Adélie penguins Pygoscelis adeliae were tested as to whether they jump with optimal energy efficiency when moving out of the sea to the land. Adélie penguins risk predation if the jump fails. Swimming penguins usually launch up the side of sea ice to a surface higher than sea level. Analysis of jumping behaviour recorded by a video camera showed that the trajectory of the centre of gravity of the birds during the aerial phase of jumping was parabolic, indicating that the success of landing depends on three parameters at the time of take-off from water: speed, angle and distance from the point of emergence to the ice edge. There was a negative relationship between distance and the take-off angle, suggesting that penguins adjust their take-off angle to the distance from the ice edge. The comparison among hypotheses revealed that penguins did not jump with optimal energy efficiency. Instead, they aimed for the refracted image of the edge of the cliff, which from underwater appears higher than it actually is. This direction-dependent rule seems to be more robust and reliable than the optimal energetic strategy.

Adult female survival as a potential proximate factor responsible for observed changes in southern elephant seal Mirounga leonina populations was investigated. We compared the survival rate estimates from mark–recapture data for female elephant seals from the Marion Island population (using program MARK) for two periods (pre- and post-decline) during the past 15 years and with estimates from another population in southern Argentina, which had increased steadily during the same period. Survival of prime-age adult females increased significantly by 6.2% during the latter part of the decline at Marion Island, and the survival of adult females at the colony in southern Argentina was 3.2% greater than at Marion Island after the stabilization. We thereby demonstrated the importance of adult female survival in population regulation and emphasized the importance of monitoring adult females in order to understand population changes in southern elephant seals. In addition, we investigated whether reproductive expenditure early on in life reduces future reproductive potential in the population at Marion Island. We did this by estimating and comparing future survival and breeding probabilities of females primiparous at different ages. The future annual survival and breeding probabilities of females breeding at a young age, was similar to those from females primiparous at an older age. There was also no reduced survival in the year following first breeding in young or older first time breeders. Reproductive expenditure in young primiparous females therefore did not entail future fitness costs relative to older primiparous females, and we found no evidence supporting the existence of various life-history strategies in terms of age of primiparity within a population of southern elephant seals.

The evolutionary history of the lion Panthera leo began in Pliocene east Africa, as open habitats expanded towards the end of the Cenozoic. During the middle–late Pleistocene, lions spread to most parts of Eurasia, North America, and may have eventually reached as far south as Peru. Lions probably evolved group-living behaviour before they expanded out of Africa, and this trait is likely to have prevailed in subsequent populations. The first lions were presumed to have been maneless, and maneless forms seem to have persisted in Europe, and possibly the New World, until around 10 000 years ago. The maned form may have appeared c. 320 000–190 000 years ago, and may have had a selective advantage that enabled it to expand to replace the range of earlier maneless forms throughout Africa and western Eurasia by historic times: ‘latest wave hypothesis’.

The home range and movements of the quokka Setonix brachyurus, a medium-sized macropodid marsupial, were investigated using radio telemetry. Fifty-eight quokkas from five remnant mainland populations in the northern jarrah (Eucalyptus marginata) forest of Western Australia were radio-collared and monitored for up to 2 years between 1998 and 2000. Mean (±SE) home-range sizes were 6.39±0.77 ha and core ranges averaged 1.21±0.12 ha. Male core home ranges were larger than those of females although not when corrected for body mass. Nocturnal ranges were larger than diurnal owing to nocturnal departures from the swamp. Ranges shifted to the edge of swamps in winter, as the swamps became inundated following rain, and toward the centre in autumn as the swamps dried. Quokka populations are thought to exist below carrying capacity at all sites and this probably caused the lack of dispersal observed in this study. We hypothesize that predation is suppressing population booms which would otherwise drive dispersal and thereby maintain metapopulation dynamics. Without dispersal to rescue unpopulated patches, we conclude that the original quokka metapopulation has collapsed and ameliatory measures are required.

Photo-identification surveys of humpback whales Megaptera novaeangliae were conducted at West Greenland during 1988–93, the last 2 years of which were part of the internationally coordinated humpback whale research programme YoNAH, with the primary aim of estimating abundance for the West Greenland feeding aggregation. The area studied stretched from the coast out to the offshore margin of the banks, determined approximately by the 200 m depth contours, between c. 61°70′N and c. 66°N. The surveys were conducted between early July and mid-August and 993 h were expended on searching effort. A total of 670 groups of humpback whales was encountered leading to the identification of 348 individual animals. Three areas of concentration were identified: an area off Nuuk; an area at c. 63°30′N; and an area off Frederikshåb. Sequential Petersen capture–recapture estimates of abundance were calculated for five pairs of years at 357 (1988–89), 355 (1989–90), 566 (1990–91), 376 (1991–92), and 348 (1992–93). Excluding the anomalously high estimate in 1990–91, the simple mean is 359 (SE = 27.3, CV = 0.076) and the inverse CV-squared weighted mean is 356 animals (SE = 24.9, CV = 0.070). These calculations lead us to conclude that between 1988 and 1993 there were 360 humpbacks (CV = 0.07) in the West Greenland feeding aggregation. Using the Cormack–Jolly–Seber model framework non-calf survival rate was estimated at 0.957 (SE = 0.028). Our data have low power (P < 0.3) to detect a trend of 3.1%, assuming the probability of a type I error was 0.05.

It is generally accepted that predation by wolves Canis lupus is one of the major factors limiting densities of woodland caribou Rangifer tarandus caribou in North America. Conversely, little is known about the role of European wild forest reindeer R. t. fennicus as wolf prey, or about the influence of wolf predation on populations of this rare subspecies. This relationship was examined in east-central Finland, where wild forest reindeer coexist with moose Alces alces at a numerical ratio of c. 1 reindeer to 1.5 moose. During the study, moose were clearly the primary prey of wolves. Reindeer were, however, an important part of wolf diet in summer, autumn and early winter, when their remains comprised roughly 20–50% of all food items identified in wolf scats. Wolves exhibited a slight preference for reindeer over moose as prey during early winter (November–December) when the reindeer were moving towards their wintering ranges. Virtually no reindeer were killed by wolves during midwinter (January–March). This held also for a pack whose territory was located in the middle of the wintering range of reindeer, where reindeer outnumbered moose during the winter months. After the approximate ratio of wolves to reindeer increased from 0.004 (1998) to 0.02 (2000), wolf predation became the most common source of mortality (50%) for reindeer. The annual net increase in the reindeer population decreased from 13% to 7% because of increased wolf predation. The results indicate that wolf predation was an important factor limiting reindeer population growth. Wolves were, however, unable to prevent entirely the number of reindeer from increasing, partly owing to behavioural traits used by reindeer during winter and calving times.

Neotropical rainforests support guilds of nectar feeding bats (Phyllostomidae: Glossophaginae) with up to six coexisting species. To analyse guild structure and mechanisms of coexistence in a Costa Rican tropical lowland rainforest, the resource use and morphology of bats were compared to the energetic characteristics of preferred nectar resources and their spatio-temporal distribution. The relative abundance of nectar-feeding bats was determined from mistnet captures over 26 months. Food items were identified by analysis of pollen loads and faecal samples. Phenology, flower density and nectar sugar content of resource plants permitted quantitative estimations of resource availability expressed as energy density (kJ ha−1 day−1) throughout the annual cycle. Four glossophagine bat species co-occurred at La Selva: two permanent residents (Glossophaga commissarisi, Hylonycteris underwoodi) and two seasonal species (Lichonycteris obscura, Lonchophylla robusta) that were found in small numbers during a period of high nectar availability. The two resident species differed in their abundance and in their temporal feeding strategies. After the main flowering peak, the common G. commissarisi shifted to a more frugivorous diet, while the rarer H. underwoodi fed on the few remaining bat-flowers. Resource plant species differed in their energy density by up to two orders of magnitude. Hylonycteris underwoodi visited more often plant species with a low energy yield than G. commissarisi. Because of its smaller body size and a wing morphology that promotes fast flight, H. underwoodi appears to be better adapted to low and scattered nectar resource levels. The two seasonal species differed greatly in body mass, which suggests different strategies for high-quality resource tracking. Large body mass in Lonchophylla robusta provides an energy buffer that permits daily commuting flights between a permanent roost and profitable foraging areas, while the small Lichonycteris obscura seems to track resources nomadically. It is proposed that energy density may be a major niche dimension that restricts access of species to certain habitats and that may profoundly influence the structure of nectar-feeding bat guilds.

Genetic differentiation among populations of the South African scrub hare Lepus saxatilis was examined using hypervariable mitochondrial DNA control region I (CR-I) sequences. Neighbour-joining analysis revealed a pattern that did not correspond to the current subspecies delineations. The CR-I sequence data delimit scrub hares into three major maternal lineages. The three phylogenetic assemblages exhibited different geographical distributions. AMOVA analyses and exact tests for population differentiation confirmed this phylogeographic partitioning. One lineage (SW) was confined to the south-western Cape, the second lineage (N) was exclusively found in the northern part of South Africa and in the neighbouring countries, and the third lineage (C) was predominant in the central parts of South Africa. This spatial distribution did not coincide with the ranges of the 10 described subspecies covered by our sampling regime. The lineages C and N overlapped in an area including eastern parts of South Africa and southern Namibia. The presence of both lineages in that area of overlap was interpreted as the result of secondary contact due to recent range expansions after the two lineages had undergone a population restriction approximately 18 000 years ago. Analyses of contemporary gene flow disclosed an exchange of migrants between N and C, which was biased towards a movement from C to N. The SW group represents a very distinct evolutionary lineage that has been isolated for more than 45 000 years. It does not exchange female migrants with the other two groups. Mismatch distribution analyses indicated sudden population size expansions in the history of all three populations.

Bone mass is dramatically compromised during periods of weightlessness, inactivity, or bed rest. Animals that hibernate reduce their body temperature, heart rate and metabolic activity, and likewise lower bone turnover activity during their immobile state. Black bears Ursus americanus, however, do not hibernate, but rather overwinter by denning during which they maintain a nearly normal functional heart rate, cardiac output and temperature. Furthermore, markers of bone turnover in black bears are maintained during denning periods. Thus, the purpose of this work was to determine if the denning state of relative immobility in black bears results in changes in bone mass and bone architectural structure. Harvested forelimbs (ulna and radius) were compared between pre- and post-denning black bears using X-ray imaging, dual energy X-ray absorptiometry, and micro-computed tomography to quantify total distal forelimb bone mineral density, cancellous bone mineral density, bone mineral content, bone volume fraction, degree of anisotropy, structure model index, and trabecular thickness. No significant differences in any of the measured parameters were found in comparing radius and ulna from autumn and spring bears in this cross-sectional sample, suggesting that black bears did not experience a significant change in bone mass or architecture during denning. The statistical power for detecting a significant difference (P ≤ 0.05) for this sample was 0.8. The specific mechanism by which the preservation of bone was attained may be related to skeletal muscle interaction or circulating systemic hormones.

Around the world, endemic species with restricted ranges seem to be at particular risk of extinction. When range size is controlled, lowland continental species may be especially at risk, more so than island or montane species. Our study aimed to investigate reasons behind the high vulnerability of endemic species. In Sri Lanka, large-scale national surveys and intensive localized fieldwork established that endemic plant and animal species are mostly associated with undisturbed rainforest habitats. On the other hand, non-endemic species used both forest and non-forest habitats almost equally, suggesting they are less likely to suffer from deleterious edge effects. To understand the different distribution of endemic and non-endemic species, our study focused on rats. Compared to the widespread species, the endemic rat species had a larger home range and fed on fewer species of forest fruit, possibly indicating greater specialization. The abundance of the endemic species was also negatively correlated with the abundance of the non-endemic species, to which it was behaviourally subordinate. Greater specialization and competitive inferiority could both contribute to the vulnerability of endemic species.

The age of 86 individuals derived from groups of female sperm whale Physeter macrocephalus involved in three mass strandings on the north and west coasts of Tasmania in 1998 was determined from the number of dentinal growth layer groups in the teeth of individuals. Dorsal total lengths were also measured. Ages of females ranged from 0.75 to 64 years, with the majority (77%) aged between 20 and 45 years. Total lengths of female sperm whales ranged from 417–1200 cm, with 68% of females 1050–1200 cm long. Constraints associated with the age structure observed in this study and the representativeness of the age structure to that of the greater population are discussed. In an effort to assess the effects of underestimation of age estimates on age-specific demographic parameters, a model simulating changes in age structure as a result of tooth wear was developed and the resulting survival rates compared to those derived from the original age structures of the samples. Survival did not change significantly between the modelled and original age structures. Also, the survival rate calculated from Australian mature female sperm whales was compared to that calculated from sperm whales caught in whaling operations from Japanese waters. Australian mature female sperm whales demonstrated significantly higher survival rates (mean=0.905±0.046 SD range: 0.856–0.986) than mature females from Japanese waters (mean=0.885±0.034 SD; range=0.859–0.970), possibly the reflection of higher fishing mortality on the Japanese whales. Growth equations using Gompertz and von Bertalanffy models were calculated for female Australian sperm whales and compared. Both models described growth in female sperm whales similarly. Changes in the demographic parameters of sperm whales with the cessation of whaling may be reflected in the growth rates of individuals and as such, these equations may provide a useful tool for monitoring continuing changes in the demographic parameters of this species. Without long-term mark–recapture studies on sperm whales in this region, the frequent stranding of sperm whales in south-eastern Australia provides an important opportunity to collect data on the life history and demography of this species.

The behaviour exhibited by the hermit crab Pagurus longicarpus in response to an empty shell varied in the presence of cues from conspecific individuals according to its familiarity or not with them. This binary discrimination was independent of the conspecific's relative size and was based on chemical signatures, an ability that this species shares with a few other aquatic invertebrates. From our results, olfaction appeared to be the dominant sensory channel in P. longicarpus' binary discrimination, but the combination of two signal components from visual and olfactory channels resulted in the enhancement of the response displayed by the receiver. Besides, crabs reacted differently when exposed to their own odour than to the odour of familiar (as well as unfamiliar) conspecifics, suggesting that recognition in this species can be more refined than a binary discrimination and that chemical ‘badges’ may be attributes of individual crabs.

The dynamics of leopard seal Hydrurga leptonyx abundance and residency were investigated both within and between years at Bird Island, South Georgia, using observations of tagged and untagged individuals. Seals belonged to an open population, and were sampled outside the breeding season and during their northward winter dispersal (April–October). Seals at Bird Island were highly transient (41–63%) with only a limited number of individuals making repeat visits across years. A minimum yearly survival for resident seals was estimated to be between 0.61 and 0.85. The physical environment played an important role in leopard seal abundance, with observations following an apparent 4-year periodicity consistent with the frequency and timing of the Antarctic Circumpolar Wave. Years of low sea-surface temperature (SST) and extensive pack-ice corresponded to a higher number of seals present, earlier arrivals, and longer periods of residency. Peak leopard seal arrivals at Bird Island corresponded to periods of low SST, which was consistent with the pagophilic nature of this species.

The grey-headed flying fox Pteropus poliocephalus, a nationally vulnerable species, is endemic to coastal eastern Australia, from Maryborough, Queensland, through New South Wales to Melbourne, Victoria. Pteropus poliocephalus forages at night, primarily on eucalypt blossom within 50 km of traditional camps (day roosts), usually in dense, riparian vegetation. Several attempts have been made to track long-distance movements of P. poliocephalus across its extensive and climatically highly variable range, but the technology has been inadequate for tracking at the required speed and scale. Satellite tracking was used to monitor movements of a 2-year-old male P. poliocephalus, trapped at the Currie Park camp in the north of the species' range (28°48′S) from January 2000, and another from its most southerly camp, Melbourne (37°50′S), from April 2000. The first flying fox moved camp 50 km noth-east to Dallas Park, where it stayed until March, then transited at least 15 other camps between 28°12′–32°44′S, before returning to Dallas Park in September. The second flying fox remained in Melbourne until July, then transited at least six other camps to 33°44′S, before returning to Melbourne in January 2001. Both animals made round trips of at least 2000 km, traversing >4° latitude, before returning to their camps of origin, while other animals remained. The study suggests that P. poliocephalus is a partial migrant that uses winds to facilitate long-distance movements, and underlines the importance of management at a national scale.

In polygynous mammals, fitness differences may reflect differences in phenotypic quality as well as experience. This study determines dominance hierarchy among female reindeer Rangifer tarandus from two experimental herds (consisting of c. 45 animals in each) during 2 consecutive years. The influence of body mass, antler size and age on social rank in the herds was investigated, first using simple regression analysis. The combined effect of body mass, age and antler size on female rank was further assessed using principal component analysis, as these three parameters were significantly correlated. The improved Laundau linearity index of c. 0.5 (Pr < 0.001) in both herds indicated that a substantial part of the hierarchies was explained by their linearity properties. Consistently, body mass, female age and antler size, as well as their combined effect (measured by the scores on the main axis, PC1), influenced social rank in all four groups. It was concluded that both body mass and age are good predictors of social rank in female reindeer, whereas antler size in comparison plays a less important role in herds with a ‘normal’ female age structure. This suggests that female antlers may have evolved in intersexual rather than intrasexual competition. The temporal variations in the importance of body mass and age, probably owing to variation in female age structure between the 2 years, calls for conservative interpretations of whether body mass or age is more important in determining social rank among female reindeer. This is confirmed by the PCA analyses, where all three variables contributed more or less equally to the first component, the size variable, which on average explained more of the variation in female rank than body mass and age, suggesting that phenotypic quality expressed as the combination of the three variables is a better predictor of social rank than the variables per se. Hence, general conclusions about social rank based on single studies including few animals may not be credible.

In primates, age determination using lines of arrested growth (LAGs) from bones has rarely been attempted, and the reliability of these structures has never been experimentally validated. In order to test skeletochronology in primates, LAGs were studied mainly in the long bones of known age Microcebus murinus, a small primate, whose potential longevity may reach 12 years. LAGs were extensively studied in 43 males and 23 females ranging from juveniles to 11-year-old adults. All individuals were born and reared in captivity. Some young individuals were injected with fluorescent dyes to quantify bone growth rates. LAGs in the diaphysis of the tibia are well correlated with age and this skeletal element appears to be the best for assessing skeletochronology in Microcebus murinus. There is strong evidence that the seasonal cycle of photoperiodicity is more important than age alone in producing LAGs.

Patterns of morphological variation in the skulls of extant bears were studied as they relate to diet and feeding behaviour. Measurements of craniodental features were used to compute indices that reflect dietary adaptations of the dentition and biomechanical properties of the skull, jaw and related musculature. Species were classified as either carnivores, omnivores, herbivores or insectivores. Differences among dietary groups were assessed with analysis of variance and discriminant factor analysis. Results demonstrated significant morphological separation among all four groups. Carnivores were distinguished by, among other features, molar size reduction, flexible mandibles and, most surprisingly, relatively small carnassial blades. In contrast, herbivores displayed, among other features, large molar grinding areas, rigid mandibles and large carnassial blades. The insectivorous sloth bear was characterized by extreme reduction of the post-canine teeth. As expected, omnivores tended to have morphology intermediate between that of carnivorous and herbivorous ursids. Comparison with previous studies revealed that bears exhibit a different set of morphological specializations for diet than other carnivoran groups. Carnivorous ursids, for example, were found to share aspects of craniodental morphology with omnivorous canids. The relatively weak adaptations for carnivory observed in bears may be the result of selection for the ability to cope with temporal fluctuations in dietary components. The giant panda Ailuropoda melanoleuca was found to have a relatively stiff jaw and great mechanical advantage of the jaw-closing muscles, features previously observed in carnivorous canids and unexpected in this herbivorous bear. Comparison of patterns of morphological variation and patterns of phylogenetic relationships among species revealed surprisingly strong congruence between morphology and phylogenetics.

The Namaqua dune mole-rat Bathyergus janetta breeds seasonally, producing up to two litters during the early summer in the Cape. Females exhibit elevated urinary progesterone and oestradiol-17β concentrations and males raised concentrations of testosterone, these correlate with the onset of the seasonal winter rainfall. The rains moisten the soil allowing excavation and provide the mole-rats with the opportunity not only to extend burrow systems but also to search for mates. An increased incidence of hindfoot drumming and visitations to neighbouring burrow systems occurs during this period. Plural occupancy of burrows by adult mole-rats was found during the onset of the winter, and pups, lactating and pregnant females were caught at the end of October and November (early Cape summer). Pups develop rapidly and their eyes open after c. 1 week. Pup body mass increases exponentially from birth to about 70 days. Inter-sibling aggression increases progressively and after c. 60 days captive-born pups had to be separated, suggesting that in the wild they would have dispersed.

The phylogenetic relationships within the family Talpidae (Mammalia, Insectivora) were inferred from parsimonious analysis of 55 morphological characters, which were collected from 43 skulls of 22 species, and which included all of the extant genera in the current taxonomy of the Talpidae. Two equally parsimonious cladograms were obtained. The Uropsilinae, Desmaninae and Talpinae subfamilies formed monophyletic clades, and the Desmaninae and Talpinae subfamilies were sister clades. The Talpinae subfamily was subdivided into the following clades: Urotrichus, Dymecodon, Scalopus–Scapanus, Parascalops–Scapanulus, Condylura, Neurotrichus, Scaptonyx and Talpini (Talpa, Euroscaptor, Scaptochirus, Parascaptor and Mogera). The Urotrichini (Urotrichus, Dymecodon and Neurotrichus) and Scalopini (Scalopus, Scapanus, Parascalops and Scapanulus) were considered to be paraphyletic, and were defined primarily by their ancestral characteristics. Since the genus Dymecodon was not clustered with Urotrichus, it was considered to be a valid genus, and not just a junior synonym of the latter. The Asian genus Euroscaptor was found to be a paraphyletic group, and deserves further systematic revision. The evolution from semi-fossorial habit to fully fossorial habit seems to have occurred three times in the Scalopus–Scapanus, Parascalops–Scapanulus and Talpini clades. The migration of the Talpinae to North America from Eurasia probably occurred on four separate occasions in the Scalopus–Scapanus, Parascalops, Condylura and Neurotrichus lineages.

In the present study the evolutionary relationships among Cordylusniger–oelofseni–cordylus are investigated using molecular sequence data to: (1) estimate the divergence within this group; (2) date the origin of melanism in the clade and to evaluate possible palaeoecological events responsible for its evolution; (3) examine the systematic relationships among the three isolated populations of C. oelofseni that were previously shown to be polyphyletic using allozyme data. The recovered topology for the combined sequence data (16S and NADH dehydrogenase component 2 (ND2)) among C. niger–oelofseni–cordylus suggests that melanism evolved only once in the clade. The application of a molecular clock to the data demonstrates that melanism evolved during the Miocene epoch, 17–15 million years ago. We believe that the development of a cold-water current and upwelling system along the south-west coast of Africa during this epoch was instrumental in the evolution of melanism in this lizard clade and possibly also in other ectotherm clades with melanistic forms occurring in the area. The results show that C. oelofseni, as presently construed, is composed of three distinct evolutionary lineages, with no shared haplotypes present and marked sequence divergence. Two of the three lineages represent undescribed species. Cordylus cordylus is composed of two distinct clades, a montane clade and a coastal lowland clade. Cladogenic activity in mountain areas of the Western Cape in South Africa has been particularly pronounced and the systematic diversity of many taxa may have been underestimated.