Density-dependent foraging behaviour of sheep on alpine pastures: effects of scale

Foraging patterns of large herbivores may give important clues as to why their life history varies depending on population density. In this landscape-scale experiment, domestic sheep Ovis aries were kept at high (80 sheep km⁻²) and low (25 sheep km⁻²) population densities during summer in high mountain pastures in Hol, Norway. We predicted an increasing use of less preferred plant species or habitat types with increasing sheep population density. Foraging behaviour was investigated by direct observations of individually marked sheep on different spatial scales, and diet composition was also assessed with microhistological analysis of faecal samples from known individuals. We found that the effects of density on foraging behaviour depended on scale and were only detected at the scale of diet choice. Use of the common grass species Deschampsia flexuosa , which provided the bulk forage (10–65% of the diet), remained constant throughout the season at low densities, but increased significantly over time at high densities. On a coarser spatial scale, neither within vegetation type nor between vegetation types, selection was affected by density, but vegetation type selection differed depending on whether the sheep were grazing or resting. Our study provides evidence of density dependence in foraging behaviour, but only at the finest spatial scale (diet choice).     

Population studies on feral American mink Mustela vison in Ireland

Populations of feral American mink Mustela vison were investigated in several study areas in Ireland, principally by capture-recapture techniques. The populations were found to be essentially self-regulating, with stress, weight loss and mortality evident during the winter and spring. The highest mean density recorded, in terms of animals per unit length of river or lake shore, was 1.4 individuals km⁻¹ with a peak density of 2.3 individuals km⁻¹, but population densities were considerably lower in most areas. Trappability and population structure, in age, sex and residency classes, varied between habitats, but population turnover was high, leading to complete replacement of individuals in two populations monitored for two years. The availability of the preferred prey, crayfish Austropotamobius pallipes appeared to be a major factor determining population size and preferred breeding areas. Control methods are either unnecessary or are likely to fail in most situations, and may be counterproductive.     

Energy density of marine pelagic fish eggs

Analysis of the literature on pelagic fish eggs enabled generalizations to be made of their energy densities, because the property of being buoyant in sea water appears to constrain the proximate composition of the eggs and thus to minimize interspecific variation. An energy density of 1.34 J μl⁻¹ of total egg volume is derived for most species spawning eggs without visible oil globules. The energy density of eggs with oil globules is predicted by     x (J μ1¹) where x is the fractional volume of the oil globule.     

Influence of host plant vs. natural enemies on the spatial distribution of a pine sawfly, Neodiprion autumnalis

Abstract .1. The pine sawfly, Neodiprion autumnalis , infests ponderosa pine, Pinus ponderosa , growing at low densities near the bottom of an altitudinal gradient in Arizona, U.S.A. The relative importance of host-plant quality vs. natural-enemy effects in determining the spatial distribution of this sawfly was examined over a 3-year period.
2. Field and laboratory bioassays were conducted on all life stages of N. autumnalis at two forest stand densities (high ≥ 23 m² ha^–1, low ≤ 7 m² ha^–1) and at two elevations (bottom slope = 2390 m, top slope = 2540 m). These experiments were used for constructing life tables of N. autumnalis that compared the effects of host-plant quality on oviposition preference and progeny performance with the effects of natural enemies at different tree densities and elevations.
3. Life-table analyses determined that mortality attributed to host-plant effects during the egg and larval stages had the largest impact on fitness between tree densities and elevations.
4. Natural enemies caused a significant reduction in progeny survival, but their effects were similar across all tree densities and elevations during egg and larval life stages. However, cocoon-stage survival did vary between tree densities and elevations due to natural-enemy effects.
5. It was concluded that the observed oviposition preference for, and higher progeny performance on, trees at low densities and bottom slope elevations were caused primarily by host-plant effects.
6. These results further the argument that heterogeneity at the resource level (i.e. bottom-up forces) determines potential outcomes of multitrophic level interactions.     

Using passive sonic telemetry methods to evaluate dispersal and subsequent movements of hatchery-reared white sturgeon in the Kootenay River

A total of 35, age 1 juvenile Kootenay River white sturgeon ( Acipenser transmontanus ), were fitted with sonic tags in 2005 and released as part of larger hatchery release groups at five sites to evaluate dispersal and subsequent movements (seven tags per site). Juvenile sturgeon released at three locations within the deep, low gradient reach (typical gradient of 0.02 m km⁻¹ and velocities of <0.4 m s⁻¹) of the Kootenay River below Bonners Ferry, ID showed substantial dispersal both up and downstream; however, downstream redistribution was more common. White sturgeon from all three release locations overlapped during dispersal, with 9% of tagged fish moving from river release sites into Kootenay Lake. The three hatchery release locations in this low gradient reach produced good dispersal of hatchery progeny into available habitats. Tagged fish released above Bonners Ferry in the shallow, higher gradient reach (typical gradient of 0.6 m km⁻¹, and velocities >0.8 m s⁻¹) at two additional sites all moved downstream of the gradient break at Bonners Ferry, ID into the lower gradient reach within 2 months of release. In total, 93% of these tagged fish relocated to the low gradient section within 25 days of release, with some fish undertaking this movement within 1 day. In general, age 1 hatchery release juveniles were mobile and capable of substantial movements.     

Home-range use, activity, and density of caracal in relation to prey density

Two male and three female caracal Felis caracal were radio-tracked over a 1-year period in arid shrub on the west coast of South Africa, by day and night over at least 130 days for each caracal, and uninterrupted for up to 120 h at a time. These results, on short-term use of space, were related to concurrent availability of prey. The use by caracal of specific plant communities showed a significant positive correlation to prey biomass of rodents. Males had much larger home-ranges (26.9±0.75 km²) than females (7.39±1.68 km²). Male home-ranges overlapped completely with those of females, whereas female ranges overlapped between 0 and 19%. Caracal were active by night and day; onset of activity was affected more by ambient temperature (T_A) than photoperiod. Caracal were active significantly longer on nights colder than 20°C. Females ceased activity at T_A > 20°C, males at T_A > 22°C. Males foraged faster than females (667 vs. 312 m h⁻¹) and moved more than twice the distance of females during an active period. Calculated density of caracal was between 0.23 and 0.47 km⁻².     

Elephants caught in the middle: impacts of war, fences and people on elephant distribution and abundance in the Caprivi Strip, Namibia

We conducted wet [26 March–4 April 2003 (Apr03)] and dry [1–8 November 2005 (Nov05)] season aerial surveys of African elephants ( Loxodonta africana Blumenbach) in the Caprivi Strip, Namibia to provide an updated status report on elephant numbers and distribution and assist with a historical analysis of elephant distribution and abundance in the Caprivi Strip. During the wet season when water was available in seasonal pans, elephants were widely distributed throughout the survey area. In contrast, during the dry season, a majority of elephant herds occurred within 30 km of the perennial Kwando, Linyanti and Okavango rivers and few herds occurred within the West Caprivi Game Reserve where water in the seasonal pans was limited. We estimated 5318 elephants for the 7731-km² survey area (0.71 elephants km⁻²) for the Apr03 wet season survey and 6474 elephants for the 8597-km² survey area (0.75 elephants km⁻²) for the Nov05 dry season survey. Based on our aerial surveys and reports of elephant numbers and distribution from historical aerial surveys and telemetry studies, civil war, veterinary fences and human activities appear to have effected changes in African elephant abundance, distribution and movements in the Caprivi Strip, Namibia since 1988 when the first comprehensive aerial surveys were conducted.     

Spatio-temporal variation in macroinvertebrate assemblages of glacial streams in the Swiss Alps

1. Changes in water chemistry, benthic organic matter (BOM), and macroinvertebrates were examined in four different glacial streams over an annual cycle. The streams experienced strong seasonal changes in water chemistry that reflected temporal changes in the influence from the source glacier, especially in water turbidity, particulate phosphorus and conductivity.
2. Nitrogen concentrations were high (nitrate-N values were 130–274 μg L^–1), especially during spring snowmelt runoff. Benthic organic matter attained >600 g m^–2 dry mass at certain times, peaks being associated with seasonal blooms of the alga Hydrurus foetidus .
3. Macroinvertebrate taxon richness was two to three times higher (also numbers and biomass) in winter than summer suggesting winter may be a more favourable period for these animals. Benthic densities averaged 1140–3820 ind. m^–2, although peaking as high as 9000 ind. m^–2. Average annual biomass ranged from 102 to 721 mg m^–2, and reached >2000 mg m^–2 at one site in autumn.
4. Taxa common to all sites included the dipterans Diamesa spp. and Rhypholophus sp., the plecopterans Leuctra spp. and Rhabdiopteryx alpina , and the ephemeropterans Baetis alpinus and Rhithrogena spp. Principal components analysis clearly separated winter assemblages from those found in summer.     

The relative densities of red (Cervus elaphus) and roe (Capreolus capreolus) deer and their relationship in Scottish plantation forests

Population densities of red and roe deer were estimated in 20 conifer plantations throughout central and northern Scotland. Ten forests were selected on the basis of culling records that were considered likely to cover the range of possible densities. Ten further forests were selected at random to investigate the relationship between red and roe deer densities. Density assessment was by standing-crop dung-group counts, on 1.5 x 100 m transects. Dung decay rates were estimated in four areas and used to convert the values to deer densities. Red deer densities were estimated to range from 0.3-35 km⁻²; roe deer densities ranged from 0.5-25 km⁻². Preferences for different structural habitats within individual forests (i.e. blocks with different age classes of tree) were inferred from dung densities. Red deer preferred unplanted ground, open-thicket, and prethicket; roe deer likewise preferred prethicket (highly) and open-thicket. Both species avoided closed-canopy habitats. Red and roe deer densities were negatively correlated and the relationship was significant when the data were log transformed ( P < 0.05).     

Influence of egg predation and physical disturbance on lake trout Salvelinus namaycush egg mortality and implications for life-history theory

The influence of egg predators and physical disturbance on lake trout Salvelinus namaycush egg mortality was investigated in situ in Lake Michigan where recruitment is below detectable levels and egg predator abundance is high. Comparisons were made with Lake Champlain where recruitment is low and egg predator abundance is also low and with Parry Sound (Lake Huron) where recruitment is moderate and egg predators are in low abundance. A multi-density egg seeding method (100 to 5000 eggs m⁻²) was used to quantify the effect of physical disturbance and egg predation on egg loss. Wind fetch was used as an index of physical disturbance and comparisons across all locations and egg densities suggested that at sites with high wind fetch (>5 km), physical disturbance may be a greater source of egg loss than predation. When analyses were limited to those sites having a wind fetch of <5 km, the percentage of eggs recovered was found to be linearly related to predator density. The strength of this relationship was based largely on egg recovery at 500 and 1000 eggs m⁻² because recovery at lower (100, 250 eggs m⁻²) and very high (5000 eggs m⁻²) densities was not significantly related to predator density. The reason for this is probably that at low egg densities, crayfish Orconectes spp., the major egg predator at most sites, had difficulty finding and consuming eggs and at high egg densities they became satiated. Egg loss was directly related to wind fetch for Lake Michigan and on average six-fold greater than for Parry Sound suggesting that without corresponding changes in fecundity and age structure, lake trout populations in large lakes like Lake Michigan are inherently less productive than those from enclosed inland waters.     

Distribution and population densities of seven species of bat in northern England

Distribution and minimum population densities for seven UK bat species known to be resident in northern England were calculated in an area covering 2500 km². The species present were pipistrelle ( Pipistrellus pipistrellus ), brown long-eared ( Plecotus auritus ), Daubenton's ( Myotis daubentonii ), whiskered ( Myotis mystacinus ), Natterer's (Myotis nattereri) , noctule ( Nyctalus noctula ) and Brandt's (Myotis brandtii). Data were collected primarily from counts at summer roosts over the period 1983 to 1990. A total of 310 bat roosts were discovered within the study area. Of the 256 roosts at which the species present was identified, the majority, 127 (49.6%) were P. pipistrellus , with a mean maternity roost size of 69.6 bats. A minimum population density of 12.6 batskm⁻² was estimated for P. pipistrellus , based on summer (maternity) roosts. The minimum population density estimate was higher than previous studies in northern England but substantially lower than those reported in Scotland (18.2 bats km-⁻²). The combined density of M. mystacinus, M. brandtii, and P. pipistrellus , which have similar foraging styles (15.8 bats km⁻²), is comparable to Scottish P. pipistrellus densities. The density of M. duubentonii was also lower than in Scotland, although the density of P. auritus was comparable. The majority of summer roosts for all species were found in buildings, except N. noctula and M. duubentonii which used bridges/tunnels or trees.     

The role of life history in the relationship between population dynamics and environmental variability in two Mediterranean stream fishes

Chub Squalius torgalensis and nase Chondrostoma lusitanicum , in a Mediterranean stream, showed important differences in life-history traits and population dynamics. Both species reached mean maturity at age 2 years. Chub lived up to age 5 years, spawned in March to June, grew at a maximum rate of 0·59 mm mm⁻¹ year⁻¹ and showed a low reproductive allocation, with fecundity and egg size increasing with body size. Nase lived up to age 4 years, spawned in January to April, grew at a maximum rate of 0·46 mm mm⁻¹ year⁻¹ and showed a high reproductive allocation, with egg size independent of body size. Both chub and nase showed moderate fluctuations in population size during 1991–1998, but differed in factors driving density at age. Density of age 1 year juvenile chub decreased following severe summer droughts and proportionate survival prevailed thereafter. Density of age 2 year adult nase decreased following severe spring floods, but neither environmental nor parental stock effects were detected for juveniles and older fishes. The results illustrated the interplay between life history and environmental variability in driving fish population dynamics, with impacts of both summer droughts and spring floods being contingent on species-specific patterns of spawning and reproductive investment.     

Effects of supplementary feeding on territoriality, breeding success and survival of pheasants

1. The effects of supplementary food in spring on subsequent pheasant breeding in an intensively farmed area in southern England were assessed by a large-scale, replicated field experiment.
2. Territorial cock pheasants were counted in April, and the breeding success and survival of radio-tagged hens were monitored in six 1-km² plots during 1994 and 1995. Total numbers of young reared and post-breeding pheasant densities were found by August counts. In 1994, three randomly selected plots were supplied with wheat grains via hoppers along woodland edges and hedgerows. The other three plots acted as controls and the treatments were reversed in 1995.
3. The density of cock territories increased significantly in food-supplemented plots (44 ± 8 km−2) in relation to control plots (29 ± 8 km−2), and the presence of hoppers significantly affected the locations of territories. However, similar proportions of territorial males acquired harems in the control and food-supplemented plots. Hen density did not increase and, consequently, the mean harem size was significantly lower with supplementary feeding.
4. Hens given supplementary food did not nest earlier and the number of nests initiated, clutch sizes and the proportion of successful nests did not differ significantly from those of controls. However, hens supplied with supplementary food re-nested significantly more quickly following the loss of a nest or brood.
5. Radio-tagged hens did not rear significantly more young with supplementary feeding. Hen survival was unchanged and post-breeding pheasant densities were no higher.
6. On present evidence, spring feeding cannot be advocated as a management technique to improve the breeding success of pheasants surviving the winter.     

Beluga whale ecology: a concern of freshwater biologists?

SUMMARY. 1. Churchill estuary beluga whales displayed their highest densities in freshwater habitat, suggesting preferential use.
2. Whale densities in freshwater were as high as 350 animals km⁻² and represented over one tenth of the estimated Hudson Bay population.
3. Future environmental assessments of hydroelectric development in northern Manitoba, Canada, must consider the effects on the beluga whales.     

Insect herbivory in an experimental agroecosystem: the relative importance of habitat area, fragmentation, and the matrix

Habitat area, fragmentation, and the surrounding matrix influence levels of herbivory in various ecosystems, but the relative importance of these effects has rarely been assessed. We compared levels of herbivory and densities of dominant arthropod herbivores (the hemipteran insects Agallia constricta, Empoasca fabae, Therioaphis trifolii, Lygus lineolaris and Halticus bractatus ) among experimental plots that varied in the area and fragmentation of clover habitat and the composition of the matrix (bare ground or grass) surrounding clover habitat. To assess levels of herbivory, we compared clover biomass within herbivore exclosures to the biomass accessible to herbivores. The area and fragmentation of clover habitat, as well as matrix composition, significantly influenced the collective densities of herbivores, although each species exhibited unique responses to habitat structure. Herbivory was strongest in plots with large (64  m²) as compared to small (16  m²) amounts of clover habitat. The difference in clover biomass between the inside and outside of exclosures increased significantly with increasing density of Empoasca fabae but was unrelated to the densities of the other herbivores, suggesting that Empoasca fabae was an exceptionally important herbivore in this system. This study supports the view that herbivore densities and herbivory generally increase with increasing area of plant monocultures, but emphasizes that levels of herbivory may be driven primarily by one or a few key herbivore species.     

Nocturnal autumn bird migration at Falsterbo, South Sweden

We investigated the patterns of nocturnal bird migration in autumn 1998 at a coastal site on the Falsterbo peninsula in south-western Sweden, by means of a passive infrared device. In total 17 411 flight paths, including track direction and altitude, of migrating birds were recorded for 68 nights from August to October. Mean migratory traffic rate per night varied between 6 and 6618 birds km⁻¹ h⁻¹, with an average of 1319 birds km⁻¹ h⁻¹. Migration at Falsterbo showed a similar seasonal pattern to that reported for central Europe, with pronounced peaks of migration and intermittent periods with relatively low migratory intensities. Weather factors explained two thirds of the variance in the intensity of bird migration. During nights with intense migration, associated with weak winds, the mean track direction was close to that in central western Europe (225°). Birds usually maintained a constant heading independent of wind directions and, in consequence, were drifted by the wind. The mean orientation clearly differed from that of the nearest coastline, suggesting that the birds did not use the topography below to compensate for wind drift.     

Effects of experimental small-scale grassland fragmentation on spatial distribution, density, and persistence of ant nests

Abstract.  1. Grassland fragmentation is expected to influence the abundance of different invertebrate species to a different extent. Fragmentation-related effects are of particular importance in species that interact with many other species.
2. The density and spatial distribution of nests of 15 ant species in experimentally fragmented calcareous grasslands at three sites in the Northern Swiss Jura mountains were examined. Fragments of different size (0.25 m², 2.25 m², and 20.25 m²) were isolated by a 5-m wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland.
3. Three years after initiation of the experiment, ant nest density did not differ between fragments and control plots. Six years after initiation of the experiment, however, ant nest density and forager abundance were higher in large fragments than in large control plots. Ant nests tended to occur more frequently along the edge of fragments than in the core area. Persistence time of nests of the most abundant species, Lasius paralienus , tended to be shorter in fragments than in control plots. Furthermore, persistence time was longer in nests situated close to the fragment edge than in nests in the core area.
4. Effects on nest density, edge effects on the spatial distribution of nests, and the relationships between nest density and environmental factors were more pronounced when only nests of L. paralienus were considered. The implications of these findings for plant and other invertebrate species are discussed.     

Population biology and stock assessment of Pristipomoides filamentosus on the Mahe Plateau, Seychelles

The population of P. filamentosus around the periphery of the Mahe Plateau was studied using data derived from commercial handline fishing operations between November 1989 and December 1990. Sex ratio was close to one. Fork length was in the range 25.6–79.8 cm: at first sexual maturity, 36–38 cm for females and 40–42 cm for males. Spawning occurred from October to April peaking between February and April and in November.
The length ( f.l. , cm)–weight (kg) relationship was W =0.00005353 × f.l. ^2.7004. The von Bertalanffy growth parameters, estimated from length frequency data, were K =0.2875 and L _inf=81.7 cm. Mortality rates, estimated from length converted catch curve analysis, were Z =0.811, F =0.277 with M =0.534 derived from Pauly's empirical formula. Jones' length cohort analysis gave an estimate of F= 0.294 for the fully exploited part of the stock, and a density of 2.59 t km⁻² for the periphery of the plateau from 75 to 150 m depth. The Lesley constant catchability model, applied to the results of intensive fishing around isolated banks, derived a mean density estimate of 2.99t km⁻². The sustainable yield was estimated to be 233–268 tonnes per annum and the catch in 1990 was 200 tonnes.     

Endurance swimming of European eel

A long‐term swim trial was performed with five female silver eels Anguilla anguilla of 0·8–1·0 kg ( c . 80 cm total length, L _T) swimming at 0·5 body lengths (BL) s⁻¹, corresponding to the mean swimming speed during spawning migration. The design of the Blazka‐type swim tunnel was significantly improved, and for the first time the flow pattern of a swim tunnel for fish was evaluated with the Laser‐Doppler method. The velocity profile over three different cross‐sections was determined. It was observed that 80% of the water velocity drop‐off occurred over a boundary layer of 20 mm. Therefore, swim velocity errors were negligible as the eels always swam outside this layer. The fish were able to swim continuously day and night during a period of 3 months in the swim tunnel through which fresh water at 19° C was passed. The oxygen consumption rates remained stable at 36·9 ± 2·9 mg O₂ kg⁻¹ h⁻¹ over the 3 months swimming period for all tested eels. The mean cost of transportation was 28·2 mg O₂ kg⁻¹ km⁻¹. From the total energy consumption the calculated decline in fat content was 30%. When extrapolating to 6000 km this would have been 60%, leaving only 40% of the total energy reserves for reproduction after arriving at the spawning site. Therefore low cost of transport combined with high fat content are crucial for the capacity of the eel to cross the Atlantic Ocean and reproduce.     

Modeling density dependence and climatic disturbances in caribou: a case study from the Bathurst Island complex, Canadian High Arctic

Peary caribou Rangifer tarandus pearyi is the northernmost subspecies of Rangifer in North America and endemic to the Canadian High Arctic. Because of severe population declines following years of unfavorable winter weather with ice coating on the ground or thicker snow cover, it is believed that density-independent disturbance events are the primary driver for Peary caribou population dynamics. However, it is unclear to what extent density dependence may affect population dynamics of this species. Here, we test for different levels of density dependence in a stochastic, single-stage population model, based on available empirical information for the Bathurst Island complex (BIC) population in the Canadian High Arctic. We compare predicted densities with observed densities during 1961–2001 under various assumptions of the strength of density dependence. On the basis of our model, we found that scenarios with no or very low density dependence led to population densities far above observed densities. For average observed disturbance regimes, a carrying capacity of 0.1 caribou km⁻² generated an average caribou density similar to that estimated for the BIC population over the past four decades. With our model we also tested the potential effects of climate change-related increases in the probability and severity of disturbance years, that is unusually poor winter conditions. On the basis of our simulation results, we found that, in particular, potential increases in disturbance severity (as opposed to disturbance frequency) may pose a considerable threat to the persistence of this species.