders along the EAAF are therefore more likely a result of changing conditions at stop‐over and wintering sites.     

The effect of conservation interventions on the abundance of breeding waders within nature reserves in the United Kingdom

Breeding populations of many wading birds have declined globally, primarily caused by habitat degradation and loss. In the UK, population declines have been particularly notable on lowland wet grasslands. In response, some areas of lowland wet grassland have been restored and are under ongoing management to improve the breeding conditions of target species. Here, we assess the efficacy of management measures using a Bayesian framework and controlling for confounding factors. We focus on four wader species, Northern Lapwing Vanellus vanellus, Eurasian Curlew Numenius arquata, Common Snipe Gallinago gallinago and Common Redshank Tringa totanus, that breed in numbers on wet grassland reserve sites in the UK. We collated annual site-specific climate variables, management information (e.g. the creation of wet features and predator control measures) and bird counts between 1994 and 2018. We found the effects of conservation actions varied between intervention types and species. For Lapwing and Redshank, excluding predators by predator-exclusion fencing, especially in combination with fox control, was generally associated with higher breeding counts. For all study species, sites with longer histories of management were associated with higher breeding numbers, with the effect of site age being particularly notable for management on former arable land. Our findings support the effectiveness of targeted conservation actions to achieve high numbers of breeding waders on lowland wet grassland reserves, and also highlight the value of consistent and reliable monitoring data.     

Overland movement and migration phenology in relation to breeding of Arctic Terns Sterna paradisaea

It is often implicitly assumed that seabirds migrate using marine environments, but this assumption is increasingly being challenged by electronic tracking data. The arrival and departure routes of Arctic Terns breeding on the North Sea coast of the United Kingdom (UK) are unknown but there has been speculation about the possibility of overland migration. Analysis of light-level geolocator data from birds breeding on the Farne Islands suggests that these birds arrived and left their North Sea colony overland via the Irish Sea, rather than taking coastal routes along the east coast of the UK and through the English Channel. In addition, some departing birds may enter the North Atlantic by crossing Ireland rather than through the Irish Sea. The direction of arrival in spring had a more southerly orientation than the direction of autumn departure. The geolocator data allow migration phenology in relation to breeding to be defined and indicated that the birds arrived around 15 days before the first eggs were laid in the colony. Departure timing may be determined by seasonal progression and not markedly influenced by breeding success. This study supports the idea that overland migration may be a more widespread and consistent strategy for seabirds than has been realized.     

Subcolony variation in phenology and breeding parameters in little auk Alle alle

In seabirds, variation in breeding phenology and success is often induced by macro- and mesoscale fluctuations in oceanographic conditions, which is recordable at the regional or colony scale. Variation in breeding parameters may also exist at the microscale—among discrete breeding aggregations and subcolonies. The aim of this study was to compare breeding phenology and parameters (hatching success, chick survival, chick body mass indices) among little auks (Alle alle) nesting in three subcolonies differing in their microtopographic and microclimatic features, situated in Magdalenefjorden (north-western Spitsbergen). The hatching phenology varied significantly among the subcolonies. This was probably due to the different duration of snow persistence in spring, as nests are occupied as soon as the snow cover melts sufficiently to allow access to them. The earliest hatching was recorded in the subcolony located on steep slopes at low altitude in the vicinity of the fjord, which favours early ice and snow melting in spring. Hatching success differed significantly among subcolonies, which could also have been due to the microclimatic features of the subcolonies. Hatching success was the lowest in the subcolony where birds started to breed while patchy snow was still persisting. Water from the melting snow could have flooded some of the nests. Alternatively, the low hatchability could have been caused by a higher frequency of less experienced breeders (e.g., first-time breeders) among the individuals nesting in this subcolony. Other breeding parameters (chick survival and growth) were generally similar in all the subcolonies.     

Mating system and timing of breeding in Holarctic waders

In passerine birds polygamous species lay eggs later than monogamous species. Yom-Tov has hypothesized that this is because polygamous males do not provision their mates with food, thereby delaying egg-laying. An indirect prediction of this hypothesis is that there should be no difference in laving dates between monogamous and polygamous birds in which mate provisioning does not occur. This prediction was tested by examining egg-laying dates in sympatric wader species that have contrasting mating systems but in which food provisioning of mates does not occur. Polygamous waders laid eggs later than monogamous waders, however, both in a comparison involving only calidridine sandpipers and in a comparison involving sandpipers and plovers. Accounting for phylogeny did not affect the direction of the results, which therefore cast doubt on the validity of the 'mate provisioning' hypothesis and indicated that features common to both passerines and waders were more likely to explain the differences in egg-laying dates between mating systems. Mating system and parental care system were strongly related, with all polygamous species being uniparental and monogamous species being biparental. It is suggested that seasonally early food supplies are of relatively low abundance, constraining species to biparental care and monogamy. By contrast, seasonally late food supplies are more abundant, making uniparental care more successful, thereby creating greater opportunities for mate desertion and polygamy.     

Diurnal variability in the detection of waders on their breeding grounds

When censusing breeding waders in large upland areas, the time of day has a major influence on the numbers of birds detected.     

Mismatches between breeding phenology and resource abundance of resident alpine ptarmigan negatively affect chick survival

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Testing the effects of recreational disturbance on two upland breeding waders

Previous work has shown that Golden Plovers Pluvialis apricaria guarding chicks utilize heavily disturbed habitat at a lower rate than surrounding areas, but that such rates of avoidance are reduced when the movement of people is restricted to a surfaced footpath. However, there remained some unanswered questions, which are addressed in this paper. First, we examine to what extent habitat avoidance is dependent upon visitor numbers, and find no evidence that Golden Plovers avoided disturbed areas at a site where visitor pressure was half that previously studied. Secondly, we examine whether these relationships between habitat occupancy and disturbance apply to other upland waders and find that Dunlin Calidris alpina habitat utilization in disturbed areas showed a non-significant increase of approximately 50% following the provision of a surfaced footpath, in a manner similar to that observed for Golden Plover. Thirdly, we examine whether the large numbers of visitors (120 per weekend day) using the surfaced footpath impact on Golden Plover breeding success, despite the lack of habitat avoidance. There was no evidence that nest location, clutch survival or chick growth rates were reduced close to the footpath. Together, these results suggest that high levels of disturbance can impact upon habitat usage by upland waders, but only in limited circumstances where visitor pressure is very high (greater than at least 30 visitors per weekend day). However, access to such areas can be permitted for large numbers of visitors without impacting upon wader reproductive performance through the provision of a well-surfaced route.     

The impact of raptors on the abundance of upland passerines and waders

The issue of predator limitation of vertebrate prey populations is contentious, particularly when it involves species of economic or conservation value. In this paper, we examine the case of raptor predation on upland passerines and waders in Scotland. We analysed the abundance of five wader and passerine species on an upland sporting estate in southern Scotland during an eight-year period when hen harrier, peregrine and merlin numbers increased due to strict law enforcement. The abundance of meadow pipit and skylark declined significantly during this time. Golden plover also showed a declining trend, whereas curlew increased significantly and there was a near significant increase in lapwings. Contrasting the local population trends of these species with trends on nearby areas revealed higher rates of decline for meadow pipit and skylark at the site where raptors increased, but no differences in trends for any of the three wader species. There was a negative relationship between the number of breeding harriers and meadow pipit abundance the same year and between total annual raptor numbers and meadow pipit abundance. Predation rates of meadow pipit and skylark determined from observations at harrier nests suggested that predation in June was sufficient to remove up to 40% of the June meadow pipit population and up to 34% of the June skylark population. This 'quasi-natural' experiment suggests that harrier predation limited the abundance of their main prey, meadow pipit, and possibly the abundance of skylark. Thus, high densities of harriers may in theory reduce the abundance of the prey species which determine their breeding densities, potentially leading to lower harrier breeding densities in subsequent years. We found no evidence to suggest that raptor predation limited the populations of any of the three wader species. We infer that concerns over the impact of natural densities of hen harriers on vulnerable upland waders are unjustified.     

Spatial variation in breeding phenology at small spatial scales: A stochastic effect of population size

Spatial variation in phenology can occur at small spatial scales over which individuals can disperse or forage within one generation. Previous studies have assumed that variations in phenological peaks are caused by differences in abiotic environmental characteristics. However, environments should generally be similar among local habitats over small spatial scales. When the local population size is small, the phenological peak of the local population should be strongly affected by the variation in timing expressed by individuals. If a regional population consists of small local subpopulations (e.g., a metapopulation), the stochastic processes regulated by population sizes may explain the spatial variation in phenology. In this study, we quantitatively evaluated the extent of the spatial and annual variations in the breeding phenology of the forest green tree frog, Rhacophorus arboreus habiting a small area (<10 km²). The spatial variation in phenological peaks among 25 breeding sites was large over 6 years. This spatial variation was not explained by differences in air temperature or water depth. Randomization tests revealed that a large portion of the spatial variation could be explained by differences in population size, without considering site-specific factors. Annual variations in phenological peaks tended to be greater for smaller populations. These results imply that the stochastic process might have caused the spatial and annual variations in the phenological peaks of R. arboreus observed in the study region. Understanding spatiotemporal variation in phenology determined by stochastic process would be important to better predict interspecific interactions and (meta)population dynamics at small spatial scales.     

Population changes of breeding waders on farmland in relation to agri-environment management

Capsule Population response of breeding waders to agri-environment management varied between management options and species; implementation has been on too small a scale to reverse national population declines.

Aims To test whether numbers of five breeding wader species have shown a more positive response between 1992 and 2005, at sites with appropriate agri-environment management, than at sites that have remained outside such schemes.

Methods Using data from 60 pairs of farmland study areas in Scotland first surveyed in 1992/93, before agri-environment scheme (AES) implementation, and again in 2005, after scheme implementation, we tested at both site and field scales whether changes in the abundance of five breeding wader species were associated with AES management options designed to benefit these species.

Results Changes in breeding wader abundance were more positive on sites in AES, especially for Northern Lapwings Vanellus vanellus and Common Redshanks Tringa totanus, even though management had not been targetted specially at breeding waders on those sites. However, AES management was associated only with modest population increase for Common Redshanks, and a reduction in the magnitude of decline for Northern Lapwings. At the field scale, there was evidence for Northern Lapwings, Common Redshanks and Common Snipe Gallinago gallinago that options which limited grazing and other agricultural activity were associated with more positive outcomes than those which also manipulated water levels.

Conclusions AES management for breeding waders slowed, and in some cases reversed, breeding wader decline at field and farm scales. These benefits were from options that limited grazing and agricultural operations during the breeding season, but not those that also aimed to raise water levels. A possible explanation is that when wetland options are applied to agriculturally marginal fields, grazing reduction or abandonment, and succession to rank vegetation cover then occur over the course of 5-year agreements, with detrimental effects for breeding waders. Verification arrangements need to be robust enough to guard against this. Levels of agri-environment provision in 2005 were too limited and too poorly targeted at remaining key areas for breeding waders to be able to halt or reverse national population declines.     

Carbon sources for lake food webs in the Canadian High Arctic and other regions of Arctic North America

We investigated the role of autochthonous and terrestrial carbon in supporting aquatic food webs in the Canadian High Arctic by determining the diet of the dominant primary consumer, aquatic chironomids. These organisms were studied in fresh waters on 3 islands of the Arctic Archipelago (~74–76°N) including barren polar desert watersheds and a polar oasis with lush meadows. Stomach content analysis of 578 larvae indicated that chironomids primarily ingested diatoms and sediment detritus with little variation among most genera. Carbon and nitrogen stable isotope mixing models applied to 2 lakes indicated that benthic algae contributed 68–95% to chironomid diet at a polar desert site and 70–78% at a polar oasis site. Detritus, originating from either phytoplankton or terrestrial sources, also contributed minor amounts to chironomid diet (0–32%). Radiocarbon measurements for the 2 lakes showed that old terrestrial carbon did not support chironomid production. Carbon stable isotope ratios of chironomids in other High Arctic lakes provided further dietary evidence that was consistent with mixing model results. These findings indicate that, in the Canadian High Arctic, chironomids (and fish that consume them) are supported primarily by benthic algae in both polar desert and oasis lakes. In contrast, our review of carbon flow studies for lakes in other Arctic regions of North America shows that terrestrial carbon and phytoplankton can be important energy sources for consumers. This study provides a baseline to detect future climate-related impacts on carbon pathways in High Arctic lakes.     

Clock gene variation is associated with breeding phenology and maybe under directional selection in the migratory barn swallow

Background

In diverse taxa, photoperiodic responses that cause seasonal physiological and behavioural shifts are controlled by genes, including the vertebrate Clock orthologues, that encode for circadian oscillator mechanisms. While the genetic network behind circadian rhythms is well described, relatively few reports exist of the phenological consequences of and selection on Clock genes in the wild. Here, we investigated variation in breeding phenology in relation to Clock genetic diversity in a long-distance migratory bird, the barn swallow (Hirundo rustica).

Methodology/Principal Findings

In a sample of 922 adult barn swallows from a single population breeding in Italy we found one very common (Q₇) and three rare (Q₅, Q₆, Q₈) length variants of a functionally significant polyglutamine repeat. Rare (2.9%) Q₇/Q₈ heterozygous females, but not males, bred significantly later than common (91.5%) Q₇/Q₇ females, consistent with the expectation that ‘long’ alleles cause late breeding, as observed in a resident population of another bird species. Because breeding date depends on arrival date from migration, present results suggest that the association between breeding date and Clock might be mediated by migration phenology. In addition, fecundity selection appears to be operating against Q₇/Q₈ because late migrating/breeding swallows have fewer clutches per season, and late breeding has additional negative selection effects via reduced offspring longevity. Genotype frequencies varied marginally non-significantly with age, as Q₇/Q₈ frequency showed a 4-fold reduction in old individuals. This result suggests negative viability selection against Q₇/Q₈, possibly mediated by costs of late breeding.

Conclusions/Significance

This is the first study of migratory birds showing an association between breeding phenology and Clock genotype and suggesting that negative selection occurs on a phenologically deviant genotype. Low polymorphism at Clock may constrain microevolutionary phenological response to changing climate, and may thus contribute to the decline of barn swallow populations.     

Phenological variation in annual timing of hibernation and breeding in nearby populations of Arctic ground squirrels

Ecologists need an empirical understanding of physiological and behavioural adjustments that animals can make in response to seasonal and long-term variations in environmental conditions. Because many species experience trade-offs between timing and duration of one seasonal event versus another and because interacting species may also shift phenologies at different rates, it is possible that, in aggregate, phenological shifts could result in mismatches that disrupt ecological communities. We investigated the timing of seasonal events over 14 years in two Arctic ground squirrel populations living 20 km apart in Northern Alaska. At Atigun River, snow melt occurred 27 days earlier and snow cover began 17 days later than at Toolik Lake. This spatial differential was reflected in significant variation in the timing of most seasonal events in ground squirrels living at the two sites. Although reproductive males ended seasonal torpor on the same date at both sites, Atigun males emerged from hibernation 9 days earlier and entered hibernation 5 days later than Toolik males. Atigun females emerged and bred 13 days earlier and entered hibernation 9 days earlier than those at Toolik. We propose that this variation in phenology over a small spatial scale is likely generated by plasticity of physiological mechanisms that may also provide individuals the ability to respond to variation in environmental conditions over time.     

Effects of food abundance and early clutch predation on reproductive timing in a high Arctic shorebird exposed to advancements in arthropod abundance

Climate change may influence the phenology of organisms unequally across trophic levels and thus lead to phenological mismatches between predators and prey. In cases where prey availability peaks before reproducing predators reach maximal prey demand, any negative fitness consequences would selectively favor resynchronization by earlier starts of the reproductive activities of the predators. At a study site in northeast Greenland, over a period of 17 years, the median emergence of the invertebrate prey of Sanderling Calidris alba advanced with 1.27 days per year. Yet, over the same period Sanderling did not advance hatching date. Thus, Sanderlings increasingly hatched after their prey was maximally abundant. Surprisingly, the phenological mismatches did not affect chick growth, but the interaction of the annual width and height of the peak in food abundance did. Chicks grew especially better in years when the food peak was broad. Sanderling clutches were most likely to be depredated early in the season, which should delay reproduction. We propose that high early clutch predation may favor a later reproductive timing. Additionally, our data suggest that in most years food was still abundant after the median date of emergence, which may explain why Sanderlings did not advance breeding along with the advances in arthropod phenology.     

Vegetation structure influences predation rates of early nests in subarctic breeding waders

Ground-nesting species are vulnerable to a wide range of predators and often experience very high levels of nest predation. Strategies to reduce nest vulnerability can include concealing nests in vegetation and/or nesting in locations in which nests and eggs are camouflaged and less easy for predators to locate. These strategies could have important implications for the distribution of ground-nesting species and the success rates of nests in areas with differing vegetation structure. However, the factors influencing the success of nest concealment and camouflage strategies in ground-nesting species are complex. Here we explore the effects of local vegetation structure and extent of nest concealment on nest predation rates in a range of ground-nesting, sympatric wader species with differing nest concealment strategies (open-nest species: Oystercatcher Haematopus ostralegus, Golden Plover Pluvialis apricaria and Whimbrel Numenius phaeopus; concealed-nest species: Black-tailed Godwit Limosa limosa, Redshank Tringa totanus and Snipe Gallinago gallinago) in south Iceland, in landscapes that comprise substantial variability in vegetation structure at a range of scales. We monitored 469 nests of these six wader species in 2015 and 2016 and ~40% of these nests were predated. Nest predation rates were similar for open-nest and concealed-nest species and did not vary with vegetation structure in the surrounding landscape, but nest-concealing species were ~10% more likely to have nests predated when they were poorly concealed, and the frequency of poorly concealed nests was higher in colder conditions at the start of the breeding season. For concealed-nest species, the reduced capacity to hide nests in colder conditions is likely to reflect low rates of vegetation growth in such conditions. The ongoing trend for warmer springs at subarctic latitudes could result in more rapid vegetation growth, with consequent increases in the success rates of early nests of concealed-nest species. Temperature-related effects on nest concealment from predators could thus be an important mechanism through which climate change affecting vegetation could have population-level impacts on breeding birds at higher latitudes.     

Food selection by calanoid copepods in response to between-lake variation in food abundance

1. Food selection experiments were conducted by acclimating calanoid copepods (Eudiaptomus spp.) in suspensions of natural seston and then adding pairs of dual-labelled (¹⁴C/³²P) algae. Each feeding trial measured selectivity between a small, high-quality reference alga, Chlamydomonas reinhardii, and a test alga that differed in size and/or food quality. The influence of food concentration on food selection was tested by using seston from two lakes with contrasting food abundance and by including treatments with filtered lake water ('starved’) and seston diluted with filtered water or enriched with cultured algae. 2. Copepods that had been starved or acclimated to natural seston with low food abundance preferred the larger of two labelled algae, regardless of the nutritional quality of the algae. In agreement with the predictions of an optimal diet model, however, copepods that were acclimated to high food conditions discriminated against low-quality foods, including digestion-resistant algae and dead algae. 3. Selectivity coefficients showed excellent agreement with a previous study involving the same taxa of copepods and labelled algae but in which the copepods had been acclimated to pairs of cultured algae rather than natural seston. Thus, these comparisons emphasize the importance of food availability in modulating copepod selectivity for foods that differ in nutritional quality and suggest that such behaviour occurs in nature.     

Patterns of inter-annual variation in the size asymmetry of growth in Pinus banksiana

A large body of literature suggests that asymmetric competition, where large individuals suppress the growth of smaller individuals by intercepting a disproportionate share of incoming light, is a dominant process in tree population development. This has not been examined extensively for long-lived tree species that accumulate growth over many years under varying growing conditions. Using dendrochronological techniques, we reconstructed annual growth and mortality rates at ten stands of jack pine (Pinus banksiana Lamb.) in Western Canada. We used these data to calculate an annual index of the size asymmetry of growth for each stand for the last 50 years. Jack pine is a shade-intolerant species found in even-aged monoculture stands, so the simple hypothesis is that large trees should consistently perform relatively better than small trees. Inter-annual variation in the index of size-asymmetric growth was positively associated with interannual variation in stand productivity at eight of ten sites. The size asymmetry of growth also showed a positive trend with age at eight of ten sites, even though all sites were in a period of declining leaf area. This should have reduced the intensity of asymmetric competition for light and reduced the size asymmetry of growth over time. Alternate hypotheses for this trend are (1) that physical collisions between crowns result in asymmetric competition for growing space because they are more damaging to small trees, or (2) that a differential build up of diseases in susceptible trees suppresses their growth, even in the absence of competition.