Seasonal and annual differences in the foraging ecology of two gull species breeding in sympatry and their use of fishery discards

Niche segregation between similar species will result from an avoidance of competition but also from environmental variability, including nowadays anthropogenic activities. Gulls are among the seabirds with greater behavioural plasticity, being highly opportunistic and feeding on a wide range of prey, mostly from anthropogenic origin. Here, we analysed blood and feather stable isotopes combined with pellet analysis to investigate niche partitioning between Audouin's gull Larus audouinii and yellow‐legged gull Larus michahellis breeding in sympatry at Deserta Island, southern Portugal, during 2014 and 2015. During the breeding season there was considerable overlap in the adults’ diet, as their stable isotope values of blood and primary feather (P1) did not differ, and their pellets were comprised mainly by marine fish species. However, Audouin's gulls presented higher occurrences of pelagic fish, while yellow‐legged gulls fed more on demersal fish, insects, and refuse. SIAR mixing models also estimated a higher proportion of demersal fish in the diet of yellow‐legged gulls. We also found differences between the two gull species in chicks’ feathers, suggesting that Audouin's gull adults selected prey with lower carbon isotope values to feed their young. Secondary feather (S8) of Audouin's gull presented higher isotope values compared to yellow‐legged gulls, indicating different foraging areas (δ¹³C) and/ or trophic levels (δ¹⁵N) between the two species in the non‐breeding season. During both the all‐year and non‐breeding periods the yellow‐legged gull showed a broader isotopic niche width than Audouin's gull in 2013, and in 2014 the two gull species exhibited different isotopic niche spaces. Our study suggests that both gull species foraged in association with fisheries during the breeding season. In this sense, a discard ban implemented under the new European Union Common Fisheries Policy may lead to a food shortage, therefore future research should closely monitor the population dynamics of Audouin's and yellow‐legged gulls.     

Seasonal variation in male testosterone levels in a tropical bird with year‐round territoriality

Testosterone is important in mediating investment in competing activities such as territoriality, parental care, and maintenance behavior. Most studies of testosterone function have focused on temperate species and less is known about the role of testosterone in territoriality or variation in mating systems of tropical species. Results of studies of tropical species with year‐round territoriality indicate that territorial aggression during the non‐breeding season is maintained with low levels of testosterone, and increased levels of testosterone in males during the breeding season may increase mating opportunities or aid in competition for mates. We studied seasonal variation in testosterone levels of male Red‐throated Ant‐tanagers (Habia fuscicauda), a socially monogamous species with year‐round territoriality and with high levels of extra‐pair matings (41% of young), to determine if testosterone levels increased during the breeding season. We captured males during the non‐breeding and breeding seasons and collected blood samples for hormone analysis. We found that mean testosterone concentrations were low during the non‐breeding season (0.18 ± 0.05 [SD] ng/ml, range = 0.11–0.31 ng/ml), and significantly higher during the breeding season (2.37 ± 2.47 ng/ml, range = 0.14–6.28 ng/ml). Testosterone levels of breeding males were not related to aggression levels as measured by attack rates toward a stuffed decoy or singing rates during simulated territorial intrusions. These results suggest that the higher testosterone levels of breeding male Red‐throated Ant‐tanagers may be important in an extra‐pair mating context, possibly in display behavior or mate attraction, but additional study is needed to clarify the role of testosterone during the breeding season.     

Age,condition and dominance‐related sexual ornament size before and during the breeding season in the black grouse Lyrurus tetrix

Male ornaments function as honest cues of male quality in many species and are subject to intra‐ and intersexual selection. These ornaments are generally studied during peak expression, however their size outside the breeding season may determine ultimate ornament size and costliness, and as such reproductive success. We investigated whether male black grouse Lyrurus tetrix eye comb size was related to age, condition and measures of male dominance before and during the breeding season. Total combined eye comb size began to increase ~70 d before the start of the breeding season. Adult males (aged ≥ 2 yr old) had consistently larger eye combs than younger males (1 yr old) both before and during the breeding season. Heavier and more dominant adult males (attending the lek more frequently and successfully reproducing) had larger eye combs. For younger males, those that were heavier had larger eye combs. Additionally, males that spent more time on the lek showed increased eye comb size as the breeding season approached. Overall we find that ornament size is positively related to dominance and condition before and during the breeding season. Since dominance is accrued through year‐round interactions in many species, the ability to maintain larger signals over prolonged periods, including outside of the breeding season, is likely to be beneficial for adults. For younger males, it is likely that they cannot sustain or are constrained from producing larger eye combs over long periods of time. They therefore prioritise growth of their ornaments later, and according to the amount of time they spend on the lek.     

Gulls can change their migratory behavior during lifetime

Migration is a widespread phenomenon among birds and is likely to be subject to strong selective pressures. Birds' annual routines and behaviors might be expected to change during their different life history stages, resulting in different, age‐related migration patterns. However, although migration has been the subject of many publications, age‐related differences in migration have received little attention. The present study examined age‐related changes in individual migration habits in lesser black‐backed gulls, Larus fuscus. We analyzed data from 10‐year (1998–2007) color‐ringing project in NW England, comprising more than 10 000 ringed individuals. Our results showed a latitudinal cline in age structure across the wintering range, with adults and gulls in their first breeding year wintering closer to the breeding grounds. Supporting this result we observed that individuals, as they get older, changed the migration behavior and winter closer to the breeding areas. Interestingly, we found no differences in survival rates across the wintering grounds. Thus differences in survival rates can not account for the latitudinal cline in age structure, and the observed findings seem to be best explained by the arrival time hypothesis, based on a mechanism whereby individuals are able to change their migratory behavior as result of the onset of sexual maturity and associated mating pressures.     

Long‐distance movements and evidence of post‐breeding elevational movements by Cassin’s Vireos

Bird migration is often framed as a straightforward journey between one breeding site and one wintering site, but recent research has shown that the reality is often more complex. Many species of birds undertake short‐distance movements separate from long‐distance migration. Such movements appear to be common in species that breed in western North America, where mountainous terrain creates a mosaic of habitats and climatic conditions. However, individual‐based tracking studies have disproportionately focused elsewhere, leaving gaps in our understanding of the year‐round movements of western species. I used tracking data from light‐level geolocators and citizen science data from eBird to study the movements of Cassin’s Vireos (Vireo cassinii) breeding in the Sierra Nevada mountains of California, USA. During three breeding seasons (2013–2015), my observations suggested that Cassin’s Vireos vacate their breeding territories during the post‐breeding period in July and August. In April and May 2016, I tagged 22 Cassin’s Vireos with light‐level geolocators and, in April and May 2017, recaptured four that had retained their geolocators. Geolocator data showed that these four birds remained in the same geographic region as their breeding territories (likely the same mountain range) during the post‐breeding period in July and August 2016, ruling out the possibility of long‐distance movements during this time. Analysis of eBird citizen science data suggested that Cassin’s Vireos undertake short‐distance molt‐migration to higher elevations in the Sierra Nevada Mountains during the post‐breeding period. Geolocator data revealed that long‐distance fall migration took place in September and spring migration in April or May, and the four birds spent the winter in different parts of the Mexican winter range of Cassin’s Vireos. These results add to the body of literature on the complex movements of migratory songbirds breeding in the mountains of western North America, an understanding that will be important for effective conservation in the future.     

Testosterone and year‐round territoriality in tropical and non‐tropical songbirds

Past studies have suggested a fundamental difference in testosterone concentrations between tropical and northern latitude male birds, with the convention being that males in the tropics express much lower levels of testosterone. However, recent comparative studies have shown that tropical males with a short and synchronous breeding season (i.e. a breeding season typical of northern species) express maximum testosterone levels similar to those of northern latitude birds. Here, we ask the converse: do northern latitude songbirds that express a defining life‐history characteristic typical of the tropics, i.e. year‐round territoriality, have an annual testosterone profile similar to that of tropical songbirds? For the few year‐round territorial species for which data are available, we found that seasonal testosterone profiles and seasonal maxima in plasma testosterone were similar between males of tropical and non‐tropical species. For example, males of both groups expressed seasonal maxima during the period when females were fertile, and testosterone levels at this time were similar. In contrast, this and other studies show that species with seasonal territories typically express maximum testosterone levels earlier in the breeding cycle, when territories are first being established. Taken together, we suggest that specific life‐history traits may play a more important role in determining testosterone profiles of tropical and non‐tropical birds than breeding latitude and encourage further studies to allow for more formal comparisons.     

Reproductive investment and parental roles in Sabine's gulls<Emphasis Type="Italic"> Xema sabini</Emphasis>

More than 90% of avian species exhibit biparental care, though parental activities are often shared unequally between the members of a pair. Among gull species (Laridae), males and females generally share parental activities, although there appear to be considerable differences between species in the relative contribution of each sex. This study examined the behaviour of male and female Sabine's gulls (Xema sabini) during the incubation period and immediately post-hatch; particularly the amount of time they each invested in breeding activities. Although considered an aberrant species in aspects of behaviour and biology, the Sabine's gull showed a high reproductive investment by both sexes, as other gull species do. Males fed females prior to egg laying and contributed equally to incubation and chick provisioning, and females contributed equally to nest defence. Overall, there was no difference between the sexes in the extent of their contributions to parental care, although there was considerable individual variation within pairs. Sabine's gulls are Arctic breeders and the extent of their contributions to parental activities could have been influenced by their extreme breeding environment and short breeding season.     

Effect of breeding performance on the distribution and activity budgets of a predominantly resident population of black‐browed albatrosses

Pelagic seabirds breeding at high latitudes generally split their annual cycle between reproduction, migration, and wintering. During the breeding season, they are constrained in their foraging range due to reproduction while during winter months, and they often undertake long‐distance migrations. Black‐browed albatrosses (Thalassarche melanophris) nesting in the Falkland archipelago remain within 700 km from their breeding colonies all year‐round and can therefore be considered as resident. Accordingly, at‐sea activity patterns are expected to be adjusted to the absence of migration. Likewise, breeding performance is expected to affect foraging, flying, and floating activities, as failed individuals are relieved from reproduction earlier than successful ones. Using geolocators coupled with a saltwater immersion sensor, we detailed the spatial distribution and temporal dynamics of at‐sea activity budgets of successful and failed breeding black‐browed albatrosses nesting in New Island, Falklands archipelago, over the breeding and subsequent nonbreeding season. The 90% monthly kernel distribution of failed and successful breeders suggested no spatial segregation. Both groups followed the same dynamics of foraging effort both during daylight and darkness all year, except during chick‐rearing, when successful breeders foraged more intensively. Failed and successful breeders started decreasing flying activities during daylight at the same time, 2–3 weeks after hatching period, but failed breeders reached their maximum floating activity during late chick‐rearing, 2 months before successful breeders. Moon cycle had a significant effect on activity budgets during darkness, with individuals generally more active during full moon. Our results highlight that successful breeders buffer potential reproductive costs during the nonbreeding season, and this provides a better understanding of how individuals adjust their spatial distribution and activity budgets according to their breeding performance in absence of migration.     

Year‐round spatial movements and trophic ecology of Trindade Petrels (Pterodroma arminjoniana)

Trindade Petrels (Pterodroma arminjoniana) are vulnerable gadfly petrels that breed on the remote Trindade Island, located ~1100 km off the Brazilian coast. Little is known about their spatial ecology, and their trophic ecology has only been described for the breeding season. We tagged four Trindade Petrels with global location sensing loggers (GLS) from October 2013 to November 2014 and sampled the blood and feathers (innermost primary and the eighth secondary) of 14 individuals to evaluate their year‐round spatial and isotopic ecology. We examined individual distributions, habitat use and suitability, activity, and isotopic values during the breeding, migration, and non‐breeding periods. Trindade Petrels used areas in the southwest Atlantic Ocean (between 10°N and 50°S in latitude) during the breeding season. They migrated through pelagic waters of the tropical Atlantic to the northwest Atlantic, where they spent the non‐breeding season. Trindade Petrels used mostly tropical to subtropical waters in areas of intermediate to high wind speeds and low marine productivity. Individuals spent more time foraging at night than during the day. During the breeding season, birds in northerly areas had higher carbon‐13 values, and birds that used more pelagic areas foraged on prey at a higher trophic level (higher nitrogen‐15 values) than those in more southern and coastal areas. Isotopic values during the breeding, migration, and non‐breeding periods differed, possibly due to differences among individuals in their at‐sea distribution throughout the year. We confirmed the non‐breeding distribution of Trindade Petrels, which was previously known only from vessel sightings and stranded birds. Our results also suggest a strong temporal segregation in the at‐sea distribution and trophic ecology between two groups of individuals, which might indicate the existence of two separate breeding populations.     

Timing of migration and residence areas during the non‐breeding period of barn swallows Hirundo rustica in relation to sex and population

We investigated sex‐ and year‐dependent variation in the temporal and spatial movement pattern of barn swallows Hirundo rustica during the non‐breeding period. Hundred and three individuals equipped with miniaturized light‐level geolocators at three different breeding areas in southern Switzerland and northern Italy provided data for the analysis. We identified a region 1000 km in radius centred in Cameroon as the main non‐breeding residence area of these three geographical populations. Five residence areas of males only were in southern Africa, south of 19°S. Most individuals occupied a single site during their stay south of the Sahara. The timing of migration broadly overlapped between sexes and all geographical breeding populations. Between the two study years there was a distinct difference of 5 to 10 d in departure dates from and arrival at the breeding sites. Remarkably, the period of residence in sub‐Saharan Africa was very similar (157 d) in the two study years, but their positions in the first year (2010–2011) were about 400 km more to the north than in the second (2011–2012). Independent of the year, individuals with sub‐Saharan residence areas further north and east had a shorter pre‐breeding migration and arrived earlier than those staying further south and west. In addition, birds breeding in southern Switzerland arrived at their breeding colony 7–10 d later than those breeding only 100 km south, in the Po river plain. Our study provides new information on the variance in migration phenology and the distribution of residence areas in sub‐Saharan Africa in relation to sex, population and year. It supports the usefulness of light‐level geolocators for the study of annual routines of large samples of small birds.     

Breeding Experience and not Age Modulates the Song Development of Pied Flycatchers (Ficedula hypoleuca)

In songbirds, the development of the species‐specific adult song involves a learning process that varies in extension. In species that incorporate new song elements throughout life (open‐ended learners), variation in male song composition could be the result of either age or breeding experience. Using data from 16 yr of fieldwork on pied flycatchers (Ficedula hypoleuca), we aimed to disclose the individual contribution of these two factors on the species song characteristics, as well as their relation with morphology and plumage color changes. Finally, we explored whether any of the song or physical features could predict the probability of males returning to the breeding site. We found that the song characteristics of the first‐time breeders did not differ between age classes, except for the total number of syllables per song, which was higher in the 1‐year‐old than in the 2‐year‐old males. However, we found that song variables associated with complexity (song and sample versatilities and repertoire size), increased significantly from the first to the second breeding season. Males showed delayed plumage maturation, with 1‐year‐old males being browner than the 2‐year‐old males independently of their breeding experience. Morphology, however, was not affected by age or breeding experience. The probability that males returned to the breeding site was not associated with song or physical features. Considering that some song learning occurs during the breeding period and that some males may skip the first breeding season, selective pressures may have been established for song complexity to be an honest indicator of breeding experience rather than age.     

Annual cycle of the black‐capped chickadee: Seasonality of singing rates and vocal‐control brain regions

Black‐capped chickadees have a rich vocal repertoire including learned calls and the learned fee‐bee song. However, the neural regions underlying these vocalizations, such as HVC, area X, and RA (robust nucleus of arcopallium), remain understudied. Here, we document seasonal changes in fee‐bee song production and show a marked peak in singing rate during March through May. Despite this, we found only minimal seasonal plasticity in vocal control regions of the brain in males. There was no significant effect of time of year on the size of HVC, X, or RA in birds collected in January, April, July, and October. We then pooled birds into two groups, those with large testes (breeding condition) and those with small testes (nonbreeding), regardless of time of year. Breeding birds had slightly larger RA, but not HVC or X, than nonbreeding birds. Breeding birds had slightly larger HVC and RA, but not X, as a proportion of telencephalon volume than did nonbreeding birds. Birds collected in July had heavier brains than birds at other times of year, and had the greatest loss in brain mass during cryoprotection. The absence of any overall seasonal change in the vocal‐control regions of chickadees likely results from a combination of individual differences in the timing of breeding phenology and demands on the vocal‐control regions to produce learned calls year‐round. © 2006 Wiley Periodicals, Inc. J Neurobiol, 2006     

Indirect genetic effects in a sex‐limited trait: the case of breeding time in red‐billed gulls

Female reproductive performance can be strongly affected by male care, so that breeding time, a trait expressed only by females, can be seen as one trait determined by both male and female genotypes. Animal model analyses of a 46‐year study of red‐billed gulls (Larus novaehollandiae scopulinus) revealed that laying date was not heritable in females (h² = 0.001 ± 0.030), but significantly so in males (h² = 0.134 ± 0.029). Heritability of breeding time in males probably reflects genetic variability in some other trait such as courtship feeding ability. In line with predictions of evolutionary models incorporating indirect genetic effects, the strong and consistent directional selection for advanced breeding time has not resulted in detectable selection response in males. Our results demonstrate that a female trait is largely determined by genetic characteristics of its mate, and hence, any evolutionary change in red‐billed gull breeding time depends critically on genetic variation in males.     

Lek fidelity and movement among leks by male Greater Sage‐grouse Centrocercus urophasianus: a capture‐mark‐recapture approach

Males in lek mating systems tend to exhibit high fidelity to breeding leks despite substantial evidence of skewed mating success among males. Although movements between leks are often reported to be rare, such movements provide a mechanism for an individual to improve lifetime fitness in response to heterogeneity in reproductive conditions. Additionally, estimates of apparent movements among leks are potentially biased due to unaccounted variation in detection probability across time and space. We monitored breeding male Greater Sage‐grouse Centrocercus urophasianus on 13 leks in eastern Nevada over a 10‐year period, and estimated movement rates among leks using capture‐mark‐recapture methods. We expected that male movement rates among leks would be low, despite predictions of low breeding success for most males, and that detection rates would be highly variable among leks and years. We used a robust design multistate analysis in Program mark to estimate probability of movements among leks, while accounting for imperfect detection of males. Male Sage‐grouse were extremely faithful to their leks; the annual probability of a male moving away from its original lek of capture was approximately 3% (se = 0.01). Detection probabilities varied substantially among leks (range = 0.21–0.95), and among years (range = 0.30–0.76), but remained relatively constant within years at each lek. These results suggest that male Sage‐grouse dispersal is either rare, or consists primarily of dispersal of sub‐adults from their natal areas prior to the breeding season. The study highlights the benefits of robust design multistate models over standard ‘live‐encounter’ analyses, as they not only permit estimation of additional parameters, such as movement rates, but also allow for more precise parameter estimates that are less sensitive to heterogeneity in detection rates. Additionally, as these data were collected using capture‐mark‐recapture methods, our approach to estimating movement rates would be beneficial in systems where radiotagging is detrimental to the study organism.     

Distribution and at‐sea behavior of Bermudan White‐tailed Tropicbirds (Phaethon lepturus catesbyi) during the non‐breeding season

The movements and behavior of many taxa of seabirds during the non‐breeding season remain poorly known. For example, although studies conducted in the Pacific and Indian oceans suggest that White‐tailed Tropicbirds (Phaethon lepturus) seldom fly more than a few thousand kilometers from nest colonies after breeding, little is known about the post‐breeding movements and behavior of a subspecies of White‐tailed Tropicbirds (P. l. catesbyi) that breeds on islands in the North Atlantic Ocean. Our objective, therefore, was to use light‐based geolocators to identify the ranges and pelagic activities of White‐tailed Tropicbirds from Bermuda during the non‐breeding periods in 2014–2015 (N = 25) and 2015–2016 (N = 16). Locations were estimated based on changes in light intensity across time, and pelagic activities were determined based on whether geolocators attached to leg bands were wet (i.e., birds resting on the water's surface) or dry (i.e., birds in flight). In 2014, birds spent late summer (July–September) near Bermuda and the British Virgin Islands; by mid‐September, most (N = 17; 68%) birds took a direct easterly route to the Sargasso Sea. In 2015, most post‐breeders (N = 15; 94%) flew east from Bermuda and to the Sargasso before the end of late summer. For both years combined, fall and winter (October–February) ranges extended as far west as North Carolina and as far east as the mid‐Atlantic Ridge. In both years, all birds were located between Bermuda and the British Virgin Islands during the spring (April–May). All birds then flew north to Bermuda in both years, with variations in timing, during April and May. We also found extensive overlap in the ranges of males and females during the non‐breeding season in both years. During the non‐breeding season, White‐tailed Tropicbirds spent 5% of night periods and 41% of day periods in flight in 2014; in 2015, birds spent 8% and 42% of night and day periods, respectively, in flight. Tropicbirds spent more time flying during the day because they hunt by day, detecting prey on the wing by sight. Overall, our results suggest that White‐tailed Tropicbirds that breed in Bermuda are diurnal, nomadic wanderers that range over an extensive area of the Atlantic Ocean during the non‐breeding season.     

Breeding territory fidelity in a partial migrant, the American dipper Cinclus mexicanus

American dipper Cinclus mexicanus populations are frequently composed of resident individuals that occupy permanent territories year round and migratory individuals that overwinter with residents but migrate to breeding territories on higher elevation creeks each spring. Between 1999 and 2004 we examined how migratory strategy (resident/migratory) and sex differences influence breeding territory fidelity of American dippers occupying the Chilliwack River watershed, British Columbia, Canada. Counter to expectation we found that the migratory strategy of American dippers did not influence whether birds breeding in one year were found on their former breeding territory in the next. Migratory strategy also did not affect the probability that known surviving dippers occupied the same breeding territory in the following year. Males and females were equally likely to be found on their former territory in the following year (females 43%, males 41%) and known survivors had similar levels of breeding territory fidelity (females 74%, males 68%). However, breeding territory fidelity of males and females varied in response to different factors. Surviving female dippers were more likely to be found on their former breeding territory in the subsequent year following a successful breeding attempt than an unsuccessful breeding attempt. Prior reproductive performance did not influence whether surviving male dippers were found on their former breeding territory. Male dippers were more likely to be found on their former territory and, if they survived, have higher breeding territory fidelity when their mate also returned to that same territory. Mate retention also influenced whether females were found on their former territory in the following year but had no effect on the breeding territory fidelity of known survivors. We argue that sex‐specific dispersal decision rules in American dippers are driven by sex differences in the predictability of breeding performance between years and sex differences in how mate retention influences subsequent reproductive success.     

Range sizes and habitat use of non‐breeding Crested Caracaras in Florida

In Florida, habitats that include the breeding territories of Crested Caracaras (Caracara cheriway) are protected, but non‐breeding individuals may be vulnerable because they may occupy different areas and habitats. We captured and radio‐tagged 58 non‐breeding caracaras in Florida from July 2006–March 2009, determined their locations during weekly flights, and used GIS and compositional analysis to evaluate range sizes and habitat use. Non‐breeding caracaras (N = 58) ranged five times more widely during breeding seasons (N = 573 locations) than during non‐breeding seasons (N = 592 locations), and ranged >250 times more widely than breeding caracaras that defend territories year‐round. The large ranges of non‐breeders suggest they may be searching for and evaluating prospective territories or breeding opportunities (territory prospecting). Pasture occupied by cattle was the most used habitat relative to availability and was used more than pasture without cattle, likely because insects associated with cattle are an important food source for caracaras. Cattle numbers in Florida are declining and, because both breeding and non‐breeding caracaras primarily occupy pasture, this may present difficulties for long‐term management. Citrus groves were also used more than expected given availability by non‐breeding caracaras, but are rarely included in nesting territories. Because pasture and citrus were often adjacent, we suggest that citrus groves may function as refugia from socially dominant breeding caracaras. Conservation and recovery efforts for Florida's caracara population are needed throughout the range of non‐breeders, and should include management that ensures availability of habitat matrices of cattle pasture and citrus groves.     

Investment in testes,sperm‐duct glands and lipid reserves differs between male morphs but not between early and late breeding season in Pomatoschistus minutus

This study of the sand goby Pomatoschistus minutus, a nest‐holding fish with paternal care, focused on gonadal investment among males of different sizes collected early and late in the breeding season. All males caught at the nest had breeding colour, whereas trawl‐caught fish consisted of males both with and without colour. The absence or presence of breeding colour was a good predictor of testes investment. Compared to males with breeding colour, males without colour were smaller in body size but had extraordinarily large testes. In absolute terms, testes mass of males without breeding colour was on average 3·4 times greater than those of males with breeding colour. Since small colourless males are known to reproduce as sneaker males, this heavy investment in testes probably reflects that they are forced to spawn under sperm competition. Contrary to testes size, sperm‐duct glands were largest among males with breeding colour. These glands produce mucins used for making sperm‐containing mucous trails that males place in the nest before and during spawning. Since both sneakers and nest‐holders potentially could benefit from having large glands, this result is intriguing. Yet, high mucus production may be more important for nest‐holders, because it also protects developing embryos from infections. There was no significant effect of season on body size, testes or sperm‐duct glands size, but colourless males tended to be less common late in the season. Possibly this may indicate that individual small colourless males develop into their more colourful counterparts within the breeding season.     

Seasonal changes in the distribution and composition of common seal (Phoca vitulina) haul-out groups

Seasonal changes in the distribution and composition of common seal haul–out groups were followed in a study area in Orkney, Scotland. A marking programme was also undertaken, using both conventional and radio–tags, to study individual movements between sites and seasonal changes in site–use. Certain haul–out sites were used only in the breeding season, while others were used during the winter. Seals were seen at one site all year round and at another during only the pre–pupping and moult period. On one island where two sites were used during the summer, there were significant differences in the sex ratio of groups at the two sites: at one site males predominated and few pups were seen; on another, nearby, mothers and pups were regularly seen, although the site was also used by males. There was also evidence for segregation of the sexes outside the breeding season. Repeated observations of marked seals showed that seals used several different haul–out sites throughout the year, and that the seasonal changes in abundance at different sites resulted from individual changes in site–use. These changes in site–use are discussed in relation to feeding movements, breeding requirements and the physical characteristics of different sites.