Goshawk predation during winter, spring and summer in a boreal forest area of central Sweden

Predation by goshawks was studied in a central Swedish boreal forest area. Data were collected in winter (January–February) 1977-81 by tracking radio-tagged goshawks, and in the breeding season (April–July) by collecting prey remains at the nest. In the breeding season birds dominated the prey, amounting to 86% of prey number and 91% of prey biomass. Wood pigeon Columba palumbus , black grouse Tetrao tetrix , hooded crow Corvus corone cornix and jay Garrulus glandarius accounted for more than 50% of the prey animals, whereas capercaillie Tetrao urogallus and black grouse accounted for more than 50% of prey biomass. There was no functional response to black grouse density fluctuations. Every year goshawks killed significantly more females than males of both capercaillie and black grouse, due to high vulnerability of the grouse hens while laying and incubating. It was estimated that during spring and early summer goshawk predation removed 25% of the female, and 14% of the male black grouse population. In winter squirrel was the dominating prey, both in terms of number (79%) and weight (56%). The proportion of squirrel in the diet was equally high both in winters of low and high squirrel density. The high proportion of squirrel in the winterdiet, as compared to the breeding season, is believed to be due to squirrels having to accept an increased predation risk in winter, in order to feed efficiently enough.     

Management of the Nile perch, Lates niloticus fishery in Lake Victoria in light of the changes in its life history characteristics

Nile perch was introduced into lakes Victoria and Kyoga in the 1950s and 1960s from Lake Albert. The changes in prey eaten and the life history characteristics of Nile perch in lakes Victoria and Kyoga from the 1960s to 1990s were examined and compared with Lake Albert. The dominant prey eaten changed from haplochromines, Caridina nilotica, Rastrineobola argentea, and Nile perch juveniles. The condition factor deteriorated from 1.4 in 1960s to 1.2 in 1990s, compared with 1.3 in Lake Albert suggesting a reduction in food supply. Therefore, exploitation of Nile perch prey should be controlled. The size at first maturity increased from 30–40 to 40–50 cm and 50–59 to 80–100 cm for males and females, which is similar to Lake Albert. Sex ratios decreased from 85–100 to 20–65 females for every 100 males suggesting that Nile perch had less capacity to replenish its stocks such that breeding females should be protected. As males mature at 50–55 cm and females at 80–100 cm, immature males of <50 cm and breeding females of >100 cm could be protected through size selective exploitation of fish of 50–100 cm using gillnets of 127–254 mm.     

Seasonal changes in territorial behaviour of hazel grouse ( Bonasa bonasia) in a temperate forest of South Korea

We studied the seasonal changes in territorial behaviour of hazel grouse (Bonasa bonasia) from December 1999 to July 2001 in a temperate forest of South Korea. The study was based on the use of radio telemetry. Territory size and degree of overlap increased after the breeding period, and index of advertisement and border disputes decreased. In spring, the territory was defended by both males and females. During autumn and winter, this behaviour was relaxed to such a degree that hazel grouse cannot be considered territorial, but as forming a home range.     

Diet composition of smooth-hound shark, Mustelus mustelus (Linnaeus, 1758), in the Gulf of Gabès, southern Tunisia

The diet of the smooth-hound shark, Mustelus mustelus , from the Gulf of Gabès (southern Tunisia, central Mediterranean Sea) was investigated with respect to fish size and season. Stomach contents were analyzed from 540 specimens with total lengths ranging from 34 to 158.5 cm. Of the total number of stomachs examined, 63 were empty (11.67%). Smooth-hound shark fed mainly on crustaceans, fishes and cephalopods. Sipunculids, polychaetes and echinoderms were occasional preys. No differences were found between the diets of males and females. Ontogenetic changes in diet of M. mustelus were apparent, with crustaceans forming a greater proportion of the diet of smaller sharks. Both teleosts and molluscs increased in importance with increasing shark size. Consumptions of polycheates, sipunculids and echinoderms were not related to predator size. Prey diversity also increased with size, with large and mobile prey species found more commonly in the diet of larger sharks. The limited overlap in the dietary compositions of juveniles, subadults and adults suggests the possibility of resource partitioning. Seasonality in food habits was in accordance with the dynamics of the predator and the prey species.     

Effects of season and body-size on the distribution and diet of the Iberian chub Leuciscus pyrenaicus in a lowland catchment

Seasonal and ontogenetic variations in space and food use by Leuciscus pyrenaicus were analysed in a Portuguese lowland catchment. Large fish occurred mostly in deep permanent-flowing sites whereas small fish occurred mostly in the shallowest intermittent-flowing site. No seasonal or size-related changes in feeding intensity were found, but the diet changed both across seasons and throughout ontogeny. The diet was dominated by aquatic prey over all seasons, but during winter and summer more plant material and terrestrial prey, respectively, were eaten. Throughout ontogeny fish shifted from soft-bodied to hard-shelled prey and decreased animal prey breadth. Mean prey size increased with fish size but the prey size spectrum was more variable for medium-sized fish than for either small or large fish. It is suggested that: (i) large fish avoid shallow drying areas owing to the risk of mortality, either by thermal and respiratory stresses or increased predation by mammals and birds; (ii) seasonal changes in diet are a response to differences in prey availability; and (iii) morphological constraints, prey handling costs and habitat partitioning are responsible for size-related changes in diet.     

Prey choice by marten during a decline in prey abundance

Summary We examined variation in diet choice by marten (Martes americana) among seasons and between sexes and ages from 1980–1985. During this period prey populations crashed simultaneously, except for ruffed grouse (Bonasa umbellus) which was common at the beginning and end of the study, and masked shrews (Sorex cinereus) which were abundant in 1983. Marten were catholic in selection of prey and made use of most available mammalian prey, ruffed grouse, passerine birds, berries, and insects. Diet niche was widest during the latter three years when prey was scare, particularly in late winter. Diet niche breadth was negatively correlated with abundance of all common prey species. Proportion of small prey species in the diet was correlated with absolute abundance of those species, but proportion of some large prey was related to their relative abundance. Diet choice varied among years and among seasons. Berries and insects were common in summer diets while large prey, particularly varying hare (Lepus americanus), were more frequent in winter diet than in summer diet. We found little evidence that any small mammal species was a preferred prey. Sexual size dimorphism between the sexes did not affect prey choice, nor did age. Reduced foraging effort in winter resulted in a wider diet niche only when prey was scarce. The only prediction of optimal foraging models fully supported by our data was a wider diet niche with reduced prey abundance. However, among the three most profitable prey species choice was dependent on the absolute abundance of the most profitable type (varying hare). We suggest that marten primarily forage for large prey but employ a strategy which results in encounters with small prey as well. These small prey are eaten as they provide energy at minimal cost, between captures of large prey.     

Survival, ranging, habitat choice and diet of the Northern Goshawk Accipiter gentilis during winter in Northern Finland

The winter ecology of the Northern Goshawk Accipiter gentilis was studied in northern Finland in 1991-95. The data consist of 26 radiomarked hawks, of which 331 day and 299 night positions were located. Average range size was 9894 ha for males (minimum convex polygon, n = 4) and 6282 ha for females ( n = 11). Ranges calculated by the 75% harmonic mean contour gave range areas of 3283 ha for males ( n = 3) and 2753 ha for females ( n = 11). Harmonic mean centres concentrated near human settlements, implying better food availability there compared with woodland areas. Some Goshawks preyed upon animals living in the city dump in late winter. Goshawks preferred deciduous and mature conifer forests and avoided open areas such as fields and bogs. They also avoided very heterogeneous sites. The winter diet consisted mostly of Mountain Hares Lepus timidus (30.9%), Red Squirrels Sciurus vulgaris (23.6%), Brown Rats Rattus norvegicus (12.7%) and forest grouse Tetraonidae (27.1%), of which Black Grouse Tetrao tetrix was the most important (14.5%). By weight, hares constituted 70% of the biomass consumed.     

Inter-decadal patterns of population and dietary change in sea otters at Amchitka Island, Alaska

After having been hunted to near-extinction in the Pacific maritime fur trade, the sea otter population at Amchitka Island, Alaska increased from very low numbers in the early 1900s to near equilibrium density by the 1940s. The population persisted at or near equilibrium through the 1980s, but declined sharply in the 1990s in apparent response to increased killer whale predation. Sea otter diet and foraging behavior were studied at Amchitka from August 1992 to March 1994 and the data compared with similar information obtained during several earlier periods. In contrast with dietary patterns in the 1960s and 1970s, when the sea otter population was at or near equilibrium density and kelp-forest fishes were the dietary mainstay, these fishes were rarely eaten in the 1990s. Benthic invertebrates, particularly sea urchins, dominated the otter’s diet from early summer to mid-winter, then decreased in importance during late winter and spring when numerous Pacific smooth lumpsuckers (a large and easily captured oceanic fish) were eaten. The occurrence of spawning lumpsuckers in coastal waters apparently is episodic on a scale of years to decades. The otters’ recent dietary shift away from kelp-forest fishes is probably a response to the increased availability of lumpsuckers and sea urchins (both high-preference prey). Additionally, increased urchin densities have reduced kelp beds, thus further reducing the availability of kelp-forest fishes. Our findings suggest that dietary patterns reflect changes in population status and show how an ecosystem normally under top-down control and limited by coastal zone processes can be significantly perturbed by exogenous events. Received: 23 March 1999 / Accepted: 19 January 2000     

Golden Eagles Aquila chrysaetos: land use and food in northeast Scotland

This paper contrasts changes in breeding numbers and breeding success of Golden Eagles Aquila chrysaetos on four areas with different land uses. On land primarily used for deer stalking but also for grouse shooting, and supporting abundant prey, the number of eagle pairs was steady in 1944-80. On deer land with fewer prey, the number of pairs declined greatly in the 1960s, when deer carrion became scarcer following increased shooting of red deer. On grouse moors incorporating deer stalking, the number of pairs declined in the 1950s due to persecution by gamekeepers, and then largely recovered as this lessened. On grouse moors with little or no stalking, the number of pairs fell after 1946 and remained low due to persecution, which has continued since then. Eagles on grouse moors bred poorly due to persecution. On deer land they were seldom persecuted by estate staff, and bred well. On an area of deer land, the mean annual number of young reared per undisturbed clutch in summer was related to the estimated weight of prey in the spring of the same year. The eagles have remained fairly pesticide-free, and bred well in 1963-65 when more contaminated birds in west Scotland bred poorly.     

The predatory behavior of wintering <Emphasis Type="Italic">Accipiter</Emphasis> hawks: temporal patterns in activity of predators and prey

Studies focused on how prey trade-off predation and starvation risk are prevalent in behavioral ecology. However, our current understanding of these trade-offs is limited in one key respect: we know little about the behavior of predators. In this study, we provide some of the first detailed information on temporal patterns in the daily hunting behavior of bird-eating Accipiter hawks and relate that to their prey. During the winters of 1999–2004, twenty-one sharp-shinned hawks (A. striatus) and ten Cooper’s hawks (A. cooperii) were intensively radio tracked in rural and urban habitats in western Indiana, USA. Cooper’s hawks left roost before sunrise and usually returned to roost around sunset, while sharp-shinned hawks left roost at sunrise or later and returned to roost well before sunset. An overall measure of Cooper’s-hawk-induced risk (a composite variable of attack rate and activity patterns) generally reflected the timing of prey activity, with peaks occurring around sunrise and sunset. In contrast, risk induced by the smaller sharp-shinned hawk did not strongly reflect the activity of their prey. Specifically, an early morning peak in prey activity did not correspond to a period with intense hawk activity. The lack of early morning hunting by sharp-shinned hawks may reflect the high risk of owl-induced predation experienced by these hawks. The net effect of this intraguild predation may be to “free” small birds from much hawk-induced predation risk prior to sunrise. This realization presents an alternative to energetics as an explanation for the early morning peak in small bird activity during the winter.     

Sexual dimorphism of body and relative head sizes in neonatal common garter snakes

Morphological and behavioral differences between sexes are commonplace throughout the animal kingdom. Body size is one of the most obvious sex differences frequently found in snakes. However, the developmental origins of size differences in many species, including snakes, are not well known. We examined post-natal variation in sexual size dimorphism in garter snakes Thamnophis sirtalis . The weights, body and tail lengths, and head sizes of male and female neonates born to mothers collected from ecologically dissimilar habitats on Beaver Island, Lake Michigan were compared. Sexual size dimorphism was prominent. Overall, males had significantly longer bodies and tails than females. Females were significantly heavier and had larger heads than male snakes. Maternal site affected head but not body measurements, perhaps due to differences in prey availability. The body condition of maternal females predicted neonatal body length. Significant litter variation suggests heritable variation in morphological traits possibly correlated with feeding success and survival.     

Diet-shifts by an estuarine goby (Pomatoschistus microps) in the face of variable prey availability

Ontogenetic shifts in diet provide a mechanism for maximising fitness throughout development and are common where predators exhibit large increases in size. In order to maximise their fitness throughout development, benthic feeding fish can show diet-shifts that centre on the transition from meiofaunal to macrofaunal prey. Here we assessed whether such a shift was influenced by natural variation in prey-size availability by comparing the sizes of prey consumed by naturally foraging common gobies (Pomatoschistus microps). We tested explicitly for the presence of an ontogenetic shift by analysing the length of prey consumed and an index for prey importance for gobies of different lengths. We also tested the match between actual diets and those predicted by a foraging model. The goby size at which the diet-shift occurred was consistent among locations that differed in their availability of prey and through temporal changes in densities and types of prey. The mean sizes of ingested prey increased for gobies > 35 mm in length and the relative importance of macrofauna increased at 30 mm. The foraging model predicted that gobies > 30 mm would eat larger prey than would smaller gobies which differed from the observed changes in prey-size at 35 mm. Availability of prey did not appear to influence the lengths at which gobies changed diet but did affect the size of prey taken after the diet-shift. A relatively large abundance of large-bodied chironomids at two sites was reflected in the mean size of prey consumed by gobies > 30 mm at these sites. Our study indicates that intrinsic mechanisms can be more important than fluctuating environments in determining prey-choice and shifts in diet, although for the common goby, variability in prey-size may have implications for prey-choice later in ontogeny.     

Comparison of wing morphology in three birds of prey: correlations with differences in flight behavior

Flight is the overriding characteristic of birds that has influenced most of their morphological, physiological, and behavioral features. Flight adaptations are essential for survival in the wide variety of environments that birds occupy. Therefore, locomotor structure, including skeletal and muscular characteristics, is adapted to reflect the flight style necessitated by different ecological niches. Red-tailed hawks (Buteo jamaicensis) soar to locate their prey, Cooper's hawks (Accipiter cooperii) actively chase down avian prey, and ospreys (Pandion haliaetus) soar and hover to locate fish. In this study, wing ratios, proportions of skeletal elements, and relative sizes of selected flight muscles were compared among these species. Oxidative and glycolytic enzyme activities of several muscles were also analyzed via assays for citrate synthase (CS) and for lactate dehydrogenase (LDH). It was found that structural characteristics of these three raptors differ in ways consistent with prevailing aerodynamic models. The similarity of enzymatic activities among different muscles of the three species shows low physiological differentiation and suggests that wing architecture may play a greater role in determining flight styles for these birds.     

Diet variation of common buzzards in Finland supports the alternative prey hypothesis

The regional synchrony of short-term population fluctuations of small rodents and small game has usually been explained by varying impacts of generalist predators subsisting on both voles and small game (the "alternative prey hypothesis" APH). APH says that densities of predators increase as a response to increasing vole densities and then these predators shift their diet from the main prey to the alternative prey when the main prey decline and vice versa. We studied the diet composition of breeding common buzzards Buteo buteo during 1985-92 in western Finland. Microtus voles were the main prey and water voles, shrews, forest grouse, hares and small birds the most important alternative prey. Our data from the between-year variation in the diet composition of buzzards fulfilled the main predictions of APH. The yearly proportion of main prey (Microtus voles) in the diet was higher in years of high than low vole abundance. The proportion of grouse in the diet of buzzards was negatively related to the abundance of Microtus voles in the field and was nearly independent of grouse abundance in the field. In addition, buzzards mainly took grouse chicks and young hares which is consistent with the prediction of APH. Therefore, we conclude that buzzards are able to shift their diet in the way predicted by the APH and that buzzards, together with other generalist predators, may reduce the breeding success of small game in the decline phase of the vole cycle, and thus substantially contribute to the existence of short-term population cycles of small game.     

DISPERSION, BREEDING AND PREY OF THE HEN HARRIER CIRCUS CYANEUS IN GLEN DYE, KINCARDINESHIRE

Hen Harriers were studied on a grouse moor in northeastern Scotland from 1970-74. Numbers in spring were fairly stable, and the 24 nesting territories occupied in 1974 were regularly dispersed with a mean nearest neighbour distance of 1.52 ± s. e. 0.09 km. Many pairs apparently failed to breed and left the area. The mean clutch size of those remaining was 4.70 ± s. e. 0.24 (n= 27) and the mean number of young fledged per successful nest was 3.11 ± s. e. 0.26 (n= 19). Young males were more likely to disperse far from the study area than young females. The variety of prey species observed from hides at five nests was very limited, and comprised mainly pipits, grouse chicks and lagomorphs. Estimates by weight suggest that grouse and lagomorphs accounted for 89% of all prey. Males brought more small items and fewer large items to nests than females, and on average provided 72% of all prey items seen during watches from the hides. On the basis of studies of prey at three nests in 1974, and counts of grouse in spring and late summer, harrier predation was estimated to have reduced the number of grouse which might otherwise have survived to late July that year by a maximum of 7.4%.     

SEXUAL SELECTION IN THE BARN SWALLOW (HIRUNDO RUSTICA). IV. PATTERNS OF FLUCTUATING ASYMMETRY AND SELECTION AGAINST ASYMMETRY

The patterns of variation in fluctuating asymmetry were studied in four morphological characters of the barn swallow Hirundo rustica. The level of absolute and relative asymmetry was larger in the secondary sexual character “outer tail length” than in three nonsexual morphological traits (wing, central tail, and tarsus length). The extent of individual asymmetry in outer tail length was negatively correlated with tail-ornament size, whereas the relationship between asymmetry of all other morphological characters and their size was flat or U-shaped. Asymmetry in outer tail length was unrelated to asymmetry in other morphological characters, whereas asymmetries in the length of wing, central tail, and tarsus were positively correlated. Male bam swallows exhibited larger asymmetry in outer tail length than females. Asymmetry of most morphological traits exhibited intermediate repeatabilities between years, with the exception of male and female outer tail length, which were highly repeatable. Tail asymmetry of offspring weakly, though significantly, resembled that of their parents. Asymmetry in wing and outer tail length was also significantly related to several fitness components. Male barn swallows that acquired a mate were less asymmetric in wing and outer tail length than unmated males. Females with more asymmetrical tails laid eggs significantly later. Annual reproductive success was unrelated to fluctuating asymmetry. Male barn swallows that survived were less asymmetric in wing and outer tail length than nonsurvivors, whereas female survivors were less asymmetric in outer tail length than nonsurvivors. These results suggest that levels of fluctuating asymmetry in barn swallows are associated with differences in fitness.     

Morphological differentiation may mediate mate-choice between incipient species of Anopheles gambiae s.s

The M and S molecular forms of Anopheles gambiae s.s. have been considered incipient species for more than ten years, yet the mechanism underlying assortative mating of these incipient species has remained elusive. The discovery of the importance of harmonic convergence of wing beat frequency in mosquito mating and its relation to wing size have laid the foundation for exploring phenotypic divergence in wing size of wild populations of the two forms. In this study, wings from field collected mosquitoes were measured for wing length and wing width from two parts of the sympatric distribution, which differ with respect to the strength of assortative mating. In Mali, where assortative mating is strong, as evidenced by low rates of hybridization, mean wing lengths and wing widths were significantly larger than those from Guinea-Bissau. In addition, mean wing widths in Mali were significantly different between molecular forms. In Guinea-Bissau, assortative mating appears comparatively reduced and wing lengths and widths did not differ significantly between molecular forms. The data presented in this study support the hypothesis that wing beat frequency may mediate assortative mating in the incipient species of A. gambiae and represent the first documentation of a morphological difference between the M and S molecular forms.     

The Evolution of Reversed Sexual Size Dimorphism in Hawks,Falcons and Owls: A Comparative Study

Many hypotheses have been proposed to account for the origin and maintenance of reversed size dimorphism (RSD, females being larger than males) in hawks, falcons and owls, but no consensus has been reached. I performed comparative analyses, using both cross-taxa data and phylogenetically independent contrasts, to investigate potential correlates of reversed size dimorphism. Using a similar set of explanatory variables, covering morphology, life history and ecology, I tested whether any trait coevolved with size dimorphism in all three groups and hence provided a general explanation for the evolution of RSD. For hawks, strong correlates were found in the foraging-variable complex, so RSD might have evolved in species hunting large and agile prey. This is consistent with the intersexual-competition hypothesis (sexes have evolved different sizes to lessen intersexual competition for food), but especially the small-male hypothesis (males have evolved to be smaller to be more efficient foragers). Evolutionary pathway analyses suggest that RSD evolved most likely as a precursor of changes in hunting strategy but as a consequence of high reproduction. The falcons showed a similar pattern: species with strong RSD hunted larger and more agile prey. The evolutionary pathway analysis supported the idea that RSD evolved before the specialisation on more agile and/or larger prey. Finally for owls, the results showed clear parallels. RSD increased with prey size, consistent with the small-male hypothesis. Evolutionary pathway analysis suggests that RSD in owls has most likely evolved before specialisation on large prey, so a small and more agile male might be advantageous even when hunting small prey. These results suggest that RSD in hawks, falcons and owls evolved due to natural-selection pressures rather than sexual-selection pressures. Co-ordinating editor: J. Tuomi     

SEXUAL SELECTION IN THE MONOGAMOUS BARN SWALLOW (HIRUNDO RUSTICA). I. DETERMINANTS OF TAIL ORNAMENT SIZE

The proportion of phenotypic variance in the length of the sexually selected tail of the monogamous barn swallow Hirundo rustica that is attributable to genetic variance was studied in the field in Denmark during a seven-year period. Tail length was on average 20% greater in males than in females. Tail length correlated with wing length, but not with other morphological traits. Tail length increased with the first molt, but remained constant during subsequent years. Changes in tail length between years, owing to molt were significantly affected by sex and by degree of infection with an haematophagous mite (Ornithonyssus bursa). There were significant differences in sexual size dimorphism between years, apparently as a result of environmental conditions in the African winter quarters during molt. Tail length was a highly repeatable morphological trait, and standardization of tail length for age effects only marginally increased repeatability. Heritability of tail length as estimated from regression of values for sons on those of their fathers was 0.59. This suggests that secondary sexual traits affected by strong directional selection still may show a statistically significant heritability.     

Grouse dynamics and harvesting in Kainuu,northeastern Finland

We study the dynamics of the capercaillie, black grouse, hazel grouse and willow grouse in Kainuu game management district in northeastern Finland in the years 1989–2004. It appears that the 6–7 year periodicity that prevailed in this region from 1960s up to 1980s has now vanished in all species. The grouse data are modelled using a linear autoregressive model with lag terms for population dynamics including grouse harvest as annual bag and an index of winter severity (winter‐time area of Baltic Sea ice cover). We use the Akaike information criterion for selecting the best model for each species; first order lag is forced to the models. It turns out that a term is needed for harvesting (with a negative coefficient) in models for all species. For the capercaillie and the hazel grouse second order lag was included, for the black grouse and the willow grouse first order lag suffices. The willow grouse is the only species where the index of winter strength (with a negative coefficient) is needed in the model.