Reliable and abundant resources are likely to favour specialization, while unpredictable environmental variation should favour a generalist strategy. The rodent population cycles of northern latitudes can be seen as both predictable and unpredictable, depending on the scale in time and space. The arctic fox Alopex lagopus is an opportunistic carnivore, but paradoxically, it seems to function as a specialist on fluctuating rodent Arvicolinae populations in most inland areas. We have studied the dietary response of arctic foxes in Sweden during 5 years of varying abundance of Norwegian lemming Lemmus lemmus, and how these changes influenced the reproductive success of the foxes. The arctic fox population on mainland Fennoscandia is threatened by extinction and the situation has deteriorated during the 1980s and 1990s because of an absence of lemming peaks. Our results showed that in all years, lemming was the main prey for arctic foxes, with 85% frequency of occurrence in summer faeces (scats). Bird remains (mainly Passeriformes) were present in 34% of the scats, reindeer Rangifer tarandus in 21%, voles and shrews in 4% and hares Lepus timidus in 2% of the scats. The occurrences of lemming, bird and larger mammal (reindeer and hare) remains in the scats varied significantly between years. Temporal variations within summer seasons and dietary differences between sub-areas, indicated that arctic foxes fed opportunistically on the alternative prey types. Den occupancy rates were positively correlated with lemming population densities during the previous winter, indicating a strong numerical response. We conclude that from a functional aspect, the arctic fox in Sweden is a lemming specialist, since lemming is the main prey and their abundance is the best predictor of arctic fox reproductive success. Other prey are used opportunistically in relation to their availability.

This study examines aspects of the nesting behaviour of Brants' whistling rat Parotomys brantsii, a medium-sized rodent indigenous to southern Africa. In the semi-arid regions in which it occurs P. brantsii breeds opportunistically, modifying its breeding period in relation to rainfall. Within a winter rainfall area, females produced up to four litters of three to four young during the winter–spring period following a gestation period of c. 38 days, with females showing a post-partum oestrus. The behaviour of females was modified by the birth of their pups, after which they spent extended periods underground, interspersed with bouts of food-collecting activity. With the emergence of pups above-ground, females increased their vigilant activity. At 5 weeks pups had developed all the important behavioural characteristics of adults, such as creating their own overnight food stores, collecting nesting material and defending a warren area against conspecifics. This was also the age at which the young dispersed from their natal nest area as well as the time at which females could reach sexual maturity, despite physical immaturity. The high reproductive output of female P. brantsii, as well as the rapid ontogeny of their young, compared to other otomyine rodent species, may represent an adaptation for maximizing reproductive potential in the semi-arid areas they inhabit.

Early mortality and pup growth during the perinatal period were investigated in pups of the subantarctic fur seal Arctocephalus tropicalis on Amsterdam Island. Mothers that were shorter in body length and expected to be younger, gave birth earlier in the pupping season, compared with longer/older mothers. Pups born early were often still-born, suggesting that shorter/younger mothers that gave birth early in the season were not able to carry their foetuses to term. Pregnant females arrived ashore 1.4 days before giving birth, regardless of the date and their body condition. There was a positive relationship between maternal body length and pup birth mass. Consistently, birth mass increased throughout the pupping period. After parturition, mothers suckled their pups during an average 8.7-day postnatal period that was significantly shorter in mothers giving birth late in the season. However, the absolute gain in pup mass was 1.5 kg regardless of the birth date, suggesting that mothers did not leave their pups before they had transferred a given amount of body reserves to them. We propose that pups born late grew faster because they were bigger at birth and because their mothers were likely to be more experienced. Mothers in good condition, nursing male pups transferred more milk and therefore greater mass to their pup, whereas mothers in poor condition were unable to do so. Such differences did not occur in smaller female pups, suggesting that pup growth was limited by maternal resources in male pups but not in female pups. The lack of a relationship between birth mass and absolute gain during the perinatal period suggests that mass at birth determined pup body mass after the perinatal period. Body mass is an important factor in growth rate and survival during the period of pup dependence.

The social structure of an Old World tent-making bat Cynopterus sphinx (Megachiroptera), was investigated in western India. A combination of census and mark–recapture data over 2 years (1996–98) was used to infer the form of the mating system, compositional stability of social groups and mode of new social group formation. The breeding population of C. sphinx was subdivided into diurnal roosting colonies, each of which contained one to five discrete roosting groups and often one or more solitary bats in adjacent roosts. Bats most frequently roosted in stem tents constructed in the flower/fruit clusters of the kitul palm Caryota urens. Temporal variation in social structure was assessed using visual census data for a subset of the study population over 3 years (1995–98) spanning six consecutive reproductive periods. The sex and age composition of diurnal roosting groups indicated a polygynous harem-forming mode of social organization, as groups invariably contained a single adult male, 1–37 reproductive females and their dependent young (n = 33 harems). Harem size averaged 6.1 adults in the wet season (n = 19, SD = 3.5) and 13.6 adults in the dry season (n = 14, SD> = 8.5). The same harem social configuration was maintained year-round, despite a high degree of synchrony and seasonality in the timing of reproduction. Juveniles of both sexes dispersed after weaning and sexually immature bats were never present in harems at the time of parturition. Adult females often remained associated as roostmates from one parturition period to the next, and group cohesion was unaffected by turnover of harem males. Adult females frequently transferred among roosts within the same colony, and harems underwent periodic fissions and fusions. The founding of new harems most often resulted from the fissioning of previously cohesive harems within the same colony. However, some harems contained disproportionate numbers of yearling females, indicating that new groups are also founded by nulliparous females of the same age cohort. A significant degree of heterogeneity in age composition among harems was revealed in the 1998 dry season, but was unrelated to age-stratification of tent roosts. Although formation of new harems may be non-random with respect to age composition of the founders, founding events are not restricted to newly created tents and often involve recolonization of previously occupied roosts.

The diet of southern African carnivorous slugs of the endemic family Chlamydephoridae is discussed. At least three species, Chlamydephorus bruggeni, C. burnupi and C. sexangulus, are predators of pill-millipedes Sphaerotherium sp. The feeding behaviour of C. sexangulus is described in detail and illustrated. It is suggested that the prey, once captured, is killed or immobilized by the introduction of a toxin through the intersegmental membrane joining the millipede's skeletal plates. The source for such a toxin may be the anterior pedal mucous gland. Predation on sphaerotheriid millipedes may be a behavioural apomorphy associated with a clade within the Chlamydephoridae, including taxa with longitudinal body keels and distinctive radula morphology, but it is evidently not confined to this clade. Whether the diet of such millipede-eating slugs is limited to millipedes is unknown, but it is probable that at least some chlamydephorids feed on more conventional prey such as snails and earthworms.

Growth in body mass in pups of the subantarctic fur seal Arctocephalus tropicalis was studied at Amsterdam Island, southern Indian Ocean during the rearing period between December 1995 and August 1996. We examined the change in the rates of mass gain and mass loss with pup age to explain the overall change in mass throughout the rearing period, when mothers alternate foraging trips at sea and attendance periods ashore. In the absence of their mothers, pups fast and lose mass. During the maternal attendance period ashore pups suck and gain mass. There was no significant difference in the rate of growth in mass between male and female pups. The rate of loss in mass during a fasting event was positively related to the initial body mass but negatively related to the age and duration of the fast of the pup. The rate of mass loss was best related to pup age by a quadratic relationship. The rate of mass loss increased until pups were 120 days old and then decreased until the end of the study period. The rate of gain in mass following a maternal foraging trip was positively related to the initial pup mass but negatively related to the foraging trip duration, and we found that the rate of mass gain decreased linearly with the age of the pup. The concomitant decrease in the rate of mass loss and mass gain for pups > 120 days old allowed pups to maintain a positive growth until they were 220 days old. After 220 days of age the rate of mass loss exceeded the rate of mass gain and the pups start to lose mass until weaning.

Oreochromis alcalicus grahami is a small cichlid fish that lives in the hot, highly alkaline, highly saline peripheral lagoons of Lake Magadi, Kenya. The fish faces profound diurnal oscillations of oxygen concentration. During the day, from photosynthetic activity of cyanobacteria (blue-green algae), the water is supersaturated with oxygen but after sunset when photosynthetic activity stops the water is virtually anoxic as a result of bacterial respiration. During the night and after explosive exercise, O. a. grahami characteristically skims the surface of the water with the mouth agape, suggesting that the buccal cavity is used as a gas-exchange organ. Contrary to expectation, however, the buccal cavity was found to be conspicuously non-vascularized: the surface epithelial lining was fundamentally of a mucus secretory type. In addition, certain cells in the deeper layers showed extensive lateral labyrinths similar to the epithelium of the renal tubules. These morphological features respectively indicated roles of secreting a protective film and regulation of ions taken across the epithelial lining of the buccal cavity. The allocation of gas-exchange to the gills and the air-bladder, osmoregulation essentially to the gills, and mucus secretion/protection to the buccal cavity displays an adaptive trade-off process in an elite animal. Effective use of the buccal cavity as a gas-exchanger would entail air-gulping followed by brief retention of it in the cavity to allow oxygen uptake. During such interval, both the gills and the air-bladder would of necessity be rendered temporarily non-functional. Skimming the top layer of water with the mouth open ensures that the gills are passively ventilated with well aerated water and the air-bladder is simultaneously used for gas-exchange, a strategy that should enhance oxygen acquisition, especially at higher ambient temperatures.

In non-native conifer plantations characterized by strong spatial and temporal variations in the availability of tree seeds in Spadeadam Forest, northern England, the home range and habitat use of red squirrels Sciurus vulgaris was very flexible. Males tended to have much larger home ranges than females and core-areas of most breeding females seemed mutually exclusive. Adult female red squirrels were found to increase their home range and core-area size in forest patches where food was less abundant. Home-range size was significantly related to home-range quality and the extent of overlap by other females. In contrast with high-quality continuous conifer forests: (1) a considerable proportion of adult males and females at Spadeadam shifted home range, (2) both sexes had much larger home ranges than reported from other habitats in Britain or Belgium. Many ranges were multinuclear, particularly from January onwards, when supplies of seeds become depleted through consumption and seed shed. Squirrels tracked the availability of conifer seeds (lodgepole pine cones throughout the study, Norway spruce cones in spring 1992 and Sitka spruce cones in autumn 1993) and intensively used several non-adjacent activity centres in temporally food-rich patches. Consequently, habitat preference changed markedly with time. The squirrels seemed to maximize nitrogen intake and to avoid the smaller seeds when possible. This resulted in an overall preference for a mixed diet of lodgepole pine and spruce seeds and avoidance of Sitka spruce seeds when Norway spruce seeds were available. These results lend support to the hypothesis of Ostfeld (1985) that when food is sparse and patchily distributed, females should develop intrasexual territoriality, concentrating their activity in food-rich patches, while males should be non-territorial and adapt their space use to the distribution of females.

The duration of activity bouts by pine martens Martes martes was studied in the pristine deciduous forests in Białowieża National Park (north-eastern Poland). From 1991 to 1996, 14 martens were monitored by radio-tracking throughout 5823 h. On average, male martens were active for 9 h/day and females for 8.5 h/day. The duration of the martens' diel activity varied greatly among bimonthly periods (from 2.8 h/day in February–March to 12 h/day in June–July). Martens decreased their activity from 13 h/day on warm days to 2.5 h/day on cooler days. In the winter months, when martens curtailed their activity, they often hunted larger prey or scavenged ungulate carcasses. On average, 69% of the martens' active time was during the night. The number of activity bouts per day varied from one to six (mean 2.6) and increased with growing ambient temperature. Activity bouts of males were significantly longer (4 h, on average) than those of females (3 h) and were not related to ambient temperature. The duration of short inactive bouts increased in the cold season and inactivity lasted longer in females than in males.

Two newt species with contrasting ecological requirements (Triturus cristatus and T. marmoratus) were radio-tracked after leaving a shared breeding pond in western France. Movements of 30 individuals were recorded over 491 days. During the first terrestrial night, the newts migrated with high directionality up to 137 m to their refuges. Burrows of small mammals were among the favourite refuges. Movements after the first night were mostly underground and over shorter distances (< 6.8 m). Sixty-four per cent of all tracked newts stayed within 20 m of the pond edge, and migrations were recorded up to 146 m away from the pond. Both species preferred areas with bushes, hedgerows and trees, and avoided pastures and open areas. Migration in the direction of a habitat type characterized by trees and underground shelters was favoured over migrations in other directions. A clear terrestrial niche separation between the two newt species was not observed. The results are discussed in relation to previous findings that T. cristatus supersedes T. marmoratus as a consequence of anthropogenic change to the terrestrial environment, affecting the species differentially.

Nephila edulis is a spider with large body size variability in males and females. Genital characters show negative allometric values compared to somatic characters. In males, the embolus (the most important structure for sperm transfer) had a significantly lower coefficient of variation than body size. This suggests that male genitalia are under stabilizing selection favouring intermediate size. Female N. edulis showed a trend similar to males regarding allometric values in genitalia. In females, however, the variation coefficient in a specific genital character crucial for successful copulation did not differ from that of indicators for overall body size. This suggests that in Nephila the genitalia of the females experience less stabilizing selection than those of the males. In male and female genitalia, the mode of selection seems to cause developmental instability not in degrees of fluctuating asymmetry but in the degree of data scatter which indicates a lower coefficient of determination.

Movements and group structure of giraffe Giraffa camelopardalis were studied in Lake Manyara National Park, northern Tanzania. The giraffe population in Manyara had increased from 60 to 85 individuals between the early 1980s and 1991. This increase may have been the result of an increase in browse availability as a result of a dramatic decline in elephant numbers, and bush encroachment following a series of anthrax epidemics that killed impala. Giraffe densities in Manyara are high compared to other areas within the Masai ecosystem, and Manyara probably serves as a dry season refuge. Females were found in small, yet unstable groups, while males associated randomly with each other. Neither males nor females were confined to single localities, although home ranges were small compared with studies in areas with low giraffe densities. In one area within the park a more stable group was found, and resident males who were probably defending a temporary harem, engaged in necking contests with immigrant males. A comparison with other studies showed that giraffe density, home-range size, mobility and group stability differ across different habitats. The tendency that more stable groups are found in high density areas might be taken as evidence for the occurrence of resource defence polygyny in such areas.

Conventional wisdom, based on properties of reagent-grade salts, states that organisms which construct their exoskeletons from calcite rather than aragonite have a greater ability to resist dissolution. However, experiments on individual bivalve microstructures show that other factors such as crystal size and the proportion of organic matrix outweigh constituent mineralogy in determining the rate of shell loss in cold waters. Moreover, the loss of shell material was minimal, corresponding to an annual loss in shell thickness of 3–40 μm depending on microstructure. These findings must cast doubt on the commonly held supposition that calcitic layers added to the outside of shells are an adaptation against dissolution in Holocene ‘Aragonite’ seawater.

This study sought to determine if the tropical loliginid squid Photololigo sp. stores energy in the form of lipid, carbohydrate or protein for reproductive investment. Individuals were examined for changes in morphometry, mantle muscle structure and concentrations of water, lipid, carbohydrate and protein in muscle tissue and the digestive gland, associated with the stage of reproductive maturation. Muscle mass was affected by reproductive maturation in females. Mature individuals were lighter for their length compared with females in the early stages of maturation. Concentrations of lipid and carbohydrate in the muscle tissue were very low, and female Photololigo sp. showed equivocal evidence of declining lipid and carbohydrate levels with egg production. There was no evidence of dramatic changes in protein concentration in the mantle muscle with reproductive maturation. Male Photololigo sp. showed a change in the digestive gland with maturation, with water content increasing and protein concentrations decreasing. The digestive gland of both male and female Photololigo sp. increased in size and contained less water with growth. There was little evidence that the storage and transfer of energy for reproduction occurred in Photololigo sp. Instead, it is probable that energy for reproduction is predominantly sourced directly from consumed food.

Parental recognition of offspring would be expected to evolve among densely populated species in which parental investment is large, and separations of the mother–pup pair are frequent. Although otariids present a well-developed system of mutual vocal recognition, evidence in phocids is weak. Furthermore, allo-suckling is prevalent in some species and may reflect confusion over the identity of pups. The vocalizations of grey seal Halichoerus grypus pups have been found to be stereotyped and individually distinctive on the Isle of May, Scotland. However, playback experiments showed that mothers fail to discriminate between the vocalizations of their pup and a non-filial pup. Moreover, widespread allo-suckling was observed. In contrast, grey seals on Sable Island, Nova Scotia, a population reproductively isolated from that of the Isle of May, show very little allo-suckling. This may reflect the presence of a recognition system that does not seem to be present on the Isle of May. During the 1999 breeding season on Sable Island, vocalizations of grey seal pups were recorded and playback experiments carried out to determine whether female grey seals respond differently to the playback of vocalizations of their own pup, a familiar pup and an unfamiliar pup. Grey seal mothers were found to make significantly more head turns and body movements towards the loudspeaker during the playback of their own pup call than during the playback of a familiar or unfamiliar pup call. In addition, there was no evidence of an effect of pup age. This suggests that female grey seals can discriminate between pup calls using the stereotyped and individually distinctive vocalizations of their pup, and that different selective pressures may be in operation between the Sable Island and Isle of May colonies.

The whiptail lizard Cnemidophorus vanzoi is a large-bodied teiid found only on two islands (Maria Major and Maria Minor), off St Lucia, West Indies. In May 1995, 42 lizards from Maria Major were introduced to the smaller uninhabited Praslin Island on the same coastline. Three years post-release, we studied abundance, demography and morphometrics of the translocated lizard population, during a 6-month period covering wet and dry seasons. Age, sex, snout–vent length (SVL), body mass (BM), tail length, tail regeneration, and overall condition (moulting, reproductive condition, cuts, external parasites) of 107 animals caught during the study are analysed in the present paper. Comparisons are also made with the source population. A body condition index (CI (BM/SVL)), sex ratio (adult males : adult females), age ratio (adults : juveniles), and sexual size dimorphism ratio (SVL adult male : SVL adult females) were calculated for the study population. Distance sampling and mark–re-sight surveys were used to estimate population size and lizard density. A total of 155 ± 26 individuals were estimated. The lizard population was found to have a high growth rate (r = 0.97–3.95). There were significant seasonal changes in lizard abundance. Seasonal differences in lizard numbers, BM and CI suggest either severe resource limitation during the dry season, or selective aestivation. A high frequency of tail autotomy may point to intense intraspecific competition as the island is relatively free from main predators such as the black rat Rattus rattus. Sex ratio, sexual size dimorphism and sexual dichromatism all indicate a territorial species in a generally non-territorial family (Teiidae). Some adult males seem to maintain juvenile colours. It is suggested that the introduced population has successfully colonized its new environment and that no significant change in the animals condition or size has occurred during the 3 years since translocation.

Two Iberian wolf Canis lupus signatus packs were studied in the north of Portugal during spring and summer 1996. The study areas are located in two mountainous areas close to the Spanish border. Both regions presented a very low density of wild ungulates and intensive livestock production. The food habits and the livestock depredation of the wolves were considered. The study material included 87 dropping samples, interviews with shepherds, carcass investigation and government reports of livestock depredation. Prey preference was measured in both areas. Wolves fed exclusively on livestock, especially goats. Wolf attacks on goats affected mostly large flocks of > 100 heads, but where there were horses, wolves preyed preferentially on horses. In both study areas, sheep was recorded as a regular prey by the government reports but never appeared in the scat analysis results. This apparent contradiction will be discussed. The wolf's dependence on livestock can be explained by the scarcity of wild prey and the high density of livestock. Conservation of wolves in such impoverished areas depends on an efficient livestock depredation management plan and the reintroduction of native prey species.

Sixteen microsatellite markers were used to determine the genetic relatedness between adult female polar bears and their accompanying cubs for three recent occurrences of natural adoption recorded in the Canadian Arctic. The estimation of the degree of relatedness was powerful and provided a high degree of separation between first order relatives and unrelated individuals with the loci used. The analysis showed that the cubs were unrelated to their adoptive mothers and that the mothers were unrelated to one another. Although the circumstances and mechanisms whereby these adoptive events occurred are unknown, the data do not support kinship theory as an explanation for the exchange of cubs. We suggest that these adoptive events probably represent instances of the misidentification of cubs by adult polar bears.

The present study provides comparative data on the types and microhabitats of animal prey captured by associating saddleback Saguinus fuscicollis and moustached Saguinus mystax tamarins. The main prey items were katydids (Tettigoniidae, Orthoptera), stick grasshoppers (Proscopiidae, Orthoptera) and spiders (Araneida), although lizards (Squamata) and frogs (Anura) were also taken. The tamarins foraged for, and captured prey from, significantly different substrates. This resulted in a significant partitioning of the prey spectrum between the two species. Sylvan katydids (Pseudophyllinae) accounted for most of the prey of both species, but the moustached tamarins captured more bush katydids (Phaneropterinae) than the saddlebacks. The degree of dietary overlap was less for animal prey than the plant-based portion of the tamarins' diet.