Differences in facial skeletal form arise in part through differences in growth and in part through differences in embryonic patterning and proportioning. This study examines post-natal facial growth, sexual dimorphism and adult differences within and between two primate species: Cebus apella and Cercocebus torquatus. The species were chosen because they are phylogenetically distant yet present roughly similar adult morphologies. The comparison of ontogenetic, sexual and phylogenetic differences between these species illuminates the relationship between pre-natal proportioning, growth, sexual dimorphism and evolutionary change in the facial skeleton.

The quantitative analytical methods used in this study are taken from the geometric morphometric toolkit. Additionally, facial growth is examined through scanning electron microscope studies of bone growth remodelling. The main findings are that adult sexual dimorphism differs markedly between the studied species. In both species, however, it arises through extension in males relative to females of a common size–shape growth trajectory plus late divergence of growth between the sexes. This similarity contrasts with the findings that growth remodelling in the facial skeleton and size-related shape changes in the face of each species differ. Thus, variations in facial form between species seem to be the result of pre-natally established differences in form plus post-natal growth divergence, whilst the different sexual dimorphisms of these species arise through the action of similar growth processes on divergent growth trajectories. The implications of these findings for the interpretation of adult variations in facial form and growth are considered.

Inter-sexual segregation between habitats or between sites within a habitat is very widespread in sexually dimorphic ungulates. As an explanation, it has been suggested that males, because of their larger forage requirements, are driven from preferred, high quality forage habitats into marginal habitats of lower forage quality but higher forage biomass by female competition (‘indirect competition hypothesis’). However, this hypothesis has hardly been tested. In the present study, we tested the indirect competition hypothesis for site segregation of red deer on the Isle of Rum, Scotland, within two vegetation communities: short Agrostis/Festuca grassland and Calluna heath. We predicted that: (1) males should choose sites within each vegetation community that are of higher forage biomass but lower forage quality, than sites selected by females; (2) the degree of inter-sexual site segregation should be highest within the vegetation community in which indirect competition was most intense. We found that the sexes were significantly segregated between sites within both vegetation communities, but males did not use sites of lower forage quality or of higher forage biomass than did females. Moreover, degree of site segregation was not significantly higher in the vegetation community in which competition was more intense (i.e. short Agrostis/Festuca grassland). We conclude that the indirect competition hypothesis cannot explain site segregation in red deer on the Isle of Rum.

The relationships between reproductive condition, level of reproductive investment and adrenocortical modulation to capture stress in marine turtles form the basis of this study. When subjected to either capture or ecological stressors, nesting marine turtles have demonstrated adrenocortical responses that are both small in magnitude, and slow in responsiveness. These observations were further investigated to determine whether this minimal stress response was a physiological strategy to maximize reproductive investment in adult green Chelonia mydas and hawksbill Eretmochelys imbricata turtles. Female green and hawksbill turtles exhibited a decrease in adrenocortical responsiveness with progressive reproductive condition. Breeding turtles exhibited most suppression of their adrenocortical response to capture compared to both non-breeding and pre-breeding female counterparts. Nesting green turtles maintained a suppressed adrenocortical response to capture throughout the nesting season despite decreased reproductive investment. In contrast, male green and hawksbill turtles were less able to modulate their corticosterone (B) response to acute capture stress. During breeding, male turtles possessed significantly greater adrenocortical responses to capture than females. These results could indicate that the large reproductive investment necessary for female marine turtle reproduction might underlie the marked decrease in adrenocortical responsiveness. This hormonal mechanism could function as one strategy by which female marine turtles maximize their current reproductive event, even though under certain situations this mechanism could entail costs to female survival.

The mormyrids are well known for their remarkable electrogenic and electrolocation capabilities and exceptionally large cerebellum that may account for much of their total oxygen consumption. Mormyrids living in oxygen-deficient waters may use oxygen efficiently, protecting the brain from hypoxia damage; and/or brain size may be reduced. This study compares the TFL, gill lamellar density, gill lamellar area, total gill surface area, and brain size of two species of mormyrids from extremely hypoxic waters in Uganda (Gnathonemus victoriae and Petrocephalus catostoma) to two open-water species (Mormyrus kannume and Gnathonemus longibarbis) from the same region. In addition, interdemic variation was considered by comparing swamp populations of G. victoriae and P. catostoma to open-water populations of the same species. Total gill surface area of all species fell within the upper range for freshwater fishes. However, there were both intraspecific and interspecific differences in gill characters. Interdemic comparisons showed larger gill size in swamp-dwelling populations. Brain size varied among species; larger brains were characteristic of species from well-oxygenated waters. Large gill surface area may permit survival of mormyrids in oxygen-stressed environments; however, mechanisms compensating for hypoxia seem to be inadequate to support a brain size as large as that seen in fish from well-oxygenated waters.

The cheetah Acinonyx jubatus is the most cursorial felid and has been described as somewhat dog-like in both the shape and the diminished degree of retraction of the claws. In this study we evaluate and investigate the osteological correlates associated with claw retraction in the cheetah through a comparison of the morphology of its middle phalanges with those of other felids and of the wolf Canis lupus. Compared to other felids, the middle phalanges of the cheetah have better-developed grooving of the distal articulatory facets in both the manus and pes, reduced to absent angulation to the distal articulatory facets in the inner and outer digits, less marked asymmetry of the shaft, and a mid-shaft cross-section that is less triangular and more circular. In all of these features, the morphology of the cheetah is intermediate between that of other felids and that of the wolf. The cheetah's distinctive morphology is autapomorphic within Felidae and similarities between the cheetah and the wolf are the result of convergence. Study of an ontogenetic series of specimens of the domestic cat suggests that the morphology of the cheetah can be explained, at least in part, as a product of heterochrony in which the development of the middle phalanx is truncated at an earlier stage than is typical of the adults of other felids. Some of the morphological differences in the middle phalanges of the cheetah can be associated with its distinctive hunting behaviour. The reduced manipulative capabilities of the forelimb associated with the evolution of cursorial adaptations seem to have limited the roles of the forepaws in both the subduing of prey and feeding.

An existing spatial population model, developed to assess the reintroduction of the European beaver Castor fiber to Scotland, was applied to predict the likely result of reintroducing beavers to Norfolk, England. A habitat analysis was conducted using the Land Cover Map (classified satellite imagery developed by the Institute of Terrestrial Ecology) to predict the location of areas suitable for beavers. The model was used to simulate population processes within these areas and dispersal between them following a range of reintroduction scenarios. To represent uncertainty regarding the probable behaviour of beavers in Norfolk, the habitat analysis and model were applied using three sets of parameter values (high, medium and low) derived from the literature. It was predicted that Norfolk could support between 18 and 40 beaver families. Using high or medium parameter values, beaver populations were predicted to expand following reintroduction and spread to new areas. Using low parameter values populations were predicted to decrease, risk extinction and not colonize new areas. This analysis is a preliminary investigation into the likely fate of beavers in Norfolk. Checking the estimates of habitat availability in the field would enable greater confidence to be placed in these predictions.

Two expeditions were carried out during September 1997 and March 1999 to confirm the current existence of Mesocricetus auratus in northern Syria. Six females and seven males were caught at different sites near Aleppo. One female was pregnant and gave birth to six pups. Altogether, 30 burrows were mapped and the structures of 23 golden hamster burrows investigated. None of the inhabited burrows contained more than one adult. Burrow depths ranged from 36 to 106 cm (mean 65 cm). Their structure was simple, consisting of a single vertical entrance (gravity pipe) that proceeded to a nesting chamber and at least one additional food chamber. The mean length of the entire gallery system measured 200 cm and could extend up to 900 cm. Most burrows were found on agricultural fields preferentially on leguminous cultures. The distribution of golden hamsters is discussed in association with historical data, soil types, geography, climate and human activities. All 19 golden hamsters were transferred to Germany and, together with three wild individuals supplied by the University of Aleppo, form a new breeding stock.

Domestic lamb mortality on an open forested and alpine summer range in the municipality of Målselv in northern Norway was investigated. Two hundred and fifty-three lambs in four of the area's six flocks were randomly fitted with mortality transmitters and monitored throughout the summer grazing season from June to September. Total losses in the six flocks were 183 (22.9%) lambs and 20 (4.2%) ewes. The carcasses of 69 lambs, 36 of which had radio-collars, and six ewes were recovered. Of the collared lambs, eight (22%) died as a result of disease and 27 (75%) were killed by predators. Implicated predators included red fox Vulpes vulpes, lynx Lynx lynx, golden eagle Aquila chrysaetos and wolverine Gulo gulo. Diseased animals included those diagnosed with pasteurellosis, disrupted intestinal functions and advanced coccidiosis. The remains of one lamb were too decomposed for necropsy, though there was no evidence that it had been attacked by a predator. Factors associated with losses were identified using logistic regression. Age of lamb at time of release on the summer range and age of dam were statistically associated with lamb losses (P<0.05). Mortality was higher among older lambs and those of year-old ewes. Parameters commonly associated with lamb size and growth were not statistically associated with mortality. The negative association between lamb mortality and age of dam may be related to the quality of maternal care provided by ewes of different ages, particularly vigilance in the habitat of predators. Similarly, the positive association between mortality and age of lamb may be related to changing lamb behaviour, specifically increased distance from ewe, as lambs grow older and more independent. Predators, when present, are a major cause of mortality in free-ranging lambs, as in most wild ungulates.

This note presents plantar micro-tattooing (subcutaneous injection of Chinese ink) as a good alternative to toe-clipping, which is often used to identify individual rodents. The method was tested under laboratory conditions on voles Microtus arvalis from birth to sexual maturity. This easy, harmless and inexpensive technique could be used in the field, and not only on pups but also on adults of different species.

The emergence of Daubenton's bats Myotis daubentonii from a priory in the north of England was recorded to assess the impact of an annual music festival. Bat emergence was related to measured environmental variables using generalized linear modelling. The temperatures inside and outside the priory were significant predictors of the numbers of bats emerging. Sunset time was a significant predictor of the time of bat emergence. The timing of bat emergence was significantly later during the music festival, but there was no significant difference in the number of bats emerging. The bats left the priory up to 47 min later on festival nights than one would expect them to emerge in the absence of the festival. The results on the patterns of bat emergence were used to make recommendations regarding the timing of the music festival in the following year to minimize detrimental impacts on the bat colony.

The paper reports on a new family of spiders from the eastern Cape region in South Africa. Chummidae, new family, are small, three-clawed spiders, recognized by the abdomen with strong dorsal scutum, provided with a frontal field of stiff macrosetae in large sockets. Further apomorphies are the absence of a fovea, the reduction of posterior and median spinnerets, the absence of major ampullate gland spigots, and the presence of branched lateral tracheae and of a hair-basket around the cheliceral fang. The type genus Chumma, is represented by two species, both known from male and female – the type species C. inquieta sp. nov. and C. gastroperforata sp. nov. The males of the latter species are remarkable for having two pairs of abdominal pockets, which play a role during mating.

The Tete veld rat Aethomys ineptus Thomas & Wroughton, 1908 is a newly recognized, widely distributed species in southern Africa. Analysis of geographic variation among samples of A. ineptus from southern Africa across a more comprehensive geographical range than has previously been considered suggests that the species has a clinal pattern of variation in which overall cranial size was positively and significantly correlated with longitude. While the suggested clinal pattern of cranial size variation may be valid, the status of some operational taxonomic units, however, may require further refinement involving additional sampling as well as other systematic techniques, such as DNA analysis, cytogenetics and geometric morphometrics. These additional studies may have to include geographic information systems, step-wise multiple regression, and trend-surface analysis involving a wide range of environmental parameters to identify factors that may explain the nature and extent of the delineated pattern of geographic variation within A. ineptus from southern Africa. Consequently, the seven previously recognized subspecies recently assigned to A. ineptus in southern Africa (A. chrysophilus tzaneenensis Jameson, 1909; A. c. pretoriae Roberts, 1913; A. c. magalakuini Roberts, 1926; A. c. capricornis Roberts, 1926; A. c. tongensis Roberts, 1931; A. c. fouriei Roberts, 1946; and A. c. harei Roberts, 1946) are only provisionally synonymized.

The variable keratin appendages of the forestomach epithelium on both sides of the reticulo-omasal orifice, emerging in early summer and disappearing towards the winter in the moose Alces alces L. were studied. During the study period, droppings of the free-ranging moose were collected randomly for faecal particle size distribution analysis. The morphological distribution, size and abundance of keratin end caps was recorded in moose that had died of various causes, and it correlated with the particle size distribution observed in faeces. The fusiform keratin filter of the reticulo-omasal orifice seems to reflect the forage quality and use. It is suggested that this filter prevents the passage of non-fragmented foliage and twig fragments of excessive size during the period of rapid and non-selective intake of high-quality mature forage and it disappears before the short growing season of easily available succulent vegetation with a high nutrient content.

The reproductive pattern of the female Angolan free-tailed bat Mops condylurus (A. Smith, 1933) was investigated at two localities in Mpumalanga. Female Angolan free-tailed bats displayed a seasonal bimodally polyoestrous reproductive pattern. Parturition in December was followed closely by a post-partum oestrus. Ovarian follicle and endometrium cycles also closely followed the breeding season. The reproductive tract displayed complete dextral dominance with the right ovary and uterus being functionally dominant. Luteal activity was maintained throughout pregnancy and healthy corpora lutea were observed until late in pregnancy. Shedding of the zona pellucida occurred in the oviduct in the region of the ampulla–isthmus junction. The embryo entered the uterus as a morula, where it was transformed into a unilaminar, free-lying blastocyst. A unique feature of the pre-implantation uterus was the formation of a distinct decidua at the implantation site on the mesometrial side. Implantation in the Angolan free-tailed bat was centric and superficial. First attachment of the trophoblast to the endometrium was mesometrial. The formation of a pre-placental pad of trophoblast cells was also initiated on the mesometrial side, while orientation of the embryonic disc was anti-mesometrial. Placental development occurred in the abembryonic hemisphere. Proliferation of the trophoblast over the embryonic hemisphere was slow and a bilaminar arrangement of the trophoblast was not observed in this region until after the primitive streak stage.

This paper provides a comparison of the landbirds of all main zoogeographical regions, based on the most recent (Sibley–Monroe) listing and classification of the world's birds. This classification arranges 9416 landbird species (i.e. excluding seabirds) into 2002 genera, 140 families and 23 orders. On this basis, the Neotropical region holds 36% of all known landbird species and 45% of genera, the Afrotropical region holds 21% of species and 24% of genera, the Indomalayan region 18% of species and 22% of genera, the Australasian region 17% of species and 23% of genera, the Palaearctic region 10% of species and 14% of genera and the Nearctic region 8% of species and 15% of genera. These major continental regions thus show 4.6-fold variation in species numbers or 9.1-fold variation in species numbers per unit area. The region of Oceania, comprising many Pacific Islands, holds only 2% of the world's bird species and 4% of genera. About 92% of all bird species on the continental parts of the Neotropical, Afrotropical and Australasian regions are endemic to those regions, compared to 64% of the Indomalayan, 54% of the Nearctic and 46% of the Palaearctic species. The Oceania region has the smallest number of endemics, but these form 87% of all species occurring naturally in this region. About 91% of all landbird species breed in only one zoogeographic region, another 8% in two regions, with the remaining 1% in three to seven regions. Only four species breed in all seven regions. Similarities in the species composition of different regions were compared using Jaccard and Simpson indices. As expected, each region shares the greatest number of species with the closest other region and the fewest species with the most remote region. As in previous analyses, the Neotropical and Australasian regions emerged as having the most distinctive avifaunas. Regions that hold large numbers of landbird species also hold large numbers of genera and families, as well as high species-per-genus and species-per-family ratios. Comparable levels of diversity thus extend through all these taxonomic categories. This implies that, whatever factors have promoted particular levels of avian diversity in the different regions, they are of long standing. As found in previous studies, species-per-genus and species-per-family ratios are lower in island than in continental avifaunas. No relationship is apparent between the size of each zoogeographical region and the numbers of species, genera and families found there; rather those regions with tropical forest have many more bird taxa overall than those without.

There are now 91 records of individual Nathusius' pipistrelle Pipistrellus nathusii examined ‘in the hand’ in the U.K., and bat detector records from a further 36 localities in Britain and Ireland. These records are distributed throughout all months of the year with clear peaks in spring and autumn suggesting migratory movements into and out of the British Isles in autumn and spring, respectively. Three maternity colonies have been located and the species must now also be regarded as a resident breeding species in Britain and Ireland.

The analysis of life-history traits suggests that the age and size of female mammals at parturition will affect the birth size and survival of their offspring. We collected data from 252 mother–pup pairs of northern fur seals on Bering Island, Russia, during the 1994 and 1996 breeding seasons to determine the interrelationships among the mothers' age and mass, the sex of the pup, the parturition date, and the length and mass of the pup at birth. Among reproductive females of 19 years, mass increased with age. The mean mass of the oldest class of females (20–23 years) was lighter than the 16–19 year age group but not different from the mean mass of the 7- to 15-year-old females; a quadratic model of mass on age gave similar results, and indicated a maximum mass at age 19 years. We suggest that this may be an effect of a higher rate of survival among leaner animals. Female pups were lighter than males at birth. There was no evidence that the sex ratio of pups differed from 1:1 over the range of observed mothers' mass and age or parturition dates. Older mothers tended to give birth earlier than younger mothers, and heavier mothers earlier than lighter mothers. The relationship of the size of pups at birth and their mothers' age was adequately described with a quadratic model, which predicted a maximum size for mothers at age 12–13 years. The size of pups at birth and the size of their mothers was described with a logistic model which predicted that the size of pups increased for mothers up to 41 kg, with no further increase for heavier mothers. The total amount of variability in birth size explained by the combined models is < one-third of the total, which implies that other influences, such as the contribution of the fathers and individual variation, are also important. The relative maternal investment, measured as the ratio of pup mass to mothers' mass, ranged from 20% for younger and smaller females to 10% for older and heavier females. In addition, relative maternal investment was found to be higher than for other pinnipeds.

The relationship between wing morphology and ability to manoeuvre through an obstacle course was examined in five species of bats in the Neotropical family Phyllostomidae. These species represent a range of morphologies and feeding adaptations: Tonatia silvicola, a surface-gleaning insectivore; Artibeus jamaicensis and A. literatus, canopy frugivores; and Carollia perspicillata and C. castanea, understory frugivores. Bats of known morphology were presented with three different obstacle arrangements chosen in random order. In order to control for wing span, spacing between obstacles was scaled to the wing span of the individual (1×, 0.75×, 0.5× the wing span of the bat). Performance was measured as the number of times a bat hit an obstacle hard enough to deflect it. Morphological variables were analysed using principal components analysis and were summarized by the first two principal components (PC): PC 1, ‘size’, with which size-related variables were strongly, positively correlated; PC 2, ‘camber’, on which variables associated with maximum depth of wing camber weighted most heavily. Obstacle course performance was significantly correlated with both size and camber at all obstacle spacings: larger bats hit more obstacles, even though obstacle spacing was scaled to wing span, and bats with better ability to camber wings hit fewer obstacles. These results show that bat species that forage in more cluttered habitats are more likely to be small in size and/or have wings able to produce the high camber necessary to maintain lift at low flight speeds.

The Asiatic wild ass Equus hemionus, or khulan, once ranged across much of Central Asia, but is now globally threatened. The largest free-ranging populations are now restricted to a 250-km wide area (range 100–400 km) across the Gobi Desert region of southern Mongolia. Over the last 23 years the population has moved further north and east into its former range. Surveys conducted in the 1970s and 1980s estimated that the Mongolian population contained fewer than 15 000 animals and was declining as a result of human exploitation and livestock competition. Aerial surveys (one in autumn 1994, two in spring 1997) were flown as line transects over portions of the khulan's range in Mongolia and ground surveys (five in spring, summer, and autumn 1994–97) were conducted by vehicle and foot. Sample sizes and areas surveyed were larger than previous surveys, and our methods were often more systematic. Population size was estimated at 33 000–63 000 wild asses in Mongolia. Animal density ranged from 4.2±1.3 to 19.1±3.2 per 100 km2. Mean group size ranged from four to 35 animals in the south-western Gobi, four to seven animals in the southern Gobi, and three to 18 in the south-eastern Gobi. Our data suggest that Mongolia is the most important stronghold for the conservation of E. hemionus. Conservation management continues to be challenging because intensive studies on khulan biology and ecology are just beginning. As a free-market economy continues to emerge in Mongolia, pressure from resource extraction interests and nomadic livestock herders to remove the khulan's protected status, permit harvesting and halt population growth and expansion, also makes implementation of research and conservation management programs more imperative.

Temperature and respiratory gas (CO2 and O2) concentrations were measured in the foraging tunnels of burrows naturally inhabited by two species of southern African mole-rats, the Cape mole-rat Georhychus capensis and the Damaraland mole-rat Cryptomys damarensis. Both species are completely fossorial and inhabit closed burrow systems. Tunnels of G. capensis burrows had a mean diameter of 8.7 cm and a depth, measured to the roof of the tunnel, of 6.2 cm; those of C. damarensis had a mean diameter of 6.5 cm and depth of 40 cm. In both species, the mean concentration of CO2 was higher, and mean concentration of O2 lower, in burrows than in the surrounding soil or in ambient air. Mean and minimum values of O2 were 20.4% and 19.8%, respectively, in G. capensis and 20.4% and 19.9% in C. damarensis; mean and maximum values of CO2 were 0.4% and 1.2% in G. capensis and 0.4% and 6.0% in C. damarensis. Temperature varied between 18.5 and 24.2 °C in burrows of G. capensis by comparison with an ambient range of 16.9 to 26.8 °C; and from 19.6 to 29.3 °C in burrows of C. damarensis by comparison with an ambient range of 8.6 to 30.8 °C. Thus a burrowing habit seems to offer both species protection from extremes of temperature without entailing the cost of a grossly abnormal respiratory environment. From a review of the relevant literature, we conclude that average concentrations of CO2 and O2 in mammalian burrows often do not differ greatly from ambient values. However, more work is needed to determine the respiratory gas concentrations in the immediate vicinity of active, burrowing animals.