Variation in male‐built nest volume with nesting‐support quality,colony, and egg production in whiskered terns

Nest building can represent an energetically costly activity for a variety of animal taxa. Besides, the determinants of within‐species variation in the design of nests, notably with respect to natural and sexual selection, are still insufficiently documented. Based on an observational study, we examined the influence of nesting conditions (nesting‐support quality, colony, laying date, and year) on male‐built nest volume and also its potential role as a postmating sexually selected display in the whiskered tern Chlidonias hybrida. This tern species is a monogamous colonial bird with obligate biparental care breeding on aquatic vegetation. Hence, large nesting platforms are expected to be a selective advantage because they would better withstand adverse environmental conditions and provide a secure structure for eggs. Nest size may also serve as a postmating sexual trait, and variation in egg production would be positively associated with nest size. We found that nest volume was adjusted to different environmental cues. A positive relationship was found between nest volume and nesting‐support quality, indicating that the leaf density of white waterlily is essential for nest stability. Variation in nest volume was not correlated to colony size but varied among colonies and years. Male‐built nest volume was also positively associated with mean egg volume per clutch but not with clutch size. The fitness consequences of building a large nest are yet to be studied, and additional investigations are recommended to better understand whether the activity of males early during breeding season (e.g., nest building and courtship feeding performance) really serves as postmating sexually selected signals.     

Indirect evidence of conspecific nest parasitism in the colonial whiskered tern (Chlidonias hybrida)

Conspecific nest parasitism (CNP) is widespread among birds and is particularly common in colonial species. Nevertheless, this reproductive tactic is remarkably little described in terns. In this study we use egg morphology as indirect evidence of CNP in the whiskered tern (Chlidonias hybrida) in 17 colonies from two French populations. We study CNP with respect to the duration of egg laying (i.e. clutch size), and also present some behavioural aspects that could facilitate CNP. From the inspection of 121 two-egg, 279 three-egg and 11 more egg-clutches, we detected a minimum %CNP of 9.2% for the two study populations, combined with large differences between the two populations (4.0% vs. 14.5% in three-egg clutches) and between colonies (0 up to 30%). %CNP was significantly higher in three-egg (9.0%) than in two-egg clutches (1.7%), suggesting that duration of egg laying is a determining factor of parasitism. The rather high %CNP detected in this species might be facilitated by (1) the observed long unattendance of nest by any of the pair members, mainly during the pre-incubation period, and (2) high synchrony in laying dates within colonies. A possible link between nest density and CNP rate was suggested. Finally, we encourage new investigations of CNP in other tern species.     

Plant-water regime management in a wetland: consequences for a floating vegetation-nesting bird, whiskered tern Chlidonias hybridus

In this study we investigated the interplay between water level management, floating macrophytic vegetation and nesting whiskered tern (Chlidonias hybridus) during 8 years (1995–2002) at a shallow macrophyte-dominated lake in western France. The specific question was to see if slight increases in the water regime of the lake (three scenarios), as part of a restoration programme, affect the timing of nesting and occupation of colonies by whiskered terns by way of changes in biomass of waterlily (Nymphaea alba) beds where colonies always establish. During the study period, egg-laying dates were progressively delayed up to 40–50 days (mid-May to early July) in relation to late appearance of suitable waterlily biomass (R ² = 0.67, p = 0.01) associated with high spring water levels (1.10–1.25 m vs. 0.95 m). Simultaneously some nesting attempts were observed in sub-optimal habitats (adjacent wet grasslands) in high flooding conditions leading either to poor breeding success or colony desertion. We concluded that there is a need for compromise between the experimental water levels used to control waterlily biomass and the nesting requirements of the whiskered tern population in this lake of international importance (260–510 pairs in 1995–2002, ≥ 1% Ramsar level).     

Reproductive parameters in relation to food supply in the whiskered tern ( Chlidonias hybrida )

It is generally accepted that breeding terns are sensitive to food supply and that their reproductive effort could be substantially affected by the availability and access to resources. In this study we examined reproductive parameters in the whiskered tern Chlidonias hybrida in relation to food supply during the courtship feeding period (food brought by males to females) over a 2-year period (2004–2005). We also studied whether the condition of the nestlings [body condition index (BCI)] was related to a proxy of the reproductive investment of the adults (the clutch size) during a season of food shortage. Behavioural observations showed a decrease in the intensity of male courtship feeding between years (2004 > 2005), and a strong shift in the relative abundance of the two prey groups (invertebrates/fish; invertebrate prey dropped from 88.0 to 49.3%) brought by males. This change in food delivery rates did not result in a delay in laying, but there was a significant difference in mean clutch size between years (2.71 ± 0.49 eggs in 2004 and 2.05 ± 0.78 eggs in 2005) without any within-year variation in relation to the laying dates. The egg size (volume and length) was related to the year (2004 < 2005), suggesting a trade-off in the quantity and the quality of eggs between the two seasons. We also found no evidence that the investment in a large clutch affected nestling BCI in the course of the food shortage season. Since many pairs (about 60%) interrupted breeding during the incubation stage, we assumed that parents that succeeded in rearing nestlings in these conditions were probably ‘high-quality’ individuals. Our results therefore showed that whiskered terns are sensitive to the varying food conditions they experienced throughout the courtship period. The diversity of prey types could be a key factor in the reproductive investment of this tern species.     

Little tern breeding success in artificial and natural habitats: modelling population growth under uncertain vital rates

As a consequence of habitat loss, breeding in man-made habitats has become increasingly common for many coastal breeding bird species. While artificial sites provide valuable substitutes, they may also be more attractive, and importantly, differ in quality from natural sites. Therefore, information on habitat specific breeding success and their potential for supporting stable populations are needed. We compared little tern (Sternula albifrons) breeding success (nest and hatching success) between natural habitat (sandy beaches) and artificial port habitat at Bothnian Bay, Finland from 2006 to 2011. We further reviewed published estimates on pre-fledging and adult survival for little terns and least terns (Sternula antillarum), and used these ranges to estimate plausible parameter spaces for population growth rates given our estimates of breeding success. Nest success was among the highest reported for little terns in the artificial habitat (82 %) while being lower in the natural habitat (58 %). This difference may have resulted from differences in colony sizes and levels of disturbance. Hatching success did not differ significantly, but the percentage of successful nests containing unhatched eggs was twice as high in the natural habitat. The parameter spaces for population growth rates indicated that the artificial habitat has good potential to sustain stable populations (66 % positive growth rate) while for the natural habitat this potential was lower (37 % positive growth rate). While our results suggest that artificial habitats can be very productive breeding sites for habitat deprived tern populations, management should concentrate on improving both habitats with emphasis on natural sites.     

Aggressive neighbors and dense nesting: nest site choice and success in high-Arctic common eiders

Minimizing the risk of nest predation has led some bird species to exploit the nest defense behavior of other species. At Nasaruvaalik Island, Nunavut, Canada, some common eiders (Somateria mollissima borealis) nest within the boundaries of Arctic tern (Sterna paradisaea) colonies, while others nest elsewhere on the island, away from the terns. We tested the effects of location (within vs. outside the tern colonies), density of common eider nests, and annual variation on the nesting parameters of common eiders. Our results suggest that nesting in association with Arctic terns does not confer an obvious benefit to eiders. Such associative nesting of eiders and terns may be the result of overlapping habitat preferences between the two species, or a general scarcity of suitable nesting habitat for ground-nesting species in the high Arctic. However, eiders nesting in higher densities with other eiders had greater nest success and lower total clutch predation, indicating a positive correlation between nest density and success.     

Contrasting heterozygosity‐fitness correlations across life in a long‐lived seabird

Selection is a central force underlying evolutionary change and can vary in strength and direction, for example across time and space. The fitness consequences of individual genetic diversity have often been investigated by testing for multilocus heterozygosity‐fitness correlations (HFCs), but few studies have been able to assess HFCs across life stages and in both sexes. Here, we test for HFCs using a 26‐year longitudinal individual‐based data set from a large population of a long‐lived seabird (the common tern, Sterna hirundo), where 7,974 chicks and breeders of known age were genotyped at 15 microsatellite loci and sampled for life‐history traits over the complete life cycle. Heterozygosity was not correlated with fledging or post‐fledging prospecting probabilities, but was positively correlated with recruitment probability. For breeders, annual survival was not correlated with heterozygosity, but annual fledgling production was negatively correlated with heterozygosity in males and highest in intermediately heterozygous females. The contrasting HFCs among life stages and sexes indicate differential selective processes and emphasize the importance of assessing fitness consequences of traits over complete life histories.     

Does peripheral dislodgement contribute to heterozygote deficiencies in blue mussels?

Differential loss of heterozygous individuals that move to the periphery of mussel aggregations, where they are at greater risk for dislodgement, has been proposed as an explanation for observed heterozygote deficiencies in blue mussels. To test the dislodgement hypothesis, correlations between heterozygosity and mussel motility, as well as characteristics of byssogenesis and byssal thread attachment strengths, were determined in a wild and a farmed population of blue mussels (Mytilus edulis) from New Hampshire, USA. Although both populations exhibited a heterozygote deficit as measured by three microsatellite loci, no relationship was found between heterozygosity and increased motility in either population. Similarly, no relationship was found between heterozygosity and byssogenesis or attachment strength. Hence, differential dislodgement is highly unlikely as a possible contributor to the loss of heterozygous individuals.     

Feral cat diet and impact on sooty terns at Juan de Nova Island, Mozambique Channel

Feral cat Felis catus predation on seabirds has been well documented; however, details regarding shifts in feral cat diet in relation to seabird availability, seabird predation rate and impact on seabird population dynamics are scarce. Here, we present data documenting a seasonal shift in feral cat diet at Juan de Nova Island, Mozambique Channel. We also quantify sooty tern Sterna fuscata predation by feral cats and examine the impact on sooty terns over both the short term (by removing individual cats from sub-colonies) and over the longer term by highlighting their influence on population growth rate ( λ ) using a deterministic matrix model. Cat diet shifted dramatically from insects, rats and mice outside the tern breeding season to primarily terns when terns were breeding. The predation rate of sooty terns at Juan de Nova was estimated at 5.94 terns cat⁻¹ day⁻¹, with a proportion of these (22%) being killed without being consumed ('surplus kills'). When only one cat was removed from each sub-colony, tern predation declined tenfold in the short term. From our matrix model, the annual growth rate for sooty terns was 1.01 in the absence of cat predation. It remained above one until a predation impact equivalent to approximately three times the estimated cat density (12.04 per km²) was incorporated. Our results demonstrate that cats preferentially predate and have an impact on breeding sooty terns at Juan de Nova, and that an increase in cat density could lead to negative effects on population growth, despite the large breeding tern population.     

Predator–prey interactions between the South Polar skua Catharacta maccormicki and Antarctic tern Sterna vittata

Antarctic terns have to co‐exist in a limited space with their major nest predator, the skuas. We conducted artificial nest experiments to evaluate the roles of parental activity, nest location and nest and egg crypsis in this simple predator–prey system. Predation on artificial (inactive) nests was higher in traditional nesting sites than in sites previously not occupied by terns, which suggests that skuas memorized past tern breeding sites. Predation on artificial nests in inactive colonies was higher than in active (defended) colonies. Parental defense reduced predation in colonies to the level observed in artificial nests placed away from colonies. This suggests that communal defense can balance the costs of attracting predators to active colonies. Within colonies, predation was marginally higher on experimental eggs put in real nests than on bare ground. Although it seems that the presence of a nest is costly in terms of increased predation, reductions in nest size might be constrained by the need for protective nest structures and/or balanced by opposing selection on nest size. Predation did not differ markedly between artificial (quail) and real tern eggs. A simultaneous prey choice experiment showed that the observed predation rates reflected egg/nest detectability, rather than discrimination of egg types. In summary, nesting terns probably cannot avoid being detected, and they cannot defend their nest by attending them. Yet, by temporarily leaving the nest, they can defend it through communal predator mobbing, and at the same time, they can benefit from crypsis of unattended nest and eggs.     

HETEROZYGOSITY,AGGRESSION, AND POPULATION FLUCTUATIONS IN MEADOW VOLES (MICROTUS PENNSYLVANICUS)

We tested whether variation in heterozygosity could produce cyclic changes in population size in meadow voles (Microtus pennsylvanicus). For this to occur, three conditions must be met: (1) populations are more outbred (heterozygotic) at high than low population density, (2) heterozygotic voles are more aggressive than relatively inbred individuals, and (3) heterozygotic voles have lower reproductive fitness, though being superior at defending resources. We found no evidence that heterozygosity varied with population size or that reproductive success varied with heterozygosity. However, the former test was indirect and relatively weak. We directly measured aggression and heterozygosity of individual voles. Aggression was significantly related to heterozygosity: higher heterozygosity correlated with more aggression in males and less aggression in females. The proportion of variance in aggression that could be explained by heterozygosity was small. These results suggest that changes in population size of meadow voles could not be driven by systematic changes in heterozygosity with population size.     

Can a simple algebraic analysis predict markers-genome heterozygosity correlations?

A current algebraic analysis on genome-wide heterozygosity estimates suggests that correlations between molecular markers and genome-wide heterozygosity, rho, depend on the ratio between the number of markers used, r, and the number of genome loci, n; that is: rho approximately square root r/n. Hence, it is unfeasible to obtain reliable estimates of genome-wide heterozygosity in species of large genome using a few markers. We cast some doubts about this analysis as it assumed that the probability that an individual was heterozygous at a locus is equal to the average heterozygosity of this locus in the population. However, we believe that individual heterozygosity at a given locus depends on individual pedigree. Because the pedigree is common for all loci of an individual, their probabilities of heterozygosity are not independent within the genome. We first performed simulations generating random genomes for 100 individuals. Among these individuals, markers and genome-wide heterozygosities correlated as expected from the above equation. However, when we simulated random mating among these individuals and in successive generations including their descendents, as occur in real populations, the correlations between markers and genome-wide heterozygosity were much higher than those predicted from algebraic analyses, and estimates of genome-wide heterozygosity improved slightly with the increment of the number of loci in the genome.     

Wie richtet eine Flußseeschwalbenkolonie (Sterna hirundo) ihr Abwehrverhalten auf den Feinddruck durch Silbermöwen (Larus argentatus) ein?

How a Common Tern (Sterna hirundo) Colony Defends itself against Herring Gulls (Larus argentatus) The subject of this study is the anti-predator behaviour of a small common tern colony near a large herring gull colony on the island of Mellum, West Germany (Fig. 1). In 1980 the number of gulls crossing this tern colony increased during the terns' chick-stage (Fig. 4). Observed predation of tern chicks was independent of tide and time of day (3., Fig. 5). The frequency of tern reactions corresponded to the number of herring gull crossings (Fig. 5, Table 1). The terns' responses increased between morning and evening (Fig. 8). Tern up-flights and attacks increased absolutely and as a percentage, with the advance of the breeding season (Fig. 3, 4). They were positively correlated with the observed chick predation and the number of pairs with chicks, most markedly with chicks older than 5 days (Figs. 3, 4; Table 1). This increased defence was maintained by fewer pairs as, by then, many had lost their own broods (Fig. 4). As the breeding season progressed, herring gulls increasingly became the main cause of tern up-flights and the object of the attacks (Figs. 9–11). The up-flights of the whole colony, which occurred frequently and spontaneously during incubation, were observed only rarely after hatching and were almost exclusively a response to herring gulls (Figs. 10, 12). The lower herring gulls flew over the colony, the more frequently common terns flew up or attacked and the more individuals were involved in these responses (Figs. 6, 13, 14). During the breeding period, communal up-flights and attacks by terns increased as a percentage (Figs. 12, 13, 15–17). Group-attacks effected changes in the gulls' flying-routes more often than did individual attacks (Fig. 18). Despite the defence behaviour and its adaptation to the predation pressure, herring gulls often succeeded in robbing chicks. This is why the breeding success of the common tern was poor (< 0.4 chicks/nest). Possible reasons for this are discussed.     

Influences of multilocus heterozygosity on size during early life

Genetic diversity has been hypothesized to promote fitness of individuals and populations, but few studies have examined how genetic diversity varies with ontogeny. We examined patterns in population and individual genetic diversity and the effect of genetic diversity on individual fitness among life stages (adults and juveniles) and populations of captive yellow perch (Perca flavescens) stocked into two ponds and allowed to spawn naturally. Significant genetic structure developed between adults and offspring in a single generation, even as heterozygosity and allelic richness remained relatively constant. Heterozygosity had no effect on adult growth or survival, but was significantly and consistently positively related to offspring length throughout the first year of life in one pond but not the other. The largest individuals in the pond exhibiting this positive relationship were more outbred than averaged size individuals and also more closely related to one another than they were to average‐sized individuals, suggesting potential heritability of body size or spawn timing effects. These results indicate that the influence of heterozygosity may be mediated through an interaction, likely viability selection, between ontogeny and environment that is most important during early life. In addition, populations may experience significant genetic change within a single generation in captive environments, even when allowed to reproduce naturally. Accounting for the dynamic influences of genetic diversity on early life fitness could lead to improved understanding of recruitment and population dynamics in both wild and captive populations.     

Near-infrared spectroscopy identifies the colony and nest of origin of weaver ants, <Emphasis Type="Italic">Oecophylla smaragdina</Emphasis>

The ability of social insects to differentiate between colony members and others is essential for the survival of the colony. It enables individuals to direct altruistic behavior towards colony mates, while protecting the colony from intruders. Colonies have a distinct chemical signature that facilitates colony-mate recognition. However, in large polydomous colonies, this signal is likely to be modified by factors unique to each nest. We demonstrate, using near-infrared spectroscopy (NIRS), that individual weaver ants, Oecophylla smaragdina, can be differentiated with respect to their colony and nest of origin. 76.5% of individuals from four colonies could be correctly assigned to their colony of origin; and 79.6% of individuals could be assigned to the correct nest (of two) within their colony. Despite the differences between nests within colonies, in most cases individuals from one nest were more similar to individuals from the other nest within the colony than they were to individuals from any nest outside the colony. Therefore, a distinctive colony identity is maintained despite differences between nests within colonies. We discuss the advantages of using NIRS as a faster and less expensive alternative to the analysis of cuticular hydrocarbons following extraction and identification with gas chromatography/mass spectroscopy. Received 26 November 2007; revised 22 January 2008; accepted 25 January 2008.     

Colony size and not nest density drives reproductive output in the Common Tern Sterna hirundo

Density is known to be an important factor in population size regulation. Several mechanisms of density limitation have been identified in colonial birds. We studied competition in Common Terns Sterna hirundo to assess whether the factor limiting reproductive output was competition for nest‐sites, which is dependent on local nest density, or density‐dependent competition for food resources, which is dependent on overall colony size using the same foraging area. We found strong associations of both colony size and nest density with reproductive output in five colonies of Common Terns in three different habitats (one marine, two freshwater). Based on detailed long‐term datasets of six separate sub‐colonies of the Banter See colony that differed in nest density, we found that reproductive success was not related to nest density but to overall colony size, possibly a result of resource depletion and food competition. We also found carry‐over effects of colony size during rearing on post‐fledging return rate. These results have important implications for the conservation management plans aimed at recovering declining populations of Common Terns.     

Aggression and nest spacing in single and mixed species groups of seabirds

When heterospecific seabirds are part of a nesting colony, there may be less opportunity for conspecifics to come in direct contact with each other, resulting in lower intraspecific aggressiveness. To determine if individuals spend less time in aggressive behavior when nesting in conspecific rather than heterospecific groups, we compared the behavior of black skimmers (Rhynchops niger) nesting with gull-billed terns (Sterna nilotica) in three mixed species subcolonies to those of black skimmers in three single species subcolonies. In contrast to our predictions, black skimmers spent significantly less time in aggressive behaviors when nesting in single species subcolonies than when nesting with heterospecifics. Although skimmers in mixed species subcolonies tended to have more aggressive interactions with skimmers than terns, this may be a function of subcolony composition; the proportions of aggressive interactions with conspecifics were similar to the proportions of conspecifics in each subcolony. However, within the mixed species subcolonies, skimmers that nested nearer to terns were involved in aggressive interactions significantly less than skimmers that nested closer to conspecifics. Also, skimmers nested closer to their nearest neighbor when it was a gull-billed tern than when it was another skimmer. Regardless of which species they nested closest to, skimmers were more aggressive towards other skimmers than to terns within the mixed species subcolonies. Distance to nearest neighbor's nest did not differ significantly between the colony types, and did not seem to influence the duration of aggressive activity in the single species subcolonies. In the mixed species subcolonies, however, the time spent in aggressive behavior increased as the distance to nearest neighbor increased. It appears that of the several benefits that have been proposed of mixed species colonies, reduced time spent in conspecific aggression is not among them. However, within a mixed species colony, an individual can reduce time spent in aggressive interactions by nesting near heterospecifics. Received: 16 September 1996 / Accepted: 24 February 1997     

Demographic effects of extreme weather events: snow storms,breeding success,and population growth rate in a long‐lived Antarctic seabird

Weather extremes are one important element of ongoing climate change, but their impacts are poorly understood because they are, by definition, rare events. If the frequency and severity of extreme weather events increase, there is an urgent need to understand and predict the ecological consequences of such events. In this study, we aimed to quantify the effects of snow storms on nest survival in Antarctic petrels and assess whether snow storms are an important driver of annual breeding success and population growth rate. We used detailed data on daily individual nest survival in a year with frequent and heavy snow storms, and long term data on petrel productivity (i.e., number of chicks produced) at the colony level. Our results indicated that snow storms are an important determinant of nest survival and overall productivity. Snow storm events explained 30% of the daily nest survival within the 2011/2012 season and nearly 30% of the interannual variation in colony productivity in period 1985–2014. Snow storms are a key driver of Antarctic petrel breeding success, and potentially population dynamics. We also found state‐dependent effects of snow storms and chicks in poor condition were more likely to die during a snow storm than chicks in good condition. This stresses the importance of considering interactions between individual heterogeneity and extreme weather events to understand both individual and population responses to climate change.     

Invasion history and demographic processes associated with rapid morphological changes in the Red‐whiskered bulbul established on tropical islands

The Red‐whiskered bulbul is a very successful invasive bird species. Morphological differences have been reported among individuals inhabiting the humid and dry coasts of Reunion Island, in a 30‐year‐old population. This suggests a capacity for rapid local adaptation which could explain the general invasive success of this species. However, the origin and invasion history of this population is unknown. It is therefore not possible to establish with certainty the cause of these morphological differences. Here, we investigated the invasion history of populations of Red‐whiskered bulbul established on Reunion Island, Mauritius and Oahu (three geographically similar tropical islands) to assess the link between invasion history and morphological changes in these populations. We first assessed the source(s) of the invasive populations. We then compared the morphology of the individuals between the invasive and native populations and between the dry and humid coasts of invaded islands. Finally, we inferred the invasion history of the invasive populations to investigate the role of neutral processes (e.g. founder effect and drift) on morphology. We found that the invasive populations have a similar origin and that the morphology of the individuals in these populations has diverged in a similar way from the native range, suggesting a convergent adaptation to tropical islands. Like on Reunion, we found differences in morphology between the dry and humid coasts on Mauritius. These morphological differences can be explained by invasion history on Reunion but not on Mauritius. Both neutral evolution and adaptation thus shape the morphology of invasive Red‐whiskered bulbuls.     

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.