Diving depths of Shags Phalacrocorax aristotelis breeding on the Isle of May

Information on maximum dive depth and the time spent at various depths was obtained from 49 Shags Phalacrocorax aristotelis. On average, the maximum dive depth was 33–35 m; the overall maximum was 43 m. Shags dived repeatedly to the same depth and spent c. 55% of the time between 25 and 34 m which indicated that they were foraging close to the seabed. About 46% of the time underwater was spent foraging and 54% travelling. Average underwater swimming speed was 1.7-1.9 m per second.     

Anisakid infection in the European shag Phalacrocorax aristotelis aristotelis

A total of 100 specimens of the European shag Phalacrocorax aristotelis aristotelis were examined for anisakid infections in the south-east North Atlantic. The taxonomic status of the anisakid nematodes was studied by structural analysis using light and scanning electron microscopy. Two species, Contracaecum septentrionale and C. rudolphii A (Nematoda: Ascaridoidea), occurred in the European shag with high values of prevalence and mean intensities of infection. These constitute new host records for both parasites, widening their known geographical distribution. Although the histopathology associated with the infections indicated that the anisakids did not have the potential to cause bird death, the parasites are capable of contributing to and hastening avian mortality in the case of heavily-parasitized cormorants and when stressed by xenobiotics and/or through a synergetic effect by other bioagressors.     

Foraging range and feeding locations of Shags Phalacrocorax aristotelis during chick rearing

We used radio-tracking techniques to determine the foraging range and feeding locations used by individual Shags Phalacrocorax aristotelis during chick rearing on the Isle of May, Scotland. The mean (±s.d.) foraging range was 7.o±1.9 km (maximum 17 km) and there were no significant sex or year differences. All feeding sites were within 7 km of land. Most (> 90%) were either within 2 km of the colony or in two discrete areas 5–13 km to the north and west. Use of the areas varied between years with both males and females making more use of the north area in 1987 than either 1988 or 1989. Birds used > 11% of the area of sea within the maximum recorded feeding range. This restricted distribution appeared to be related to water-depth and bottom sediment type. Shags fed most frequently in water 21–40 m deep, with a bottom of either gravel and sand, or rock with thin patchy sediment cover.     

Dietary changes of Mediterranean Shags Phalacrocorax aristotelis desmarestii between the breeding and post-breeding seasons in the upper Adriatic Sea

Capsule?Shags move between breeding and non-breeding areas and this is associated with a significant change in diet.

Aims?To determine whether the diet of Shags nesting on islets off the Croatian coast is the same as their diet after the post-breeding move to the Gulf of Trieste.

Methods?Diet was determined by the analysis of 611 regurgitated food pellets.

Results?A total of 23 988 prey items were identified in the sample of pellets. Post-breeding Shags in the Gulf of Trieste focused on demersal and relatively immobile Gobiidae (81.5% by number, 87.1% by biomass). The most frequent prey species was Gobius niger (70.8% by number). In the breeding season at Oruda island, Croatia, the diet was more varied. Breeding Shags fed on bentho-pelagic, mobile prey such as Atherina boyeri (28.4% in frequency), Serranus hepatus (16.1%) and Crenilabrus tinca (12.0%), while Gobiidae had a dietary frequency of only 18.1%. With respect to biomass the most important prey were Crenilabrus tinca (19.0%) and Serranus hepatus (18.4%).

Conclusion?We suggest that the movement of Shags within the Adriatic Sea is driven by dietary requirements. Most previous studies of Shag diet have shown that Shags tend to have a more specialized diet during the breeding season, concentrating upon demersal prey species. However, we have found that birds breeding at the Croatian study colony show dietary diversity. We suggest that lack of dietary specialization is a facultative response to local prey abundance, and is probably the result of over-fishing of demersal species in the areas around the breeding locations in which the birds find suitable sites and are little disturbed by human activity. Shags may move immediately after breeding to the Gulf of Trieste because demersal species are likely to be more abundant there. As a consequence, the diet becomes more specialized and is then more similar to the diet of other populations of Shags.     

Factors influencing food-load sizes brought in by Shags Phalacrocorax aristotelis during chick rearing

Weights of food loads brought back to the colony by 26 Shags Phalacrocorax aristotelis rearing chicks were determined using the water-offloading procedure. Loads consisted almost entirely of lesser sandeels Ammodytes marinus. Load size was extremely variable, ranging from 8 to 208 g with a mean load weight of 106 g. Data on foraging behaviour collected concurrently demonstrated that Shags brought back heavier loads when they were feeding farther away from the colony and when brood biomass was larger. These two variables together explained 70.3% of the variation in load size. We postulate that adults had already digested their own food requirements by the time they arrived back at the nest, and the contents of the loads were therefore primarily for the young. On 67% of trips, Shags caught, on average, more than one fish per dive. This estimate ignores the food requirements of the adult, and therefore true prey-capture rates must have been higher.     

Philopatry and colony fidelity of Shags Phalacrocorax aristotelis on the east coast of Britain

Shag Phalacrocorax aristotelis colonies from the Firth of Forth south to the Fame Islands (SE Scotland and NE England) were surveyed in 1982 and 1983 for nesting Shags ringed as chicks or adults in the previous 20 years. On average, 5% of 863 Shags ringed as chicks were breeding away from their natal colony, and only 1% of 401 marked adults had left their breeding colony. The proportion p of chicks that settled at or beyond distance D from the natal colony followed the relationship p = 0.263D-^0.771, implying that more Shags moved 10 km or beyond to breed than expected from diffusion models or a constant-rate dispersal model; adjusted for birds that settled outside the study area, the relationship was p = 0.314D^-0771. There was no effect of colony size on emigration rate, but females showed a greater tendency to settle away from the natal colony than did males. At the natal colony, more males (69%) than females (47%) nested within 300 m of their birthplace. Two mechanisms appear to determine where a Shag settles to breed: a navigational one governing the return to the natal colony, plus the competitive process of nest-site establishment.     

Age-related effects of a nonbreeding event and a winter wreck on the survival of Shags Phalacrocorax aristotelis

During 1993, about half the adult Shags Phalacrocorax aristotelis alive on the Isle of May, southeastern Scotland, failed to breed. The proportion not breeding was highest (66%) for individuals aged 14–21 years and lowest (12%) for those aged 3–6 years. In February 1994, a prolonged spell of onshore winds resulted in a wreck of these adults such that overwinter survival was reduced from the normal 88% to 14.7%. The age-specific pattern of survival over this winter differed from previous years, with birds aged less than 7 years and more than 16 years surviving less well and middle-aged individuals surviving better than expected. Whether an individual had bred or not in 1993 had little effect on the chances of survival of the youngest and oldest individuals, as most died. However, among Shags aged 12–15 years, those which had bred survived markedly less well than those which had not, indicating that there had been some cost involved to birds in breeding in 1993.     

Estimating dispersal distributions at multiple scales: within‐colony and among‐colony dispersal rates,distances and directions in European Shags Phalacrocorax aristotelis

Knowledge of the rate, distance and direction of dispersal within and among breeding areas is required to understand and predict demographic and genetic connectivity and resulting population and evolutionary dynamics. However dispersal rates, and the full distributions of dispersal distances and directions, are rarely comprehensively estimated across all spatial scales relevant to wild populations. We used re‐sightings of European Shags Phalacrocorax aristotelis colour‐ringed as chicks on the Isle of May (IoM), UK, to quantify rates, distances and directions of dispersal from natal to subsequent breeding sites both within IoM (within‐colony dispersal) and across 27 other breeding colonies covering 1045 km of coastline (among‐colony dispersal). Additionally, we used non‐breeding season surveys covering 895 km of coastline to estimate breeding season detection probability and hence potential bias in estimated dispersal parameters. Within IoM, 99.6% of individuals dispersed between their natal and observed breeding nest‐site. The distribution of within‐colony dispersal distances was right‐skewed; mean distance was shorter than expected given random settlement within IoM, yet some individuals dispersed long distances within the colony. The distribution of within‐colony dispersal directions was non‐uniform but did not differ from expectation given the spatial arrangement of nest‐sites. However, 10% of all 460 colour‐ringed adults that were located breeding had dispersed to a different colony. The maximum observed dispersal distance (170 km) was much smaller than the maximum distance surveyed (690 km). The distribution of among‐colony dispersal distances was again right‐skewed. Among‐colony dispersal was directional, and differed from random expectation and from the distribution of within‐colony dispersal directions. Non‐breeding season surveys suggested that the probability of detecting a colour‐ringed adult at its breeding location was high in the northeastern UK (98%). Estimated dispersal rates and distributions were therefore robust to incomplete detection. Overall, these data demonstrate skewed and directionally divergent dispersal distributions across small (within‐colony) and large (among‐colony) scales, indicating that dispersal could create genetic and demographic connectivity within the study area.     

Immediate and delayed effects of a gale in late spring on the breeding of the Shag Phalacrocorax aristotelis

At the beginning of May 1982, a westerly gale destroyed or damaged 49% of 761 Shag Phalacrocorax aristotelis nests on the exposed west side of the Isle of May, southeast Scotland; 518 nests on the sheltered east side acted as controls against which to assess subsequent effects of the gale on Shag reproduction and recruitment.
On sites affected by the gale, 90% of pairs rebuilt their nest and laid a second clutch, on average 18 days after the loss of the first one. First-time breeders nested closer to the sea, suffered greater damage to their nests, and deserted their damaged nests more readily after the gale than did experienced ones; a quarter of potential recruits to the west side were deterred from breeding in that year.
By the end of the season, the number of chicks fledged per pair was 31% lower on the west side than on the east. The difference was caused partly by nest desertion, partly by greater hatching failure of full clutches in apparently undamaged nests and especially by lower fledging success of pairs that renested. Irrespective of breeding experience, delayed early breeders produced more chicks than late breeders nesting at the same time, showing that late breeders were of poorer intrinsic quality than early breeders. This was one reason for a seasonal decline in Shag productivity, although environmental factors also played a role.
During April-July, gales like that of May 1982 occur on average once every 5 years on the Isle of May. Despite the disruption that they may cause to nesting Shags, the impact on the long-term growth of the colony is small.     

Influence of body composition on the metabolic rate of nestling European shags (Phalacrocorax aristotelis)

During the early development of avian nestlings, their mass-specific resting metabolic rate (RMR) changes in a biphasic pattern with the peak value often being much higher than that expected for an adult bird of similar body mass. In the present study we examined the possible influence of variations in the size of internal organs in “setting” the high RMR and peak metabolic rate (PMR) during development in a large altricial species, the European shag (Phalacrocorax aristotelis). Thermoneutral RMR and cold-exposure induced PMR were measured in nestlings 15 days old, the age at which the highest RMR occurred during development. Body mass averaged 414 g. Mean values of RMR and PMR were 5.75 W and 9.08 W, respectively; the RMR value corresponds to approximately 250% of the expected value for an adult non-passerine bird of similar body mass. The masses of all the organs measured (breast and leg muscles, heart, liver, intestine, and kidney) varied isometrically with total body mass. However, large chicks had a significantly lower fractional water content than small chicks, suggesting that the former had achieved a higher level of functional maturity. In contrast to what has been suggested for adult birds in general, the heart and kidney masses of shag nestlings were not significantly correlated with the metabolic rates. The intestine length, in contrast, was highly and positively correlated with both the RMR and the PMR, i.e. intestine length was a better predictor of RMR and PMR than was total body mass. In addition, liver mass was positively correlated with RMR. The results of the present study suggest that the liver in particular may play a key role in establishing the high, mass-specific RMR which is attained during development in bird chicks. Our results also support previous suggestions that early in their development, altricial chicks mainly allocate energy to the growth of `energy-processing' organs (such as the intestine and liver) rather than to `energy-consuming' organs. Accepted: 3 March 1999     

Weak large‐scale population genetic structure in a philopatric seabird,the European Shag Phalacrocorax aristotelis

Quantifying population genetic structure is fundamental to testing hypotheses regarding gene flow, population divergence and dynamics across large spatial scales. In species with highly mobile life‐history stages, where it is unclear whether such movements translate into effective dispersal among discrete philopatric breeding populations, this approach can be particularly effective. We used seven nuclear microsatellite loci and mitochondrial DNA (ND2) markers to quantify population genetic structure and variation across 20 populations (447 individuals) of one such species, the European Shag, spanning a large geographical range. Despite high breeding philopatry, rare cross‐sea movements and recognized subspecies, population genetic structure was weak across both microsatellites and mitochondrial markers. Furthermore, although isolation‐by‐distance was detected, microsatellite variation provided no evidence that open sea formed a complete barrier to effective dispersal. These data suggest that occasional long‐distance, cross‐sea movements translate into gene flow across a large spatial scale. Historical factors may also have shaped contemporary genetic structure: cluster analyses of microsatellite data identified three groups, comprising colonies at southern, mid‐ and northern latitudes, and similar structure was observed at mitochondrial loci. Only one private mitochondrial haplotype was found among subspecies, suggesting that this current taxonomic subdivision may not be mirrored by genetic isolation.     

Nutritional needs for egg formation in the Shag Phalacrocorax aristotelis

Egg production does not impose a major food need in the Shag Phalacrocorax aristotelis because the eggs are small, are formed slowly and are laid at 3-day intervals. I used dye-dosing of females, laying time and, after fixing and staining the yolk, daily ring counts to estimate the amounts of protein and energy needed each day to produce a clutch of three eggs. Maximum daily nutrient needs during egg formation were only 1.15 g per day additional protein and 34 kJ per day energy. Yolk formation times of 40 eggs were 13.5 (±1.2 s.d.) days. Based on yolk-ring counts and laying dates of 15 eggs that contained dye, the lag period between yolk completion and laying was 3.1 ± 0.06 days. The distinct light and dark rings of the stained yolk resulted from differences in the transparency of the yolk spheres. In the dark rings, the spheres were relatively clear, so more depth of stained yolk could be seen than in the light rings, which reflected the incident light.     

Strong temporal consistency in the individual foraging behaviour of Imperial Shags Phalacrocorax atriceps

Individual consistency in foraging behaviour can generate behavioural variability within populations and may, ultimately, lead to species diversification. However, individual‐based long‐term behavioural studies are particularly scarce in seabird species. Between 2008 and 2011, breeding Imperial Shags Phalacrocorax atriceps at the Punta León colony, Argentina, were tracked with GPS devices to evaluate behavioural consistency during their foraging trips. Within a breeding season, individuals were highly consistent in the maximum distances they reached from the shore and the colony, as well as in the time invested in flight and diving across consecutive days during early chick rearing. In addition, each individual had its specific foraging area distinct from the foraging area of other individuals. Comparing between early and late chick rearing in the same season, individuals were consistent, to a lesser degree, in the maximum distance they reached from the colony and the shore, increasing in consistency later on in the season. Within the season, females were more consistent than males in the maximum distance they moved from the colony and the shore, the sexes segregated in their foraging areas and individual females were segregated from one another. Twenty‐eight individuals tracked in different breeding seasons were marginally consistent in their trip durations and maximum distance reached from shore across seasons. Among seasons, foraging locations differed between sexes and among individual females. Individuals from this colony exhibited consistency over time in several aspects of foraging behaviour, which may be due to a combination of individual characteristics such as learning abilities, breeding experience or health, as well as targeted prey type and stability of the environment at this location.     

From cradle to early grave: juvenile mortality in European shags Phalacrocorax aristotelis results from inadequate development of foraging proficiency

In most long-lived animal species, juveniles survive less well than adults. A potential mechanism is inferior foraging skills but longitudinal studies that follow the development of juvenile foraging are needed to test this. We used miniaturized activity loggers to record daily foraging times of juvenile and adult European shags Phalacrocorax aristotelis from fledging to the following spring. Juveniles became independent from their parents 40 days post-fledging. They compensated for poor foraging proficiency by foraging for approximately 3 h d(-1) longer than adults until constrained by day length in early November. Thereafter, juvenile foraging time tracked shortening day length up to the winter solstice, when foraging time of the two age classes converged and continued to track day length until early February. Few individuals died until midwinter and mortality peaked in January-February, with juvenile mortality (including some of the study birds) five times that of adults. In their last two weeks of life, juveniles showed a marked decline in foraging time consistent with individuals becoming moribund. Our results provide compelling evidence that juveniles compensate for poor foraging proficiency by increasing foraging time, a strategy that is limited by day length resulting in high winter mortality.     

Breeding performance and timing of breeding of inland and coastal breeding Cormorants Phalacrocorax carbo in England and Wales

Capsule Breeding performance was higher at inland colonies than on the coast due to higher nestling survival during late development.

Aims To compare breeding performance of inland and coastal breeding Cormorants in England and Wales and to provide breeding performance data for Cormorants for future demographic studies.

Methods Breeding performance and timing of breeding was monitored at six inland and four coastal colonies during 1997 and 1998. We compare clutch and brood size and egg and nestling survival.

Results Breeding performance was higher at inland colonies because of higher nestling survival during the later stages of nestling development, from 15–56 days.

Conclusions Greater and earlier food availability at inland colonies, resulting in earlier and more protracted breeding is the most probable explanation for differences in breeding performance. A more protracted breeding season would reduce competition for food and enhance breeding performance, which may be especially apparent during late chick development when energy demands are greatest.     

Extra-pair paternity in the shag, Phalacrocorax aristotelis as determined by DNA fingerprinting

The frequency of chicks resulting from extra-pair copulation in the shag, Phalacrocorax aristotelis , was measured by DNA fingerprinting. DNA fingerprints were taken from both sexes of 15 pairs and their chicks (28) in a subcolony on the Isle of May, UK. It was found that 18% of the chicks had extra-pair paternity, and one chick (3.5%) was not the offspring of either member of the pair, implying either a polygynous male whose second female was fertilized by another male or adoption. Although no observations of courtship and copulation were made in the same season, observations in a previous year on the same colony of shags showed that 14.1% of the copulations by males were not with the female with whom that male reared young. The similarities and differences are discussed between these results on the level of extra-pair copulations and of extra-pair paternity and those of other studies where both observations of extra-pair copulations and various measures of the degree of extra-pair paternity have been made.     

Sexual size dimorphism and male combat in snakes

Summary This paper reviews published literature on snakes to test the hypothesis that large male size, relative to female size, evolves because of the advantage it confers in male combat. Analysis of the data reveals a high correlation between the occurrence of male combat, and sexual dimorphism in which the male is the larger sex. This correlation holds (i) within the total sample of snake species (n=224), (ii) within the family Colubridae (n=134), and (iii) in a comparison between the eight families of snakes for which data are available. These results strongly support the hypothesis that large male size is an adaptation to intrasexual competition. The analysis also shows that females are larger than males in about 66% of snake species, that male combat is known in only about 15% of species, and that both sexual size dimorphism and the incidence of male combat tend to be distributed along taxonomic lines.