A central challenge to the study of animal aerodynamics has been the measurement of forces generated by flapping wings. Relative to wings of other birds, hummingbird wings are of particular interest in that the smaller species operate in more viscous regimes (5000<Re<10 000) for which substantial drag and reduced lift:drag coefficients might be expected. Lift and drag forces were measured on mounted hummingbird wings and wing models spinning in continuous tipwise revolution about the wing base. Lift coefficients tended to increase as wing models became more realistic (i.e. with sharpened leading edges and with substantial camber). Lift:drag ratios of real wings were substantially higher than those of wing models, suggesting morphological contributions of feathers to lift enhancement and drag reduction. At Re=5000, high values of the lift:drag ratio (8–16) at low angles of attack suggest that wings of hummingbirds are exceptionally good at producing lift.

Much of the narial anatomy of the enigmatic antelope Saiga tatarica has been described by previous workers. However, the anatomy of the nasal cavity and the causally associated osteological correlates of proboscis structure require closer attention, because these data are integral for both a more comprehensive understanding of saiga functional morphology and more robust reconstructions of proboscis structure in fossil taxa. Saiga and outgroup specimens were subjected to X-ray computed tomographic (CT) imaging, gross dissection and skeletonization. The nasal cavity of saiga is characterized by an enlarged nasal vestibule and basal conchal fold. Many structures (e.g. turbinates, lateral cartilages, mucosal folds, nasolacrimal duct) are retracted caudally to a small area in the caudodorsal part of the nasal cavity proper. The enlarged vestibule is associated laterally and ventrally with paired sacs. The nasal septum is largely membranous and contains a large patch of cavernous tissue that serves as a dynamic baffle modifying the flow of inspired air. Bones comprising the narial margin have modified attachment sites for buccinator group muscles and the reduced lateral cartilages. The premaxilla is greatly modified by the enlarged musculature associated with nasolabial fusion. Maintenance of the topological relationships of narial structures compared to bovid outgroups has resulted in a nasal cavity with much larger area for seromucous glands of the vestibule as well as narial musculature capable of controlling the aperture of the nasal cavity. Maxillolabial muscles and the lateralis nasi act together both to compress the nasal cavity and to control the dilation of the nostrils such that air flow through the cavity is highly modified relative to bovid outgroups. The lateral vestibular recess is an outpocketing of the nasal vestibule that produces supplementary seromucous secretions and seems to have no homologue among outgroups. The enlarged nasal vestibule, lateral vestibular recess, repositioned basal fold, and septal cavernous mass are regarded as a coordinated adaptation to dusty habitats, such that nasal air flow can be dynamically regulated allowing for collection of inspired particulates in the vestibule and thus cleansing of air destined for the lungs.

Aspects of the physiology of the arid-occurring otomyine rodents of southern Africa (Parotomys brantsii, Parotomys littledalei and Otomys unisulcatus) were examined, including: (1) renal function; (2) thermal physiology. Parotomys littledalei exhibited a maximum predicted urine-concentrating ability similar to other arid-occurring rodents. Maximum concentrating abilities were moderate for O. unisulcatus and low for P. brantsii. The extension of P. littledalei into the driest regions, in which it occurs allopatrically, suggests the importance of renal function in determining the distributional limits of these species. The thermophysiology of these otomyine species demonstrated a mixed ability to cope with aridity. Typical of arid-occurring diurnal rodents, all became hyperthermic at high ambient temperatures. However, none exhibited several additional thermophysiological characteristics common to arid-occurring rodents. Thus, resting metabolic rate was not lower than expected for body size, while lower critical temperatures were relatively low and the thermal neutral zone was broad. Nevertheless, the upper limit of thermoneutrality of P. littledalei was lower than that of P. brantsii or O. unisulcatus. The failure of P. littledalei to occur in hotter areas, in which P. brantsii occurs, seems to be related to its thermal physiology. Whereas renal function or thermophysiology may influence the arid extension of P. brantsii and P. littledalei, the high vulnerability of its refuges to fire damage may limit the distribution of O. unisulcatus. Reviewing the thermophysiology of the Otomyinae from more mesic habitats, we argue that their thermophysiology is more closely related to refuge type than aridity. Compared to the apparently specialized physiological mechanisms used by omnivorous or granivorous desert rodents, our study suggests behavioural adaptations such as refuge type and food selection are more important to the survival of these herbivorous desert rodents.

Many species exhibit skewed sex ratios at birth. Here we investigate the relationships between environmental and maternal variables (as surrogates for maternal condition) and foetal sex in African buffalo Syncerus caffer and elephant Loxodonta africana of the Kruger National Park, South Africa. Using logistic regression no significant effect was found of year, maternal lactational status, maternal age, rainfall or density on foetal sex ratio. Using a subset of our data, it was also concluded that maternal body condition did not affect foetal sex ratio in buffalo. Our analyses failed to support hypotheses predicting that mothers will skew the sex ratios of their offspring in relation to their body condition. In this study, buffalo and elephant produced offspring with a sex ratio close to parity. Our results are discussed in light of the implications for testing such hypotheses in analyses of population level.

Embryos of four ovoviviparous species of Australian onychophorans were examined to establish the process of their development, their reproductive cycle and to estimate the gestation period. The process of development is the same in all four species with, in the early embryo, segment halves separated by broad bands of extra-embryonic ectoderm and the embryos growing by the posterior addition of segments. Seven developmental stages (I–VII) can be clearly identified from external morphology. Analysis of the developmental stages of the embryos present in females collected throughout a calendar year has allowed estimation of the time taken for each developmental stage and demonstrated that there is an annual reproductive cycle with a total gestation period of c. 12 months. Stages I and VII are prolonged, the latter related to the late completion of midgut development. The simultaneous presence of stage I and stage VII embryos at certain times of year indicates an overlap of generations within the uteri, which is more extensive for some species than others. As the stage VII embryos of one cohort complete their development, enlarged oocytes are released from the ovaries to become fertilized as they pass the seminal receptacles and enter the uteri to start their embryonic development to form the next cohort. Cephalofovea clandestina, Phallocephale tallagandensis and Ruhbergia bifalcata have head structures in males, Euperipatoides rowelli does not. The presence of embryos at all times of year in the uteri of mature females has implications for mating and sperm storage that are discussed.

A distinctive new striped bandicoot of the genus Microperoryctes (M. aplini new species) is described and compared with Microperoryctes murina, another little-known bandicoot with which the new species has been confused in the past. The smallest of all living bandicoots, M. aplini is apparently restricted to high-elevation forests in the Arfak Mountains of the Vogelkop Peninsula of western New Guinea. The similarly small M. murina is known only from high-elevation forests on Gunung Sumuri, a peak in the Weyland Range of western New Guinea. The two species differ markedly in fur colour and ornamentation, pelage texture, external proportions, and quantitative and qualitative craniodental aspects, and may not be immediately related to one another.

Comparative studies require information on phylogenetic relationships, but complete species-level phylogenetic trees of large clades are difficult to produce. One solution is to combine algorithmically many small trees into a single, larger supertree. Here we present a virtually complete, species-level phylogeny of the marsupials (Mammalia: Metatheria), built by combining 158 phylogenetic estimates published since 1980, using matrix representation with parsimony. The supertree is well resolved overall (73.7%), although resolution varies across the tree, indicating variation both in the amount of phylogenetic information available for different taxa, and the degree of conflict among phylogenetic estimates. In particular, the supertree shows poor resolution within the American marsupial taxa, reflecting a relative lack of systematic effort compared to the Australasian taxa. There are also important differences in supertrees based on source phylogenies published before 1995 and those published more recently. The supertree can be viewed as a meta-analysis of marsupial phylogenetic studies, and should be useful as a framework for phylogenetically explicit comparative studies of marsupial evolution and ecology.

As a solitary species, giant pandas Ailuropoda melanoleuca have a chemical communication system that allows for avoidance with conspecifics throughout most of the year and facilitates breeding during their brief reproductive period. To date, most studies have focused on adults during the breeding season, but much remains to be learned about developmental and seasonal effects on chemical communication. Here, using odour discrimination methodology, we examine these contextual influences on chemosignalling in giant pandas for the first time. During the breeding season both adult and subadult giant pandas overtly discriminated conspecific sex using chemical cues in urine. Male urine was consistently investigated more than female urine by pandas across all age–sex categories. This preference for male urine was observed for both adult and subadult urine donors, indicating that adult levels of reproductive hormones are not necessary for the production of sex-specific urinary odour cues. By contrast, giant pandas did not overtly discriminate sex during the non-breeding season. This finding underscores that negative results for discrimination studies need to be interpreted with caution, as many contextual factors may influence overt expression of this ability. Evidence points to seasonal changes in motivation, rather than in the availability of sex-specific chemical cues in urine, as the causal factor for changing odour discrimination patterns. These and other findings are discussed in the light of the potential role that discrimination of odour plays in the regulation of competitive and reproductive interactions in nature.

Adult sex change and juvenile sexual differentiation in the protogynous epinepheline Cephalopholis boenak were demonstrated in captivity to be influenced by social factors. Adult sex change in C. boenak occurred in two directions, female to male and male to female. The presence or absence of a larger male plays an important role in adult female sex change; female(s) did not change sex in the presence of a larger male, but sex change occurred after the removal of the larger males in the same social groups. In male pairs, either the larger or the smaller male changed sex. Male to female sex change has not been reported previously in Cephalopholis, and only rarely in epinephelines. This is the first report of direct male sexual differentiation from juveniles (i.e. primary male development), through manipulating the number of juveniles, in a protogynous epinepheline. All isolated, single, juveniles differentiated directly as males, and male to female sex ratios did not differ significantly from 1:1 in all experimental social groups of two to four juveniles. Differentiating males grew significantly faster than differentiating females and undifferentiated juveniles during the 57-week experimental period. The role of growth rate in sex determination in C. boenak is not known but clearly plays a part in juvenile sexual differentiation and merits further investigation. Social factors influencing bi-directional adult sex change and juvenile sexual differentiation are a reflection of plasticity of sexual expression in C. boenak, in particular, and the Serranidae in general.

Mimicry is a widely documented phenomenon in coral reef fishes, but the underlying relationships between mimics and models are poorly understood. Juveniles of the surgeonfish Acanthurus pyroferus mimic the coloration of different pygmy angelfish Centropyge spp. at different locations throughout the geographic range of the surgeonfish, while adopting a common species-specific coloration as adults. This study examines the ecological and behavioural relationships between A. pyroferus and one of its models, Centropyge vroliki, in Papua New Guinea. Surgeonfish underwent a transition from the juvenile (mimetic) coloration to the adult (non-mimetic) coloration when they reached the maximum size of the angelfish. As typical of mimic–model relationships, mimic surgeonfish were always less abundant than their model. Spatial variation in the abundance of mimics was correlated with models, while the abundance of adults was not. We show that juvenile surgeonfish gain a foraging advantage by mimicking the angelfish. Mimic surgeonfish were always found within 1–2 m of a similar-sized individual of C. vroliki with which they spent c. 10% of their time in close association. When in association with angelfish, juvenile surgeonfish exhibited an increase of c. 10% in the amount of time spent feeding compared to when they were alone. This foraging benefit seems to be explained by reduced aggression by the territorial damselfish Plectroglyphidon lacrymatus, which dominates the reef crest habitat. While adult A. pyroferus and all other surgeonfish were aggressively displaced from damselfish territories, mimic surgeonfish and their models were attacked less frequently and were not always displaced. Stomach contents analysis showed that the diet of C. vroliki differed substantially from P. lacrymatus, while that of A. pyroferus was more similar to the damselfish. We hypothesize that mimics deceive damselfish as to their diet in order to gain access to food supplies in defended areas.

Nipponoclava gigantea is restricted to the tropical Western Pacific waters of southern Japan at depths of between 5 and 70 m and lives in coarse sand and gravel. In many respects, N. gigantea appears to be similar to other endobenthic penicillids in that there is a true shell, the margins of which, via the intermediary of a narrow ‘saddle’, are united with an adventitious tube. This has a large watering pot anteriorly and pleated ruffles, representing either growth or repair events, posteriorly. Nipponoclava gigantea differs from other penicillids, however, in that its true shell is large, i.e. ∼20 mm high as compared to 2–3 mm, and the angle of valve splay is ∼80° as opposed to 180°. Internally the shell has a wide pallial line and shallow pallial sinus and there are anterior adductor muscle scars. There are possibly posterior adductor muscle scars also. These are not present in the adult animal, however, although the anterior adductor is. Also present are paired anterior and (possibly) posterior pedal retractor muscles, the latter having a nervous union with the visceral ganglia to create simple stretch or tonus proprioreceptors. It is argued that N. gigantea provides clues about how the tube-dwelling lifestyle has evolved in the various taxa of the Penicillidae. It is suggested that through a process of heterochrony, in which the animal produces its adventitious tube at progressively earlier and earlier developmental stages, the true shell has become actually and relatively smaller and smaller and, hence, more splayed in more derived penicillids. Nipponoclava gigantea is thus an important link between an hypothetical lyonsiid ancestor and the more specialized penicillids and helps us to understand the adaptive radiation of this group of tube-dwelling bivalves.

The spatial organization of one clan of spotted hyenas Crocuta crocuta in the centre of the Etosha National Park, Namibia, is described during the dry and the wet seasons. The clan comprised 11 adults and sub-adults and occupied a territory of 160 km2 in the dry season and 320 km2 in the wet season. The dry season territory contained a low density (one animal/km2) of resident herbivores, such as gemsbok Oryx gazella, kudu Tragelaphus strepsiceros, giraffe Giraffa camelopardis, steenbok Raphicerus campestris and ostrich Struthio camelus, and a higher density of migratory species (12 animals/km2), principally springbok Antidorcas marsupialis, zebra Equus burchelli and wildebeest Connochaetes taurinus. These migratory species were the main prey of clan members. At the start of the wet season, the migratory herbivores migrated to the north-west, resulting in a considerable decline in the density of prey in the area used by clan members during the dry season. In response to this decline in prey, clan members followed the migratory herds and shifted the focus of their activities to an area grazed by migratory herbivores during the wet season. There existed a strong spatial relationship between the hyena density and the migratory prey density in both the dry and wet season. We postulate that the considerable enlargement of the clan territory in the wet season is a response to both the migratory movements of prey and an increase in the dispersion of prey during the wet season.

Nesting hawksbill turtles Eretmochelys imbricata and their eggs and young were studied on Cousine Island, Seychelles from 1995 to 1999. Numbers of females known to nest (tagged individuals) on Cousine was 146. An average of 75% of emerging adults were intercepted, and this rose to 81.6% in 1999. Mean carapace lengths and widths of adult females were 81.2×60.8 cm, and included some of the longest carapace lengths known for hawksbill turtles. There were poor correlations between adult weight and carapace length or width, but better correlations between adult weight and the product of carapace length and width. Width to length carapace proportions averaged 0.75. The mean weight of nesting females before their first recorded nesting on Cousine was 65.6 kg (51.2–83.0 kg). These were the first recorded weights for nesting hawksbills from the Seychelles. Individuals lose 8.5–15% of their body weight after laying three to five clutches. Mean clutch size was 176.7, and clutch sizes for any season showed great variation for some individuals. Clutch sizes tended to decrease after the main nesting month for the population but not necessarily for the individual. The largest clutch size of 264 is the largest recorded for hawksbill turtles worldwide. Turtles with longer carapaces tended to lay larger clutches than those with shorter carapaces, and heavier turtles generally laid heavier clutches than did lighter turtles. Eggs were usually (85.6%) somewhat elongate, measuring on average 36.3×35.5 mm, and weights averaged 25.5 g. Very few ‘pea’ eggs were found. The average hatchling weight was 14.5 g and the average carapace length and width were 39.2 and 30.1 mm, giving a mean ratio of carapace length to width of 0.77.

Predation by wolves Canis lupus on roe deer Capreolus capreolus was studied by scat analysis in five areas of the Province of Arezzo, north-eastern Tuscany, Italy. In the intensive study area (ISA) roe deer represented 19.1% of mean per cent volume (MPV) while in the other areas its use ranged between 10.9% and 53.4% of MPV. A low degree of variation was found in both annual and seasonal use of roe deer, although seasonal differences were more marked in those areas where roe deer use was lower. In ISA, roe deer was negatively selected among species: the Ivlev's electivity index ranged between −0.41 and −0.89. Analysing intra-specific selection, in ISA <1-year-old individuals were preferred by wolves, mainly during the fawns' first months of life. This trend was confirmed in the other areas, where fawns represented more than 50% of the relative number of roe deer prey. In ISA, the use of roe deer (as MPV) was not correlated with its density or with that of the main prey, wild boar, and in almost all the other areas no relation between use and density of roe deer was founded. However, in the one area where both roe deer density and use were the highest, these two variables seemed to exhibit a comparable trend. The use of roe deer was negatively correlated with the percentage of forest cover among all the study areas.

Variation in size-assortative pairing was studied in relation to sexual selection on male body size in the amphipod Gammarus pulex, using both transverse and longitudinal surveys of natural populations. In addition, the influence of male–male competition on size-assortative pairing was tested in the laboratory. In both surveys, the intensity of sexual selection was positive and significant for male body size but not for females, and size-assortative pairing was positive and significant. The magnitude of size-assortative pairing, however, varied significantly between populations. The magnitude of size-assortative pairing was positively correlated with the intensity of sexual selection on male body size in both surveys. In the laboratory, the strength of size-assortative pairing decreased with decreasing male:female ratio. Taken together our results point to a role of interference competition between males in promoting patterns of size-assortative pairing in G. pulex, although alternative explanations (i.e. female resistance) cannot be excluded.

The ‘diaphragm’ of Xenopus laevis has close anatomical relations to the lower end of the oesophagus. In mammals, the crural diaphragm acts as a pinch valve at the gastro-oesophageal junction and is an important component of the gastro-oesophageal reflux barrier. The present study analysed the effect of amphibian ‘diaphragm’ contraction on oesophageal pressure using a superfused in situ oesophago-diaphragmatic preparation of large female Xenopus. Three-dimensional reconstruction of the oesophageal pressure profile was performed using four-port oesophageal infusion manometry. Bilateral electrical stimulation of the nerves supplying the ‘diaphragm’ of Xenopus increased the pressure volume vector of 5 mm of oesophagus (centred around the insertions of the diaphragm) from 20.4±16 to 553.6±232 mm.mmHg2 (mean±sd). This was a statistically significant increase and statistically significantly higher than that evoked by electrical stimulation of both vagi (28.1±30.7 mm.mmHg2). The amphibian ‘diaphragm’ seems to be functionally similar to the mammalian crural diaphragm. By analogy, we suggest that the original role of the diaphragm was not respiratory but gastrointestinal.

The skin of the emu Dromaius novaehollandiae is described in its entirety by light microscopy for the first time, therefore providing a basis for comparison with the integument of volant birds. The epidermal and dermal layers have a similar arrangement to other avian species. The concentration of lipid spheres in the basal layer of the epidermis is unique to emus. Although no differences in collagen density with depth of the dermis were observed, collagen bundle diameter is significantly smaller in the stratum superficiale compared to the stratum compactum (P=0.001). The extensive stratum laxum, comprised predominantly of adipose tissue, may be an evolutionary adaptation to support the large feathers and also enhance insulation against ambient subzero temperatures. Males have a significantly thicker cellular epidermis (P=0.024) and stratum compactum (P=0.04) than females. In contrast, females have denser collagen (P=0.04) within the stratum superficiale and stratum compactum.

Nesting hawksbill turtles Eretmochelys imbricata were studied on Cousine Island, Seychelles from 1995 to 1999. Mean overall incubation time averaged 58.1 days. Clutch size did not influence incubation periods. Following hatching, hatchlings took an average of 3.8 days to leave the nest. Mean hatchling emergence time was 60.4 days. Hatching success averaged 64.3% for the four seasons. During 1996–97 there were heavy, unexplained egg losses due to crab predation. Excluding that season, average hatching success was 70.9%. Overall hatchling success was 61.1%, translocated nests having a better success (69.3%) than natural nests (57.7%). Overall egg and hatchling losses were 35.8% and 3.1%, respectively, most egg losses being attributed to ghost crabs Ocypode cordimana (16.3% overall) and unknown causes (17.0% overall), which were mainly non-viable eggs. The high level of nest manipulation was believed to be justified by increasing the number of hatchlings entering the sea.

Circulating testosterone levels were measured at 195, 318 and 437 days of age in 76 male garter snakes (Thamnophis sirtalis) born to 26 wild-caught females. Testosterone levels increased significantly with increasing mass at all three ages and differed significantly among families at 318 and 437 days. ‘Full-sib’ estimates of heritability were near one at 318 and 437 days but these estimates may be inflated by maternal effects, dominance and epistasis. Evidence of heritable variation in circulating testosterone levels suggests that the evolution of traits affected by this hormone (reproductive, territorial and agonistic behaviour; body size and shape) might be mediated through changes in this hormonal control mechanism.

Sexual dimorphism is a common phenomenon among animals. The usual cause cited for sexual dimorphism in animals is sexual selection acting through female choice or male–male combat. Natural selection acting to reduce resource competition between the sexes, however, is an important alternative evolutionary scenario, but this possibility has received little empirical study. Here this issue is addressed by examining the relationships among body size, head shape and the functional aspects of diet in the adult male and female cottonmouth snake Agkistrodon piscivorus. In this species, males are larger in overall body and head size. Whereas an analysis of gross head measurements (simple linear head dimensions) shows little dimorphism in head shape, a more detailed analysis of head shape (using digital images of the snakes' heads) revealed some subtle, yet functionally significant, differences in head shape between adult male and female cottonmouths. Specifically, male cottonmouths have longer quadrate bones, and have greater lateral surface areas than females. Male cottonmouths also consumed relatively taller prey (prey size relative to snake body size) than conspecific females, and the sexes consumed significantly different proportions of prey. Because the size of the quadrate bone is a strong determinant of maximum gape in snakes, we suggest that the observed shape differences may reflect functional differences in maximum gape between similarly sized male and female cottonmouths. In turn, such differences in maximum gape width may explain why males consume taller prey than similarly sized females.