Age‐dependent dispersal and habitat choice in black‐tailed godwits Limosa limosa limosa across a mosaic of traditional and modern grassland habitats

Whether to disperse, and where to, are two of the most prominent decisions in an individual's life, with major consequences for reproductive success. We studied natal and breeding dispersal in the monogamous black‐tailed godwit Limosa limosa limosa in the Netherlands, where they breed in agricultural grasslands. The majority of these grasslands recently changed from wet herb‐rich meadows into well‐drained grassland monocultures, on which godwits have a lower reproductive success. Here we examine habitat selection with a multistate mark–recapture analysis. Habitat transition probabilities between meadows and monocultures were estimated on the basis of 1810 marked chicks and 531 adults during seven years in a 8500 ha study area. Young and adult godwits may differ in habitat selection because: 1) adults may have gained experience from previous nest success where to settle, 2) younger individuals may find it harder to compete for the best territories. Both young and adults moved at a higher rate from the predominant monocultures to meadows than the other way around, thus actively selecting the habitat with better quality. However, dispersal distance of adults was not affected by previous nest success. The average dispersal distance from place of birth of godwits breeding for the first time was ten times larger than that of adult godwits. That godwits breeding in their second calendar year arrived and laid at similar dates and were equally able to select territories in areas with high breeding densities, suggests that young birds were not competitively inferior to adults. Although on monocultures reproduction is insufficient to maintain constant populations, birds sometimes moved from meadows to monocultures. This explains why even after 30 years of land‐use intensification, godwits still breed in low‐quality habitat. The adjustment to changing habitat conditions at the population level appears to be a slow process.     

Intronic variation at the CHD1‐Z gene in Black‐tailed Godwits Limosa limosa limosa: correlations with fitness components revisited

Recently, Schroeder et al. (2010, Ibis 152: 368–377) suggested that intronic variation in the CHD1‐Z gene of Black‐tailed Godwits breeding in southwest Friesland, The Netherlands, correlated with fitness components. Here we re‐examine this surprising result using an expanded dataset (2088 birds sampled from 2004 to 2010 vs. 284 birds from 2004 to 2007). We find that the presence of the Z* allele (9% of the birds) is not associated with breeding habitat type, egg size, adult survival, adult body mass or adult body condition. The results presented here, when used in synergy with the previously reported results by Schroeder et al., suggest that there might be a tendency towards female adults with the Z* allele laying earlier clutches than adult females without the Z* allele. The occurrence of the Z* allele was also associated with a higher chick body mass and return rate. Chicks with the Z* allele that had hatched early in the breeding season were heavier at birth than chicks without the Z* allele and chicks with the Z* allele that had hatched late. Collectively, the results suggest that variation in the CHD1‐Z gene may indeed have arisen as a byproduct of selection acting on females during the egg fase and on chicks during the rearing stages of the reproductive cycle.     

Linking intronic polymorphism on the CHD1‐Z gene with fitness correlates in Black‐tailed Godwits Limosa l. limosa

We show that variation in an intronic length polymorphism in the CHD1‐Z gene in Black‐tailed Godwits Limosa l. limosa is associated with fitness correlates. This is the second example of the CHDZ‐1 gene being correlated with fitness, a previous study having established that Moorhens Gallinula chloropus carrying the rare Z* allele have reduced survival. In Godwits, however, carriers of the Z* allele (374 bp) fared better than those with the more frequent Z allele (378 bp) with respect to body mass, plumage ornamentation, reproductive parameters and habitat quality. The Z* allele was found in 14% of 251 adult birds from nature reserves, but was absent from 33 birds breeding in intensively managed agricultural lands. Males and females with the Z* allele had less extensive breeding plumage, and females had a higher body mass, bred earlier and had larger eggs. There were no significant differences in annual survival between birds with and without the Z* allele. DNA isolated from museum skins demonstrated that this polymorphism was present at low frequency in 1929. We speculate that strong asymmetrical overdominance may explain the low frequency of the Z* allele and that genetic linkage to causal genes might be an explanation for the phenotypic correlations. Our findings suggest a degree of cryptic genetic population structuring in the Dutch Godwit population.     

Carry‐over effects and compensation: late arrival on non‐breeding grounds affects wing moult but not plumage or schedules of departing bar‐tailed godwits Limosa lapponica baueri

In the annual cycle of migratory birds, temporal and energetic constraints can lead to carry‐over effects, in which performance in one life history stage affects later stages. Bar‐tailed godwits Limosa lapponica baueri, which achieve remarkably high pre‐migratory fuel loads, undertake the longest non‐stop migratory flights yet recorded, and breed during brief high‐latitude summers, may be particularly vulnerable to persistent effects of disruptions to their rigidly‐timed annual routines. Using three years of non‐breeding data in New Zealand, we asked how arrival timing after a non‐stop flight from Alaska (>11 000 km) affected an individual godwit's performance in subsequent flight feather moult, contour feather moults, and migratory departure. Late arrival led to later wing moult, but godwits partially compensated for delayed moult initiation by increasing moult rate and decreasing the total duration of moult. Delays in arrival and wing moult up to 34–37 d had no apparent effect on an individual's migratory departure or extent of breeding plumage at departure, both of which were extraordinarily consistent between years. Thus, ‘errors’ in timing early in the non‐breeding season were essentially corrected in New Zealand prior to spring migration. Variation in migration timing also had no apparent effect on an individual's likelihood of returning the following season. The bar‐tailed godwits’ rigid maintenance of plumage and spring migration schedules, coupled with high annual survival, imply a surprising degree of flexibility to address unforeseen circumstances in the annual cycle.     

Adverse effects of agricultural intensification and climate change on breeding habitat quality of Black‐tailed Godwits Limosa l. limosa in the Netherlands

Agricultural intensification is one of the main drivers of farmland bird declines, but effects on birds may be confounded with those of climate change. Here we examine the effects of intensification and climate change on a grassland breeding wader, the Black‐tailed Godwit Limosa l. limosa, in the Netherlands. Population decline has been linked to poor chick survival which, in turn, has been linked to available foraging habitat. Foraging habitat of the nidifugous chicks consists of uncut grasslands that provide cover and arthropod prey. Conservation measures such as agri‐environment schemes aim to increase the availability of chick foraging habitat but have not yet been successful in halting the decline. Field observations show that since the early 1980s, farmers advanced their first seasonal mowing or grazing date by 15 days, whereas Godwits did not advance their hatching date. Ringing data indicate that between 1945 and 1975 hatching dates advanced by about 2 weeks in parallel with the advancement of median mowing dates. Surprisingly, temperature sums at median mowing and hatching dates suggest that while the agricultural advancement before 1980 was largely due to agricultural intensification, after 1980 it was largely due to climate change. Examining arthropod abundance in a range of differently managed grasslands revealed that chick food abundance was little affected but that food accessibility in intensively used tall swards may be problematic for chicks. Our results suggest that, compared with 25 years ago, nowadays (1) a much higher proportion of clutches and chicks are exposed to agricultural activities, (2) there is little foraging habitat left when chicks hatch and (3) because of climate change, the vegetation in the remaining foraging habitat is taller and denser and therefore of lower quality. This indicates that for agri‐environment schemes to make a difference, they should not only be implemented in a larger percentage of the breeding area than the current maxima of 20–30% but they should also include measures that create more open, accessible swards.     

BirdOriTrack: a new video‐tracking program for orientation research with migratory birds

Experimental research on the orientation of migratory songbirds is traditionally conducted using orientation funnels or automatic registration cages that record the directional activity of individual birds on paper or a computer. Most traditionally used funnel and cage designs do not permit investigators to observe detailed behavior of the birds and, therefore, we have gained little insight into the actual behavior of birds while they are exhibiting migratory restlessness and making directional choices. Such behavior can only be studied by direct observation or by video filming. Here, we present BirdOriTrack, a video‐tracking program for extracting time‐resolved, positional data of birds (and potentially other animal species) to determine their orientation relative to the center of a circular cage/funnel. With relatively inexpensive cameras, recording equipment, and cages, orientation experiments can easily be conducted and analyzed using BirdOriTrack. The program is designed to be flexible, allowing analysis of orientation behavior of birds of any size in different cage designs and in both controlled laboratory settings and field‐based studies. To demonstrate the program's utility, we show the results of preliminary field experiments on several species of migratory birds captured at a migration monitoring station. BirdOriTrack is freely available at http://canmove.lu.se/birdoritrack .     

Clock gene polymorphism,migratory behaviour and geographic distribution: a comparative study of trans‐Saharan migratory birds

Migratory behaviour is controlled by endogenous circannual rhythms that are synchronized by external cues, such as photoperiod. Investigations on the genetic basis of circannual rhythmicity in vertebrates have highlighted that variation at candidate ‘circadian clock’ genes may play a major role in regulating photoperiodic responses and timing of life cycle events, such as reproduction and migration. In this comparative study of 23 trans‐Saharan migratory bird species, we investigated the relationships between species‐level genetic variation at two candidate genes, Clock and Adcyap1, and species’ traits related to migration and geographic distribution, including timing of spring migration across the Mediterranean Sea, migration distance and breeding latitude. Consistently with previous evidence showing latitudinal clines in ‘circadian clock’ genotype frequencies, Clock allele size increased with breeding latitude across species. However, early‐ and late‐migrating species had similar Clock allele size. Species migrating over longer distances, showing delayed spring migration and smaller phenotypic variance in spring migration timing, had significantly reduced Clock (but not Adcyap1) gene diversity. Phylogenetic confirmatory path analysis suggested that migration date and distance were the most important variables directly affecting Clock gene diversity. Hence, our study supports the hypothesis that Clock allele size increases poleward as a consequence of adaptation to the photoperiodic regime of the breeding areas. Moreover, we show that long‐distance migration is associated with lower Clock diversity, coherently with strong stabilizing selection acting on timing of life cycle events in long‐distance migratory species, likely resulting from the time constraints imposed by late spring migration.     

Movements and Conflicts in a Flock of Foraging Black‐Tailed Godwits (Limosa limosa): The Influence of Feeding Rates on Behavioural Decisions

We studied movements and conflicts within a small flock of free‐living black‐tailed godwits foraging on benthic invertebrates in a brackish lagoon. To interpret our results in the framework of foraging theory, we studied the influence of individual feeding rate on the decisions to move and to attack flock companions. Birds changed their position within the flock more often when their intake rate was low and sometimes attacked conspecifics to supplant them from their feeding place. Aggressors significantly avoided front attacks and were almost always successful. They attacked individuals having higher feeding rates than themselves and their own feeding rate significantly increased after the attack, although victims were not chased off to particularly poor sites. Our results suggest that aggressors could obtain reliable information about the quality of the foraging site they coveted by observing their victim’s feeding activity before attacking. Although aggression seemed to be caused by a low intake rate, we show that displacing another bird was more time‐consuming than independent foraging. We conclude that it was not the most profitable behaviour in terms of energy intake. Foraging site displacement probably also had social functions, such as reinforcement of social status in a flock of birds preparing for pre‐breeding migration.     

Advances in tracking small migratory birds: a technical review of light‐level geolocation

Light‐level geolocation data loggers, or geologgers, have recently been miniaturized to the extent that they can be deployed on small songbirds, allowing us to determine many previously unknown migration routes, breeding locations, and wintering sites. Use of geologgers on small birds has great potential to help address major research and conservation questions, but the method is not without its shortcomings. Among these shortcomings are the need to recapture birds after they have carried a device throughout a migration cycle and the potential for the devices to affect survival and behavior. We examined return rates of birds with geologgers in published and unpublished studies and found no evidence of a general negative effect of geologgers on survival, although there were a few individual studies where such an effect was evident. From these same studies, we found that most currently used harness materials are equivalent in terms of failure rates, and the most reliable geologgers are those made by the British Antarctic Survey (although these were also the largest geologgers used in the studies we examined). With regard to analysis methods, we believe there is much room for improvement. Use of online archiving of both data and analysis parameters would greatly improve the repeatability and transparency of geologger research.     

Occurrence and genetics of black‐eyed migratory locusts,Locusta migratoria (Orthoptera: Acrididae)

Black‐eyed Locusta migratoria appeared in albino locusts as a result of crossing between a short‐winged strain originating from Tsushima Island, Japan, and an albino strain originating from Okinawa Island. The black eye trait was recessive to the white eye trait because the crosses between black‐ and white‐eyed albino locusts produced only individuals with white eyes in the F₁ generation. In the F₂ generation, black‐ and white‐eyed individuals appeared in a ratio of 1:3, indicating that the black eye trait was controlled by a simple Mendelian unit. The black eye trait showed no genetic association with other traits including wing morph, adult body dimensions and classical morphometric ratios such as hind femur length / head width and forewing length / hind femur length.     

Efficacy of a fipronil bait in reducing the number of fleas (Oropsylla spp.) infesting wild black‐tailed prairie dogs

Bubonic plague (Yersinia pestis) is a deadly zoonosis with black‐tailed prairie dogs (Cynomys ludovicianus) as a reservoir host in the United States. Systemic insecticides are a promising means of controlling the vectors, Oropsylla spp. fleas, infesting these prairie dogs, subsequently disrupting the Y. pestis cycle. The objective of this study was to conduct a field trial evaluating the efficacy of a grain rodent bait containing fipronil (0.005%) against fleas infesting prairie dogs. The study was performed in Larimer County, CO, where bait was applied to a treatment area containing a dense prairie dog population, three times over a three‐week period. Prairie dogs were captured and combed for fleas during four study periods (pre‐, mid‐, 1^st post‐, and 2^nd post‐treatment). Results indicated the use of bait containing fipronil significantly reduced flea burden. The bait containing fipronil was determined to reduce the mean number of fleas per prairie dog >95% for a minimum of 52 days post‐initial treatment application and 31 days post‐final treatment application. These results suggest the potential for this form of treatment to reduce flea population density on prairie dogs, and subsequently plague transmission, among mammalian hosts across the United States and beyond.     

Short‐distance partial migration of Neotropical birds: a community‐level test of the foraging limitation hypothesis

Partial migration of tropical birds was long believed to be driven by variation in food abundance. Recent evidence from a partially‐migratory species suggests that in contrast, limited foraging opportunities at high elevations during severe wet season storms drives the most metabolically‐challenged individuals down to elevations where rainfall is lighter. Here, I test community‐level predictions of this hypothesis by examining the relationship between high‐elevation rainfall in the second half of the year and counts of migrant birds in lowland forest during late December each year from 1990–2009. I contrast results derived from analysis of all migrant species with both analyses of only the frugivorous migrants, and analyses of resident species. Counts of migrant species were on average positively associated with montane rainfall with differences of up to 72% in the numbers of birds counted in drier or wetter years. Frugivores and smaller birds responded more strongly to variation in rainfall compared to the broader migrant species pool. Interestingly, counts of resident species were also higher following wetter montane wet seasons. Results of analyses exploring the cause of resident responses were not consistent with climatic effects on breeding productivity or short‐term weather effects on detectability. Results were, however, consistent with cryptic down‐slope migration of individuals breeding at higher elevations augmenting lowland resident populations in wet years. These results suggest that changes in rainfall amount, storm intensity, and timing of severe weather events would lead to large increases in or losses of an important behaviour.     

Determining fine‐scale migratory connectivity and habitat selection for a migratory songbird by using new GPS technology

Migratory aerial insectivores are among the fastest declining avian groups, but our understanding of these trends has been limited by poor knowledge of migratory connectivity and the identification of critical habitat across the vast distances they travel annually. Using new, archival GPS loggers, we tracked individual purple martins Progne subis from breeding colonies across North America to determine precise (< > 10 m) locations of migratory and overwintering roost locations in South America and to test hypotheses for fine‐scale migratory connectivity and habitat use. We discovered weak migratory connectivity at the roost scale, and extensive, fine‐scale mixing of birds in the Amazon from distant (> 2000 km) breeding sites, with some individuals sharing the same roosting trees. Despite vast tracts of contiguous forest in this region, birds occupied a much more limited habitat, with most (56%) roosts occurring on small habitat islands that were strongly associated with water. Only 17% of these roosts were in current protected areas. These data reflect a critical advance in our ability to remotely determine precise migratory connectivity and habitat selection across vast spatial scales, enhancing our understanding of population dynamics and enabling more effective conservation of species at risk.     

Using habitat selection theories to predict the spatiotemporal distribution of migratory birds during stopover – a case study of pink‐footed geese Anser brachyrhynchus

Understanding how animals select for habitat and foraging resources therein is a crucial component of basic and applied ecology. The selection process is typically influenced by a variety of environmental conditions including the spatial and temporal variation in the quantity and quality of food resources, predation or disturbance risks, and inter‐ and intraspecific competition. Indeed, some of the most commonly employed ecological theories used to describe how animals choose foraging sites are: nutrient intake maximisation, density‐dependent habitat selection, central‐place foraging, and predation risk effects. Even though these theories are not mutually exclusive, rarely are multiple theoretical models considered concomitantly to assess which theory, or combination thereof, best predicts observed changes in habitat selection over space and time. Here, we tested which of the above theories best‐predicted habitat selection of Svalbard‐breeding pink‐footed geese at their main spring migration stopover site in mid‐Norway by computing a series of resource selection functions (RSFs) and their predictive ability (k‐fold cross validation scores). At this stopover site geese fuel intensively as a preparation for breeding and further migration. We found that the predation risk model and a combination of the density‐dependent and central‐place foraging models best‐predicted habitat selection during stopover as geese selected for larger fields where predation risk is typically lower and selection for foraging sites changed as a function of both distance to the roost site (i.e. central‐place) and changes in local density. In contrast to many other studies, the nutritional value of the available food resources did not appear to be a major limiting factor as geese used different food resources proportional to their availability. Our study shows that in an agricultural landscape where nutritional value of food resources is homogeneously high and resource availability changes rapidly; foraging behaviour of geese is largely a tradeoff between fast refuelling and disturbance/predator avoidance.     

chifish: a computer program testing for genetic heterogeneity at multiple loci using chi‐square and Fisher's exact test

chifish is a 32‐bit Windows/DOS program evaluating divergence at multiple gene loci. It tests the hypothesis of no difference at any locus both by means of Pearson's traditional chi‐square and by using Fisher's method of combining P values obtained by Fisher's exact test. Input data are read from a file formatted for genepop . Commonly used population genetics software do not perform chi‐square tests, and the simultaneous application of both techniques aids in situations where poor power of the ‘exact approach’ may prevent detection of true differentiation (e.g. few populations and few alleles per locus).     

Influence of device accuracy and choice of algorithm for species distribution modelling of seabirds: a case study using black‐browed albatrosses

Species distribution models (SDM) based on tracking data from different devices are used increasingly to explain and predict seabird distributions. However, different tracking methods provide different data resolutions, ranging from < 10 m to > 100 km. To better understand the implications of this variation, we modeled the potential distribution of black‐browed albatrosses Thalassarche melanophris from South Georgia that were simultaneously equipped with a platform terminal transmitter (PTT) (high resolution) and a global location sensor (GLS) logger (coarse resolution), and measured the overlap of the respective potential distribution for a total of nine different SDM algorithms. We found slightly better model fits for the PTT than for GLS data (AUC values 0.958 ± 0.048 vs 0.95 ± 0.05) across all algorithms. The overlaps of the predicted distributions were higher between device types for the same algorithm, than among algorithms for either device type. Uncertainty arising from coarse‐resolution location data is therefore lower than that associated with the modeling technique. Consequently, the choice of an appropriate algorithm appears to be more important than device type when applying SDMs to seabird tracking data. Despite their low accuracy, GLS data appear to be effective for analyzing the habitat preferences and distribution patterns of pelagic species.     

Egg size and offspring quality: a meta‐analysis in birds

Parents affect offspring fitness by propagule size and quality, selection of oviposition site, quality of incubation, feeding of dependent young, and their defence against predators and parasites. Despite many case studies on each of these topics, this knowledge has not been rigorously integrated into individual parental care traits for any taxon. Consequently, we lack a comprehensive, quantitative assessment of how parental care modifies offspring phenotypes. This meta‐analysis of 283 studies with 1805 correlations between egg size and offspring quality in birds is intended to fill this gap. The large sample size enabled testing of how the magnitude of the relationship between egg size and offspring quality depends on a number of variables. Egg size was positively related to nearly all studied offspring traits across all stages of the offspring life cycle. Not surprisingly, the relationship was strongest at hatching but persisted until the post‐fledging stage. Morphological traits were the most closely related to egg size but significant relationships were also found with hatching success, chick survival, and growth rate. Non‐significant effect sizes were found for egg fertility, chick immunity, behaviour, and life‐history or sexual traits. Effect size did not depend on whether chicks were raised by their natural parents or were cross‐fostered to other territories. Effect size did not depend on species‐specific traits such as developmental mode, clutch size, and relative size of the egg, but was larger if tested in captive compared to wild populations and between rather than within broods. In sum, published studies support the view that egg size affects juvenile survival. There are very few studies that tested the relationship between egg size and the fecundity component of offspring fitness, and no studies on offspring survival as adults or on global fitness. More data are also needed for the relationships between egg size and offspring behavioural and physiological traits. It remains to be established whether the relationship between egg size and offspring performance depends on the quality of the offspring environment. Positive effect sizes found in this study are likely to be driven by a causal effect of egg size on offspring quality. However, more studies that control for potential confounding effects of parental post‐hatching care, genes, and egg composition are needed to establish firmly this causal link.