Few have served the Zoological Society of London longer or more faithfully than Alexander Cave who died on 17 May at the great age of 100.

As a Fellow, his record is outstanding: Member of Council, Vice President, Silver Medallist of the Society, Honorary Research Associate; and withal, an unending stream of original papers on various aspects of comparative anatomy. Unending? Well, almost, for he was still being published in his 95th year. Nor should be forgotten the great volume of work that he did in refereeing papers for the Journal of Zoology – an onerous and time-consuming task that he dealt with meticulously.

In Mediterranean environments browse is expected to be a major component of the diet of red deer Cervus elaphus during the summer when most grasses senesce. In this study the hypothesis is tested that, if browse is an essential food resource at this time, the sex with the larger body size may have a greater physical ability to reach the tree canopy and include a higher proportion of browse in its diet. The organic matter digestibility (OMD) and composition of the diet of a population of red deer males and females were measured between July and September, in 1996 and 1997, in an area of Portugal with a Mediterranean type of climate, using the n-alkane technique. Direct observations of animals browsing on tree canopies and shrubs were made also during the same period. The OMD of the diet was significantly different between sexes and between years and was higher in males than females both in 1996 (means ± sem for males and females, 0.65 and 0.58 ± 0.026, respectively) and 1997 (means for males and females, 0.59 and 0.48 ± 0.045, respectively). The proportion of cork oak Quercus suber in the diet was also significantly different between sexes and between years, with males having a higher proportion of cork oak in their diets in 1996 (males and females, 0.30 and 0.16 ± 0.037, respectively) and 1997 (males and females, 0.41 and 0.23 ± 0.064, respectively). A significantly lower proportion of the herbage layer was found in the diet of males in 1996 (males and females, 0.46 and 0.64 ± 0.048) but not in 1997. The number of observations of males browsing on tree canopies was significantly higher than females. It is concluded that there are differences between sexes in the composition and quality of the diet and that a larger body size may confer advantages in situations where browse is an essential food resource.

We describe diurnal resting dens (setts) used by Eurasian badgers Meles meles L. in Coto del Rey, Doñana National Park, south-western Spain, and analyse the factors that determine sett location and selection in relation to territory outlines, soil type and vegetation structure. Setts were located by tracking radio-tagged badgers daily. Badger setts were mainly underground burrow systems with, on average, 2.6 badger entrances. Frequently, badgers constructed setts by enlarging existing rabbit warrens. Setts were located almost everywhere, but badgers preferred easily dug, well-drained soils with good vegetation cover within foraging habitats. A logistic regression model showed that badgers select sites with high surrounding shrub density, large shrubs covering the burrow and close to the centre of the territory (in Mediterranean scrubland habitat). The small size of badger setts in Coto del Rey is common to other low density areas, and probably relates to the extent of use (i.e. age of the burrows and number of badgers using them) of the burrow system. The number of setts per badger territory is related positively to territory size. The similarity of sett densities between this low density population and high density populations, points to a minimum number of setts being required to maximize territory use, and to offer easily-available protection. The preservation and improvement of Mediterranean scrubland vegetation, as well as of large and old shrubs, are important for the future conservation of the badger in Mediterranean landscapes. In these areas, and given the impossibility of distinguishing main setts, the estimation of absolute densities in population surveys using main sett numbers should be avoided in favour of relative indices of abundance.

Discontinuous individual variation among members of the same sex, in the same life stage, within a population, is an important phenomenon for our understanding of more fundamental issues in evolutionary biology. One set of morphological characters is conspicuously rare in the literature on intrasexual polymorphisms: genitalia. In this paper we report the first case of female genital polymorphism, the first not only for spiders but apparently for animals in general. We argue that the overwhelming use of genitalia in species identification practiced by many invertebrate taxonomists presents a strong bias against discovering polymorphisms in genitalia. Further methodological and practical aspects of invertebrate taxonomy are discussed that, taken together, make the discovery of genital polymorphisms unlikely, no matter whether or not the phenomenon is rare.

Piscivorous birds consume diets which are rich in highly-polyunsaturated n-3 fatty acids; these play vital roles in embryonic development but are very susceptible to oxidative damage. The effects of such diets on the fatty acid composition and antioxidant content of the yolk were investigated in the northern gannet Morus bassanus, the great skua Catharacta skua, the American white pelican Pelecanus erythrorhynchos and the double-crested cormorant Phalacrocorax auritus. The phospholipid fraction of the eggs of these four species contained high proportions of the n-3 fatty acid, docosahexaenoic acid, which formed 7.5–11.3% (w/w) of the fatty acids of this fraction. The presence of eicosapentaenoic and docosapentaenoic acids also contributed to the total n-3 content of the phospholipid. The n-6 polyunsaturate, arachidonic acid, formed between 8% and 19% (w/w) of the phospholipid fatty acids. For the pelican and cormorant, this is consistent with the consumption of freshwater fishes in which arachidonic acid may be a significant acyl constituent. This finding is, however, more difficult to explain for the gannet and skua which largely consume marine fish with a low arachidonic acid content. The yolks of all four species contained relatively high concentrations of vitamin E (90.2–302.3 μg g−1 wet yolk) which was mainly present as α-tocopherol. The eggs of the pelican and cormorant were especially enriched in carotenoids (150.9 and 115.7 μg g−1 wet yolk, respectively).

Studies of the grey seal Halichoerus grypus mating system, using focal animal sampling constrained by study site location and size, limit the measurement of variation in male mating tactics and success. Using this method, the mating tactics of grey seal males have been classified as either ‘tenured’ or ‘transient’. Preliminary evidence is presented of wider variation in male mating tactics and success using mobile focal-animal techniques that are not constrained by site, and of the effects of age on behaviour when accounting for body mass. The study was conducted during the breeding seasons of 1997 and 1998 at Sable Island, Nova Scotia. Six young (11–12 years) and five old (20–25 years) males were captured at the beginning of the breeding season, weighed and fitted with a VHF transmitter and time-depth recorder. Each male was located daily, and a count of late-lactating females within a 10 m radius recorded as a measure of mating potential. Female guarding and observed copulatory attempts were used as a measure of mating success. Mean initial body mass of young males and old males was similar, 291 ± 13 kg and 298 ± 14 kg, respectively. All males changed location during the study period. Young males changed sites more often and moved shorter distances between sites than old males. Estimated mating success varied from probably no matings to a minimum of seven; there were no differences between young and old males. Maximum duration of stay at a site and the number of late-lactating females at a site significantly influenced mating success. All the males except for two young males departed on diving excursions during the breeding season and engaged in deep diving behaviour. Diving behaviour did not differ between the age classes. Our results indicate that the current classification of mating tactics in grey seal males is an oversimplification. Preliminary data show that age may account for some of the variation in male behaviour on land and mating success, but not for behaviour at sea.

Bridled nailtail wallabies Onychogalea fraenata are endangered, medium-sized, nocturnal macropodids that persist at only one location in central Queensland, Australia. Characteristics of juvenile development, shelter use, anti-predator behaviour and maternal care were investigated in the wild using trapping, radio-tracking and spotlighting observations. Timing of developmental stages was identical to the pattern previously found in captivity, except for age at weaning which was much earlier in the wild. After young had left the pouch permanently at 17 weeks of age and weighing c. 800 g, they always spent the day concealed in dense cover, generally >200 m from their mothers. Juveniles were also alone in >50% of observations at night, and stayed closer to cover than did adult females. Young became independent of their mothers 7–8 weeks after permanent exit from the pouch and weighing c. 1800 g. Females with dependent juveniles changed their behaviour in ways likely to reduce predation on young. They reduced their home ranges, stayed closer to cover and became more wary than other females. Juveniles differed from adult females in their habitat use, anti-predator behaviour and shelter site preferences. Juveniles were more likely than adults to respond to threats by standing still or lying flat on the ground, whether or not they were in concealing cover. Juveniles used a wider range of smaller shelters than adults, and were less likely to use solid shelters such as hollow logs during the day. Because bridled nailtail wallabies have a ‘hider’ strategy of maternal care and the young rely on crypsis, successful breeding in the wild requires dense vegetation cover.

Most species of birds of paradise (Paradisaeidae) are endemic in the rainforests of New Guinea. There are also a few species in the northern Moluccas and in northern Australia. Bowerbirds (Ptilonorhynchidae) are centred in New Guinea but are more widespread in Australia. The two families have often been regarded as sister groups, but in recent studies Paradisaeidae appear as sister to Corvidae, a worldwide family which is notably depauperate in New Guinea and Australia. This indicates vicariance of a worldwide ancestor, rather than invasion of New Guinea. Other families in the superfamily Corvoidea include Ptilonorhynchidae (basal), Campephagidae (Africa to New Guinea and Fiji), and Cracticidae (Australia to New Guinea). Biodiversity levels in Paradisaeidae and Ptilonorhynchidae were assessed from literature records by counting numbers of species in grid cells 1° latitude by 1° longitude. Birds of paradise are equally diverse in the Mendi square and the Mount Hagen–Wahgi Valley–Jimi Valley square. Bowerbirds are most diverse in the Mount Hagen–Wahgi Valley–Jimi Valley square. This area lies on one of the main tectonic boundaries in New Guinea, the former margin of the Australian craton, and is geologically distinctive in having several diverse accreted terranes juxtaposed there, including an ophiolite complex. It also includes the western slopes of Mount Wilhelm, one of the highest mountains in the New Guinea orogen. Paradisaeidae have secondary centres of diversity in the southern part of the New Guinea orogen (south of the former craton margin), while Ptilonorhynchidae have secondary centres north of the craton margin on the accreted terranes, and also in eastern Australia. Within New Guinea the two distributions correlate closely with the geological interpretation of the orogen as comprising southern (craton) and northern (accreted terrane) components. Within Australia, Paradisaeidae have two species per degree square in northern Cape York Peninsula (on the old Australian craton), and only one elsewhere in eastern Australia, whereas Ptilonorhynchidae have a clear Australian massing further south in Queensland, on the accreted terranes of the Tasman orogen.

Repeated measurements of captive-born Gaboon vipers Bitis gabonica from an inbred stock examined the degree to which an animal's size and shape are affected by food intake. We also used the level of asymmetry in dorsal coloration as an index for possible genetic (inbreeding?) effects. Both of these factors, and the interaction between them, affected phenotypes of the young snakes. Snakes raised with abundant food differed from their less well-fed siblings not only in size, but also in body mass relative to snout–vent length, head length relative to snout–vent length, head width relative to head length, and fang length relative to jaw length. Hence, our data show that body proportions (including the feeding apparatus) can be influenced by the environment after birth.

Primate quadrupeds are said to use relatively large limb excursions for mammals of their body size. Until recently, this claim was based on a comparison of hindlimb excursion data derived from small samples of primates and non-primates. Using video recordings collected at zoos and primate research centres, the present study documents this contrast on much wider samples of quadrupedal mammals. The results indicate that while on average hindlimb excursion is relatively larger in quadrupedal primates, this contrast is somewhat less dramatic than first reports suggested. Comparisons between the data reported here and previously collected forelimb excursion data reveal a surprising asymmetry between the fore- and hind excursions for most mammalian species. Most commonly, forelimb excursion exceeds that of the hindlimb. We suggest that this is related to a complementary asymmetry in limb length (forelimbs shorter than hind) for the purpose of achieving equal step lengths for both pairs of limbs. Relatively large hindlimb excursions in primates have been related to a mechanism that reduces stresses on the forelimbs and then recovers mechanical energy during gait. We suggest that large excursions of both the fore- and hindlimbs are linked to other alterations in gait parameters, such as step length, contact time, and limb compliance, that have been adopted in quadrupedal primates to facilitate locomotion along slender arboreal substrates.

The functioning of female genitalia in many arthropods, especially the site and mode of sperm storage, may have a strong impact on the pattern of sperm usage, leading to differential male fertilization success and even varying offspring sex ratios. The female genital morphology of the linyphiid spider Pityohyphantes phrygianus was investigated, as this species was shown to possess an intriguing mechanism of sex ratio determination, possibly related to genital morphology. By means of SEM and serial semi-thin sectioning, two distinctly different spermathecae plus an additional sperm sac, equipped with several valves, were found on each side of the genital apparatus. Sperm is stored under different conditions, as one of the spermathecae exhibits large gland cells of different types that discharge their products into the spermatheca through long secretory ducts, whereas the other spermatheca is surrounded by a simpler type of gland cell. Furthermore, there are folds instead of closed ducts leading to the spermathecae, and from the spermathecae to the oviduct. The latter fertilization-fold runs inside the copulatory chamber and enters the oviduct from the outside. The mechanisms for sex ratio bias in spiders are discussed. The occurrence of two types of spermatheca of which one is twisted, provides good evidence for the involvement of female genital morphology in sex ratio control of P. phrygianus offspring. Data are compiled on species for which information is available on sex ratio and female genital morphology. We put forward a hypothesis that in species with a skewed sex ratio, there is interaction between female genital morphology and behaviour, i.e. change of body position before, during or immediately after sperm transfer.

Both intensity and type of habitat management in grasslands and heathlands affect spider communities. With high intensity management, spider communities often lack diversity and are dominated by a few r-selected species affiliated with bare ground. Low intensity management produces more complex communities introducing more niches for aerial web spinners and climbing spiders. The preferred management will be site-dependent and may not be appropriate for all spiders in all situations, particularly for some rare or threatened species. Providing natural cover is recommended when using extreme forms of management or intensive grazing (particularly by sheep). In extreme cases, or where trampling is heavy, the litter layer should be conserved. We advocate research and survey before and after major management implementation. Habitat management for spiders should not be considered alone, but integrated into a holistic plan. Management for spiders may conflict with rare plant conservation and small reserves should examine the viability of providing two contrasting regimes.

The taxonomic relationship between two toothed South African river crabs, Potamonautes warreni and P. unispinus, is unclear. The problem stems from the widespread variation in carapace dentition patterns amongst P. warreni individuals over its biogeographic range, where single toothed individuals may appear similar in carapace morphology to P. unispinus. Ten populations of P. warreni and 18 populations of P. unispinus were collected and the morphometric and genetic differentiation between the two taxa quantified. Patterns of morphometric and genetic variation were examined using multivariate statistics and protein gel electrophoresis, respectively. Principal component analyses of carapace characters showed that the two species are morphologically indistinguishable. However, discriminate functions analyses and additional statistical results corroborate the morphological distinction between the two taxa. Allozyme electrophoresis of 17 protein coding loci, indicated a close genetic similarity between the two species (I = 0.92). A fixed allelic difference at one locus (LT-2) and extensive genetic variability at another locus (PGM-1) indicate that two gene pools are present and that the two taxa are genetically isolated. Intraspecific genetic I values for both species were >0.97 and indicated no apparent genetic structuring on a micro or macro-geographic scale. The variation in carapace dentition among P. warreni populations possesses no genetic basis and may possibly be the product of ecogenesis. The value of dentition patterns in the systematics of river crabs is discussed. Dentition patterns among river crab species appear to be conserved and reliable as species specific diagnostic markers, but should ideally be used in combination with other morphological data sets and genetic evidence.

During reproduction, dung beetles of the genus Onthophagus typically bury portions of dung removed from the dung pad. Even though females are capable of successfully constructing and provisioning nests when unaccompanied by males, extensive levels of co-operation between sexes have been reported. In some species, males show a clear dimorphism in body plan and only large horned males are known to assist females. In O. taurus this form of paternal investment is facultative and depends on social context (namely the presence of other males). We studied relationships between body size and paternal investment in five Onthophagus species (i.e. O. coenobita, O. fracticornis, O. illyricus, O. taurus and O. vacca). Two types of tests were established. In test A, a large and a small male were placed with a single, medium-size female; in test B, a large and a small female were placed with a single large male. In controls one male was placed with one female. Individuals were maintained for 10 days. Then the enclosures were opened to assess the position of each beetle. When two mates were found in close vicinity or in the same tunnel provisioning dung, a co-operative behaviour was assumed. In competitive tests, small males and small females never co-operated with their potential mates whilst large and medium-size individuals often co-operated with mates inside the same tunnels. No differences in frequencies of co-operation were found between large and medium-size females. Small females and small males scarcely co-operated with large or medium-size mates in controls. These results suggest that large individuals outcompete the small ones. They also suggest that paternal investment depends on body size of both mates. Large males of all species assist medium-size and large females, whereas small males do not. However, large males do not invest much parental effort on small females that, in fact, are poorly assisted. We assume that large individuals of both sexes are reproductively superior to small ones. The propensity of both small females and small males to excavate was lower than that of larger individuals; accordingly, digging capacity could be a proximate factor explaining facultative paternal investment in these species.

The foraging behaviour of Brünnich's guillemots Uria lomvia was studied in a high-arctic fjord system (Kongsfjorden) in western Spitsbergen. The physical oceanographic characteristics of the water change from the head of the fjord and westward into the Greenland Sea, and are reflected in the vertical profiles of water temperatures in different parts of the area. Nine chick-rearing Brünnich's guillemots were instrumented with temperature-depth recorders, which generated vertical temperature profiles of the dives. These were compared to synoptic measurements of the water temperature characteristics of the region. This method was used to locate the foraging areas of the Brünnich's guillemots and to study the foraging site fidelity of individual birds. The results showed that only three of the nine birds foraged outside Kongsfjorden during the study period, and only 26 of the 186 dive bouts (14%) were conducted outside the fjord, 48–58 km from the colony. Most dives were probably made only a few kilometres from the colony. The data indicate that the birds showed strong fidelity to foraging areas at spatial scales of 1–20 km. However, the birds sometimes moved between feeding areas characterized by different vertical temperature profiles. The guillemots made 2229 dives during the study period and spent c. 10% of their time under water. The diving depth averaged 45 m, and the dive duration averaged 97 s. The deepest dive recorded was 136 m and lasted 196 s. We did not find any diel rhythm in the diving depths of the Brünnich's guillemots. Also, we found no diel pattern in diving frequency. These findings contradict the predominance of night-time diving observed in studies of guillemots undertaken further south.