Group size and composition are correlated with population density in the group-territorial blue korhaan (Eupodotis caerulescens)

The blue korhaan (Eupodotis caerulescens) is a poorly known bird endemic to the grasslands of southern Africa. It lives in permanently held territories defended by groups of individuals. During 2001 and 2002, 61 transects were carried out along 544 km of roads in Mpumalanga province (South Africa) to collect data on territory density and group size and composition in this species. Habitat, altitude and topography variables were measured to evaluate their influence on population density. Average territory density was 1.56 ± 0.19 (SE) territories per 5 km and maximum densities attained 5.6 territories per 5 km. Within its area of occurrence, higher densities occurred along transects with larger proportion of short grassland (either burned or grazed) and flatter topography. Mean group size was 3.03 ± 0.11 birds (n = 119), with 87.4% of the groups having one to four birds. Most common group composition was one male and one female‐like (34.5%), followed by two male and one female‐like (17.2%) and two male and two female‐like (13.8%). Single individuals were always males, and the male/female‐like ratio peaked at a group size of three and decreased thereafter. Mean group size increased with territory density (r_s = 0.42, P = 0.007, n = 41). There were no differences in group size across habitat types. The occurrence of habitat saturation and cooperative breeding in this species could explain the observed results.     

Territory quality determines social group composition in Ethiopian wolves Canis simensis

1. We contrast the value of four different models to predict variation in territory size as follows: resource density (the ideal free distribution), population density, group size and intruder pressure (relative resource-holding potential). In the framework of the resource dispersion hypothesis, we test the effect of resource abundance and spatial variation in resource distribution on the age/sex composition of social groups. 2. We explore these drivers of territory size and group size/composition in Ethiopian wolves Canis simensis in the Bale Mountains, Ethiopia, using fine-scale distribution maps of their major prey species based on satellite-derived vegetation maps. 3. The number of adult males is correlated with territory size, while prey density, wolf population density and intruder pressure are not associated with territory size. On average, each additional adult male increases territory size by 1.18 km(2). 4. Prey abundance increases with territory size (average biomass accumulation of 6.5 kg km(-2)), and larger territories provide greater per capita access to prime foraging habitat and prey. 5. The age/sex composition of wolf packs is more closely related to territory quality than territory size. Subordinate adult females are more likely to be present in territories with greater proportions of prime giant molerat Tachyoryctes macrocephalus habitat (i.e. >80% of Web Valley territories and >20% in Sanetti/Morebawa), and more yearlings (aged 12-23 months) occur in territories with greater overall prey biomass. 6. Wolf packs with restricted access to good foraging habitat tend to defend more exclusive territories, having a lower degree of overlap with neighbouring packs. 7. The greater per capita access to prey in large groups suggests a strong evolutionary advantage of collaborative territorial defence in this species, although the relative costs of territorial expansion vs. exclusion depend upon the spatial distribution of resources. We propose a model whereby territory size is determined by the number of adult males, with the presence of subordinate females and yearlings dependent on the quality of habitat, and the abundance and distribution of prey, incorporated within territory boundaries.     

Avian top predator and the landscape of fear: responses of mammalian mesopredators to risk imposed by the golden eagle

Top predators may induce extensive cascading effects on lower trophic levels, for example, through intraguild predation (IGP). The impacts of both mammalian and avian top predators on species of the same class have been extensively studied, but the effects of the latter upon mammalian mesopredators are not yet as well known. We examined the impact of the predation risk imposed by a large avian predator, the golden eagle (Aquila chrysaetos, L.), on its potential mammalian mesopredator prey, the red fox (Vulpes vulpes, L.), and the pine marten (Martes martes, L.). The study combined 23 years of countrywide data from nesting records of eagles and wildlife track counts of mesopredators in Finland, northern Europe. The predation risk of the golden eagle was modeled as a function of territory density, density of fledglings produced, and distance to nearest active eagle territory, with the expectation that a high predation risk would reduce the abundances of smaller sized pine martens in particular. Red foxes appeared not to suffer from eagle predation, being in fact most numerous close to eagle nests and in areas with more eagle territories. This is likely due to similar prey preferences of the two predators and the larger size of foxes enabling them to escape eagle predation risk. Somewhat contrary to our prediction, the abundance of pine martens increased from low to intermediate territory density and at close proximity to eagle nests, possibly because of similar habitat preferences of martens and eagles. We found a slightly decreasing trend of marten abundance at high territory density, which could indicate that the response in marten populations is dependent on eagle density. However, more research is needed to better establish whether mesopredators are intimidated or predated by golden eagles, and whether such effects could in turn cascade to lower trophic levels, benefitting herbivorous species.     

Effect of competitor abundance on feeding territoriality in a grazing fish,the ayuPlecoglossus altivelis

The grazing fish, ayu,Plecoglossus altivelis Temminck & Schlegel, establishes feeding territorialiry during the young stage. The population density fluctuates from year to year by more than a hundredfold, but the determinant of territory size is less well known. The feeding territoriality of ayu was examined under simulated habitat conditions where fish density was manipulated and food resources were renewable. Fish competed for algae attached to the substrata and were divided into residents with territories, and floaters without territories. By experimental alteration of fish density the number of residents increased with density and rerritory size decreased with density. Floaters intruded into territories in a school to feed on algae, which induced overt aggression of the resident and reduced the productivity of algae growing there. Both the intruding frequency of floaters over territorial areas and their feeding pressure on algae increased at higher floater density. Floaters functioned to shift cost-benefit relationships for various territory sizes. They acted as food competitors to restrict territory size below a maximum through competitive interference. Although the growth rate of residents was inversely related to fish density, residents grew faster than floaters in each group. Under a given set of competitor abundances, economic defensibility determined territory size.     

An experimental analysis of territory size in juvenile steelhead trout

I experimentally manipulated levels of food abundance and density of competitors to determine how these factors influence the territory size of juvenile steelhead trout, Oncorhynchus mykiss. Steelhead trout were held in artificial stream channels and I followed cohorts that were fed at one of three levels of food abundance and stocked at one of three levels of fish density. By measuring territory size over a 2-month period, while the fish were growing, I was also able to assess the effects of body size in determining the size of a territory. Defended and foraging areas were similar in absolute size, but the frequency of space use was different for defence than for foraging. As predicted, territory size decreased with increasing levels of food abundance and increased with decreasing levels of fish density. In addition, territory size increased with increasing body size even after controlling the effects of food abundance and competitor density. In comparison to previous studies, territory size of steelhead trout changed more dramatically in response to changing levels of food and competitors. For territorial animals with indeterminate growth, territory size is not only adjusted as a trade-off between the costs and benefits of defence, but also with respect to body size due to increasing metabolic demands as individuals grow. Copyright 2000 The Association for the Study of Animal Behaviour.     

Population density and intra- and interspecific competition of the African Fish Eagle Haliæeetus vocifer in Kyambura Game Reserve, southwest Uganda

Population density and intra- and interspecific competition of the African Fish Eagle Haliæeetus vocifer were studied in Kyambura Game Reserve, southwest Uganda. Density varied significantly between 1.75 and 3.25 individuals per km of shore for the breeding biotopes along the Kazinga Channel and crater lakes and between 0.45 and 0.58 for unsuitable biotopes such as the Kyambura Gorge or the bush grassland. The age-class distribution showed extremely high percentages of immatures in these unsuitable biotopes (50–95%) as a consequence of intraspecific competition for space and food. Some immatures survived in the bush grassland, feeding on lion and leopard kills and very possibly on birds. Each territory was up to five times larger than breeding pair territories along the Kazinga Channel (5.1 km² in comparison with 0.9 km²). Interspecific competition for space was likely between immature African Fish Eagles and other species of raptors. Ecological segregation with regard to space was assessed between these species in the bush grassland.     

Microhabitat use by marsh-edge fishes in a Louisiana estuary

Synopsis We used a drop sampler to characterize use of the marsh-edge ecotone by small fishes along two transects running inland from the Gulf of Mexico for ca. 25 km in Louisiana's Barataria-Caminada Bay System. Monthly sampling was stratified among upper, middle, and lower reaches and within reaches to characterize fish responses to salinity, depth, distance from shore, substrate, dissolved oxygen concentration, temperature, turbidity, velocity, and emergent stem density. In 681 quantitative samples, covering 658 m², collected between October 1987 and October 1989, we collected 57 fish species and 16 864 individuals, primarily larvae and juveniles. The 15 most abundant fishes, comprising 97.7% of all individuals, were concentrated near the marsh edge (i.e., 0 to 1.25 m distance). Some significant differences within species for seasonal variables (e.g., temperature and dissolved oxygen concentration) reflected the ephemeral duration of early life history stages. Other differences reflected ontogenetic microhabitat shifts (e.g., depth and distance from shore). Within ecological groups, characterized as demersal residents, nektonic transients, and demersal transients, spatial and temporal segregation reflected the particular habitat requirements of each species. In a principal component analysis of microhabitat use, the first three components were interpreted as seasonal, depth-and-distance, and salinity axes, respectively. The array of species and size classes in principal component space reflected the complex dimensionality of microhabitat use. The high density of fish larvae and juveniles near the marsh edge confirmed the importance of the marsh-edge ecotone as a nursery for many estuarine-dependent fishes.     

Spatial and temporal distribution of Antarctic fur seals (Arctocephalus gazella) on the breeding grounds at Bird Island,South Georgia

Summary The number of Antarctic fur seals (Arctocephalus gazella) in the area of the Scotia Sea has been increasing. Observation of their distribution on the breeding grounds are important to help design and interpret censuses. These seals are highly polygynous. In the study area at Bird Island, South Georgia, females give birth and are mated in densely packed colonies located along the shore close to the tideline. Males establish territories in this area, but are also found further inland. This study examined the density-dependent processes regulating the instantaneous size of the male breeding population; the distribution of males in relation to space and the number of females available to be mated and the effect of gregarious behaviour of females on male dispersion. Males were only territorial on the beaches in the areas where most females gave birth and subsequently had their post-partum oestrus. There was an apparent lower (19–20 m²) and upper (40 m²) limit to territory size. Males were excluded from the beach areas when the average density on the beaches was greater than 5 males per 200 m². An asymptotic density of 10–11 males per 200 m² was reached on the beaches and 4–5 males per 200 m² elsewhere. These two asymptotic densities may represent the upper and lower limits of density for a territorial system of dispersion. A model of the temporal changes in female numbers suggested that the total number of females occupying an area of beach during the mating period was approximately twice the number at the peak of the season. There was no indication that males compensated for low female density by increasing territory size. Females and pups became more dispersed after the end of the mating period. It is suggested that one function of gregariousness in females is as a mechanism for mate selection. This study also has implications for methods used to measure population size.     

Determinants of territory size in the pomacentrid reef fish,Parma victoriae

Summary Factors governing the size of territories defended by the pomacentrid reef fish, Parma victoriae, were investigated, prompted by contradictory predictions in the literature concerning the effects of food supply and competitors. Observations were carried out over the non-breeding period (March–October) on a medium density population in which territories were partially contiguous. The territory size of adult fish varied between 3 and 26 m², and was inversely correlated with local densities of conspecifics. The same range in territory size was found for both males and females, which did not differ in the time they spent on territory defence and foraging activities. No correlation existed between territory size and the abundance of algal food, body size, age or time spent on territory defence. Also, there was little variation in territory size over time, despite seasonal changes in the abundance of food algae.Experimental reduction of food supplies on isolated territories of males and females had no effect on territory size. In a higher density habitat an experiment was carried out in which population density and food abundance were simultaneously manipulated. This showed that territory size was primarily determined by intraspecific interactions, as territories exhibited considerable increases in size upon removal of neighbours. No changes in the size of defended areas resulted from either artificial increases or decreases of food levels. There were also no changes in the time spent on defence of territories, foraging time or feeding rates associated with food manipulations or territory expansion, which suggested that food was not a limited resource. This conflicted with current theories proposed to explain territory defence and expansion. It is hypothesized that intraspecific interactions constrain territory size well below the optimum in terms of the abundance of preferred food algal species.     

Components of breeding performance in two competing species: habitat heterogeneity, individual quality and density-dependence

Density‐dependent breeding performance due to habitat heterogeneity has been shown to regulate populations of territorial species, since the progressive occupation of low quality territories as breeding density increases may cause a decline in the mean per capita fecundity of a population while variation in fecundity increases. Although the preemptive use of sites may relegate low quality individuals to sites of progressively lower suitability, few studies on density dependence have tried to separate the effects of territory quality from individual quality, and none have simultaneously considered the effects of heterospecific competitors. Using two long‐term monitored populations, we assessed the relative contribution of habitat heterogeneity and bird quality (in terms of age) on the productivity of sympatric golden Aquila chrysaetos and Bonelli's eagles Hieraaetus fasciatus under different scenarios of intra‐ and inter‐specific competition. Productivity (number of offspring fledged) varied among territories and average annual productivity was negatively related to its variability in both species and populations, thus giving some support to the habitat heterogeneity hypothesis. However, the effect of habitat heterogeneity on productivity became non‐significant when parental age and local density estimators were included in multivariate analyses. Therefore, temporal changes in bird quality (age) combined with intra‐ and interspecific competition explained variability in territory productivity rather than habitat heterogeneity among territories per se. The recruitment of subadult breeders, a surrogate of mortality in eagles, strongly varied among territories. Habitat heterogeneity in productivity may thus arise not because sites differ in suitability for reproduction but because of differences in factors affecting survival. Territories associated with high mortality risks have a higher probability of being occupied by young birds, whose lower quality, interacting with the density competitors, leads to a reduction of productivity. Site‐dependent variability in adult survival and interspecific competition may be extensive, but so far largely overlooked, factors to be seriously considered for the site‐dependent population regulation framework.     

The effect of local stock density on social behavior and sex change in the Mediterranean labrid Coris julis

Synopsis Two stocks of Coris julis, a protogynous hermaphrodite labrid fish, were studied in the field in Corsica (France) to observe the effect of stock density variations on the timing of sex change. A higher density was associated with lower terminal phase male's territory size and an earlier sex change whilst the proportion of terminal phase males in the stock increased. As well, the average number of females courted per terminal phase male and the percentage of spawning with non-territorial male interferences remained similar in the two situations. These results indicate that territory availability should be considered as an important cue for wrasse sex change.     

A long-term large-scale study of the breeding biology of the Spanish imperial eagle (Aquila adalberti)

We present data from a 17-year study of the population biology of a growing population of Spanish imperial eagles Aquila adalberti across most of its breeding range. The objective of this study was to analyse the effects of age, supplemental feeding and rabbit haemorrhagic disease (RHD) on several breeding parameters of this population of eagles. Average clutch size was 2.2 eggs per clutch, and the average incubation time was 41.7 days per clutch. Fledging occurred an average of 76.8 days after hatching, the length of the fledgling period was not correlated to clutch size. The annual average percentage of pairs laying eggs was 88%. A significant reduction in the percentage of pairs laying eggs in the period 1992–1994 (from 91 to 81%) coincided with most of the eagles’ territories being affected by the rabbit epizootic disease, RHD, which reduced their food supply significantly. Average productivity was 1.23 chicks per monitored territory, average breeding success was 1.40 chicks in a clutch per territory and the average fledging rate was 1.69 chicks per territory with hatching success. The main known causes of breeding failure during incubation were nest collapse and human disturbance. During chick-rearing, total or partial chick losses were mainly caused by siblicide, disease, malnutrition or nest collapse. In 26.2% of the 1372 monitored breeding attempts, at least one of the breeding birds was a subadult (the male in 56.1% of the cases, the female in 15.5% and both sexes in 28.4% of cases). In cases of mixed-aged pairs (n = 205), 70.7% were the result of a substitution, and 29.3% were the result of the forming of a new pair. Egg laying took place significantly earlier and breeding success was higher in territories occupied by adults than in those occupied by subadults. Breeding parameters were higher in high-quality (rabbit-rich) territories than in low-quality (rabbit-poor) territories, but only for those territories occupied by adults. The values obtained in the territories occupied by adults were only significantly higher than in those of the subadults in high-quality territories. Age and territory quality thus simultaneously affected reproductive output.     

The feeding habits of six pelagic and predatory teleosts in eastern Cape coastal waters (South Africa)

A four-year study of the feeding habits of six pelagic teleost species in eastern Cape coastal waters of South Africa is analysed. These predators are Lichia amia, Pomatomus sallatrix, Seriola lalandi, Atractoscion aequidens, Katsuwonus pelamis and Thunnus albacares. All species occur in inshore coastal water (< 1 -8 km from shore) for at least part of the year. The tunas were also sampled from the edge of the continental shelf (40–50 km from shore). The use of squid 'beaks' and fish otoliths allowed accurate prey identification and size estimates. Variations in prey taken are attributed to differences in locality, predator species and size. Three pelagic fish species are important to all the predators: Sardinops ocettata, Etrumeus teres and Engraulis capensis. The inshore squid, Loligo reynaudi, is another major prey item. Offshore tuna samples revealed a different prey spectrum, viz. oceanic squid and saury, Scomberesox saurus, being taken frequently.     

The activity budget and use of territory by a tropical blenniid fish

Both sexes of Ophioblennius atlanticus (Valenciennes) maintain permanent territories, intermixed without pattern. They occur at a density of about 1.9 individuals per m² on shallow coral rock and comprise a significant portion of the benthic fish biomass. They are diurnal; about 60% of their time is spent resting, 15% swimming and 8.5% feeding. Feeding is concentrated in the afternoon and is time-minimized. Territory is used exponentially, about 50% of time being spent in about 15% of the territory. The relatively infrequent use of peripheral parts of a territory suggest that it is potentially compressible or expansible. This, in turn, leads to a conclusion that an optimum size can only be defined as a range and mat, normally, a territory includes more resources than the minimum for survival. Competition may be reduced between aegis corrivals of different species.     

Alongshore dispersal and site fidelity of juvenile plaice from tagging and transplants

Juvenile plaice Pleuronectes platessa ( n  = 1281) were tagged and released at two locations 300 m apart on a 1 km long sandy beach. Most (>90%) of the fish were recaptured within 100 m of the release site (shown by the colour of the tag), with very few caught >200 m distance after 6 weeks. The changing spatial distribution of marked fish was adequately reproduced by a simple dispersal model with a single parameter: a 78% probability of remaining in a 100 m wide zone from one day to the next, with a 22% chance that fish move into an adjacent zone. In a subsequent similar study at the same beach, fish were either released at the point of capture ( n  = 881) or transplanted to the alternate site ( n  = 910) 100 m distant. After 6 weeks, transplanted fish moved along the shore towards their sites of original capture. Fish replaced at the point of capture showed no such movement along the shore. Further modification of the dispersal model to allow for a distinction between dispersal from home sites and from sites away from the original point of collection, was sufficient to reproduce the behaviour of the populations of both transplanted and control treatment groups. The likelihood of dispersal from home sites was much less than that seen at sites away from home. Juvenile plaice thus have a degree of long‐shore site fidelity not expected of a fish with strong depth‐related migration behaviour in a relatively homogenous habitat.     

Distribution, movements, and mortality of anadromous arctic char, Salvelinus alpinus L., in the Cumberland Sound area of Baffin Island

During 1972 the downstream movement of anadromous artic char, Salvelinus alpinus L., from two rivers flowing into Cumberland Sound, Baffin Island, began during the middle of May and was completed within 2 weeks. This movement took place during both the day and night. Upon reaching saltwater, many char older than 9 years (longer than 20 cm) began to migrate along the shore of the Sound at a rate of 0–6–0-9 km/day with the result that the average maximum distance travelled from the natal river was 40–50 km. Char 6–9 years old, 10–20 cm in length, remained much closer to the rivers while those younger than 5 years never entered saltwater. During the period of saltwater residence, the fish frequently moved into the intertidal zone and freshwater. Maturing fish of both sexes were occasionally found in saltwater. The upstream migration to the lakes began during the second week of August and was completed within 5–6 weeks. Females tended to ascend the rivers before males, although fish of all different size and age migrated upstream simultaneously. Migration occurred during both the day and night. The maximum distance anadromous char moved from saltwater was 40 km. The char were found only in freshwater between May and September and were normally located near the bottom of lakes at a depth of 15–40 m. During this period, large scale movements were probably very limited. Average annual mortality was estimated at 16.0% per year with the highest rates (25–30%) occurring at ages 10 and 15–17 years. Many fish probably died of old age but physical deterioration during and after spawning was also an important cause of death.     

Influence of Landscape Geomorphology on Large Wood Jams and Salmonids in an Old-growth River of Upper Michigan

We investigated the structure of large wood jams (LWJ) and their use by brook trout (Salvelinus fontinalis, Mitchill) and other fish in four geomorphically-distinct sections of the Little Carp River, a small river flowing through an uncut, old-growth, northern hardwood-conifer forest along the south shore of Lake Superior, Upper Michigan. We characterized nine LWJ per section and then electroshocked fish at three randomly selected LWJ per section. Structural characteristics of LWJ (e.g., total volume of wood, number of logs) varied with geomorphology at the scale of approximately one km. Differences in the abundance of fish associated with LWJ were not statistically significant among LWJ and non-LWJ portions of stream across all study reaches. Factors that explained most variability in the proportion of salmonids at LWJ (valley constraint, volume and number of pieces in the jam) reflected both large-scale geomorphology and characteristics of LWJ. If emulating an old-growth system is the goal for restoring habitat, attention should be given to the correlation of LWJ with larger-scale geomorphology of the reference river. However, it cannot be assumed that LWJ restoration will necessarily increase brook trout abundance near LWJ in a system similar to the Little Carp River as we observed low overall correlation between brook trout abundance and LWJ.     

A RADAR STUDY OF TERN MOVEMENTS ALONG THE COAST OF GHANA

During spring and autumn 1970 and 1971 the movements of terns off the coast of Ghana near Accra were studied by radar. In autumn, dense movements were recorded within 22 km of the coast both day and night from July to early December. Fluctuations in the magnitude of the movements suggest that the terns were on passage through the Gulf of Guinea. Simultaneous to these movements were roosting flights into shore roosts and both movements appeared independent.
In contrast, no spring movements along the coastline were detected either by radar or visually from the land. Although small tern flocks probably moved westwards within 22 km of the coast, but outside their detectable radar range, the results suggest that the majority of terns return through the Gulf of Guinea by a different route from that taken in autumn.
The difference between spring and autumn movements correlates well with the seasonal variation of the terns' food sources. In autumn, upwellings result in an abundant fish supply (sardines) in the surface waters close to land, whereas in spring, fish is abundant only well off shore such as at fronts (convergences) and upwellings (divergences) both of which have been located as far as 600 km from the coast.     

Small fish communities in two regions of the Liangzi Lake, China, with or without submersed macrophytes

Small fish communities were compared between the vegetated and vegetation-free regions of the Liangzi Lake, a shallow lake along the middle reach of the Yangtze River, China. Fish were sampled using 10 × 10 m² block nets and poisoning. Three samples were taken from either the near shore area or lake centre of each region. A total of 19 fish species were collected; all species occurred in the vegetated region but only 12 occurred in the vegetation-free region. The dominant small fish were Carassius auratus auratus in the vegetated region and Ctenogobius giurinus in the vegetation-free region. Diversity, density and biomass of small fishes were significantly higher in the vegetated region than in the vegetation-free region in both near shore and lake centre areas. In the vegetated region, density and biomass of small fishes was significantly higher, while species diversity significantly lower in the near shore area than in the lake centre. In the vegetation-free region, density of small fishes was significantly higher in the near shore area than in the lake centre area, but species diversity or biomass was unaffected by location.     

Recovery of the Chesapeake Bay Bald Eagle Nesting Population

Abstract We conducted annual aerial surveys throughout the tidal reach of the Chesapeake Bay, USA, between 1977 and 2001 to estimate population size and reproductive performance for bald eagles (Haliaeetus leucocephalus). The population increased exponentially from 73 to 601 pairs with an average doubling time of 8.2 years. Annual population increase was highly variable and exhibited no indication of any systematic decline. A total of 7,590 chicks were produced from 5,685 breeding attempts during this period. The population has exhibited tremendous forward momentum such that >50% of young produced over the 25-year period were produced in the last 6 years. Rapid population growth may reflect the combined benefits of eliminating persistent biocides and active territory management. Reproductive rate along with associated success rate and average brood size increased throughout the study period. Average reproductive rate (chicks/breeding attempt) increased from 0.82 during the first 5 years of the survey to 1.50 during the last 5 years. Average success rate increased from 54.4% to >80.0% during the same time periods. The overall population will likely reach saturation within the next decade. The availability of undeveloped waterfront property has become the dominant limiting factor for bald eagles in the Chesapeake Bay. Maintaining the eagle population in the face of a rapidly expanding human population will continue to be the greatest challenge faced by wildlife biologists.