A meta-analysis of the duetting patterns of insect taxa that signal by bioluminescence, substrate vibration, or sound was used to test the prediction that call length by the initiating male influences the latency of reply of the female. There was a significant and positive relationship between these two measures. Although tests on the entire dataset did not consider phylogeny, when all members of one sub-family of bushcricket, the Phaneropterinae, were isolated there was a significant and positive relationship between call length and reply latency. Two explanations are suggested. First, the female must wait for the conclusion of a long and potentially variable message; there is uncertainty as to when the call has finished before she can reply. Second, is the requirement of the female to process information; a longer male call, with more information requires more processing than a brief call. The authors suggest that long reply latencies may be vulnerable to intrusion by competing mates. One defensive tactic of potential female partners is to insert a trigger pulse that indicates to the female when the long and complex call has concluded. This behaviour may be considered as acoustic mate guarding and one example is given of this behaviour.

In lizards and Sphenodon, often the fourth toes of individuals with intact tails have more subdigital lamellae on the right than on the left side, and the opposite situation frequently occurs in individuals with injured tails. The difference between intact and injured individuals in morphological directional asymmetry was statistically significant (P<0.05) in 11.4% among 193 species from various lizard families. Lizard families varied in extent and direction of association, but no phylogenetic constraints were detected within genera. Statistical significance was greater in samples from homogenous geographic origin than from heterogeneous ones. Among gekkonid species, the difference was stronger in those with cursorial (terrestrial) habits, than in those with scansorial (rupestral or arboreal) habits. In Scincidae, loss seems more often lethal in left-footed than in right-footed individuals. Statistically significant associations between morphological left-side dominance and tail injury exist also in three independent lineages with reduced limbs (Anguidae, Scincidae and Teidae). Hence such association is probably not a result of limb function. Rather, left-side dominance seems to be the symptom of an unknown, perhaps organism-wide, detrimental trait. Polymorphism in morphological dominance existed in all species, suggesting advantages and disadvantages in different situations to both phenotypes. We propose the hypothesis that an inversion of side dominance may occur in a single trait without systematic inversion of side dominance in all traits of the body. Inversion in a single trait causes incompatibility in multiple-trait functions. Such a mechanism, rather than cultural conventions, could increase accident proneness also in left-handed Homo sapiens, and could explain increased proneness to accident and warfare mortality in left-handed men, beyond the possible involvement of cultural factors.

Sexual differentiation was studied at the histological level using a mixture of 30 families of sea bass Dicentrarchus labrax. Most of the fish (93%) differentiated into males as usually observed in farmed populations. All testes were differentiated when the males reached 12 cm and no more undifferentiated fish were found from 419 days post-fertilization (p.f.). In 28% of the males, among the biggest, sexual differentiation had already begun at 168 days p.f. (8.3–9.5 cm) and these fish started spermatogenesis in their first year of life. The other males differentiated later and remained immature at the end of their first year of life. Ovaries could be identified at the histological level from the age of 168 days p.f. (7.9–9.0 cm) and the females became significantly longer than the males from the age of 191 days p.f., i.e. during the process of ovarian differentiation. In the studied group, 62% of the males developed intratesticular oocytes. Such intersexuality had no consequence on growth rate. Intratesticular oocytes were also recorded in testes of wild males originating from Atlantic (Britain and Gulf of Gascogne) and West Mediterranean showing that juvenile intersexuality is not restricted to farmed populations but is a widespread phenomenon in sea bass.

In many social species, relationships within groups seem to be non-random but related to variables such as rank, kinship or sexual attractiveness. The endangered African wild dog Lycaon pictus is a social carnivore that lives in large, stable packs, and intra-pack associations might be expected to display similar patterns. We investigated patterns of coalition formation (support during dominance interactions, and partnership interactions) and resting associations between members of a captive pack of 19 wild dogs. The social organization of the captive pack was similar to that of free-ranging packs in many respects. Polyadic (group) incidents of coalition support were also observed in a free-ranging pack. Patterns of coalition formation in the captive pack were related to rank. Most aggressive interactions involved high-ranking individuals (particularly the alpha, beta and third-ranking males) and coalitionary support tended to reinforce the existing hierarchy. However, there was at least one example of support influencing a successful rank challenge. Support was affected by potential risks and benefits, the latter including dominance through association and revolutionary alliances. An even stronger pattern overlaid associations between pack members: coalitions and resting associations were strongest between members of the same age–sex cohort, and may have enabled the eventual dominance of younger pack members over adults. Among adults, coalitionary associations were sometimes overridden by intersexual relationships. The results from this captive pack suggest that wild dogs are sensitive to differences in competitive ability. This information, in conjunction with strong affiliative bonds between littermates, is used to manoeuvre for position in the social hierarchy. It may also be important during dispersal, in encounters with other dispersing groups of the same sex. Although most features of the social structure of the captive pack were comparable to those of free-ranging packs, aspects such as the influence of relatedness on coalition formation still need to be explored.

Southern fur seals Arctocephalus australis in Peru have declined gradually over the past decade, and declined dramatically (72%) as a result of low food availability during the severe El Niño in 1997–98. In 1999, seals abandoned some historically important breeding sites. This is particularly alarming because new sites were not colonized. Our objective was to examine how habitat features and human disturbance influenced whether sites were currently used, abandoned or apparently not used in the past by fur seals for breeding. Data were collected on 14 variables at 70 potential breeding sites at three guano reserves in Peru. Discriminant analysis revealed significant multivariate differences among sites currently used for breeding, abandoned sites and unused sites (F=5.97, P<0.00001), and the model classified 74% of sites correctly. Currently used sites were less likely to have human disturbance and more likely to have offshore islands, stacked rocks, tide pools and abundant shade. Separate discriminant analyses for each reserve produced similar results. Habitat associated with thermoregulation (e.g. shade or pools) may be more important to fur seals in Peru, which breed at lower latitudes and are at greater risk of overheating on land than other populations. Habitat with minimized human access may be especially important to seals in small populations in which individuals may perceive themselves as more vulnerable because of decreased vigilance and dilution effects. Seals in our study selected breeding habitat with stacked rocks, which create shade and tide pools for thermoregulation and make human access difficult; but pups might suffer higher mortality in this habitat. We hypothesize that fur seals in Peru may exhibit an Allee effect, whereby suitability of habitat varies with population abundance.

Morphological classification of snail shells is sometimes arbitrary and misleading in correct species delimitation. This topic is investigated in four nominal species of the minute gastropods of the genus Vallonia (V. pulchella, V. excentrica, V. costata and V. enniensis). First, individuals are classified according to their shell morphology. Second, variation in ribosomal internal transcribed spacer 1 DNA (ITS-1 rDNA) is analysed. Using a cladistic approach, the nucleotide positions of the ITS-1 are elucidated on apomorphic and plesiomorphic character states (outgroup=Acanthinula aculeata; Valloniidae). The molecular data yield evidence that the morphospecies V. excentrica is a paraphyletic group, comprising just a loose arrangement of individuals with similar shells. Data on fossil shells and putative divergence time are also elucidated. In the monograph on Vallonia it has been suggested that the split between the branch of V. costata and the branch of V. pulchella, V. excentrica and V. enniensis could have been in the upper Cretaceous. Based on the ITS-1 sequence alignment and the number of variable nucleotide positions, it is suggested that the last common ancestor of both branches lived at some time during the Miocene or even the Pliocene. Therefore, the fossil stem lines of Vallonia should be newly interpreted.

This study describes the changes in stored body fat in Svalbard barnacle geese Branta leucopsis over the non-breeding period, and uses behavioural patterns to explain the variation in body fat stores. It was predicted that foraging effort would: (1) increase at low food densities; (2) increase when days were short; (3) decrease in smaller flocks; (4) be bimodally distributed throughout the day for long days, but constant for short days. Time constraints were found to be the major driving force behind foraging decisions during the shortest days of mid-winter, whereas food density was found to drive decisions during longer days. Field observations of fat stores showed that fat was rapidly accumulated at the start and end of the non-breeding period, but that fat stores remained constant during mid-winter. It is suggested that day length prevented a positive fat storage rate in mid-winter through the limitation of foraging time, even though foraging effort was high during this period. During a single day, evidence for a bimodal foraging routine was found, where feeding activity was concentrated in the early morning and late afternoon periods. This pattern was found in the full range of day lengths, which suggests that even for short days, feeding must be interrupted so that other essential activities can be conducted. It is concluded that the behavioural choices of barnacle geese were constrained by environmental conditions, and that these behavioural patterns allow the variation in fat stores to be explained.

The middle ear structures of nine species of golden moles (family Chrysochloridae) were examined under the light microscope. Auditory structures of several of these species are described here for the first time in detail, the emphasis being on the ossicular apparatus. Confirming previous observations, some golden moles (e.g. Amblysomus species) have ossicles of a morphology typical of mammals, whereas others (Chrysospalax, Chrysochloris, Cryptochloris and Eremitalpa species) have enormously hypertrophied mallei. Golden moles differ in the nature and extent of the interbullar connection, the shape of the tympanic membrane and that of the manubrium. The stapes has an unusual orientation, projecting dorsomedially from the incus. It has been proposed that hypertrophied ossicles in golden moles are adapted towards the detection of seismic vibrations. The functional morphology of the middle ear apparatus is reconsidered in this light, and it is proposed that adaptations towards low-frequency airborne hearing might have predisposed golden moles towards the evolution of seismic sensitivity through inertial bone conduction. The morphology of the middle ear apparatus sheds little light on the disputed ordinal position of the Chrysochloridae.

The highveld mole-rat, Cryptomys hottentous pretoriae is a social, subterranean, co-operative breeder with a high reproductive skew. The relationship between reproductive status and reproductive physiology in male highveld mole-rats was investigated. Testicular morphometrics, histological parameters, plasma testosterone concentrations and sperm motility parameters were studied in 31 males (14 reproductive and 17 non-reproductive males). Reproductive males were significantly larger than non-reproductive males, with testicular mass and volume corrected for body mass considerably larger for the reproductive males. Circulating plasma testosterone concentrations of reproductive males were not significantly higher than non-reproductive males (reproductive males 10.3±1.8 nmol l−1vs non-reproductive males 7.3±1.74 nmol l−1). Sperm motility parameters were measured, but no significant differences were found between reproductive and non-reproductive males. All 14 reproductive males had motile sperm, whereas 13 of the 17 (76%) of non-reproductive males possessed motile sperm. A typical ejaculate of a reproductive male contains 48.3% motile sperm characterized by a high percentage of flagellar defects, whereas the non-reproductive male has an ejaculate containing 45.6% motile sperm with a high percentage of head defects. It is apparent that non-reproductive males are not physiologically suppressed from reproducing. Moreover, the non-reproductive males are excluded from incestuous matings as a result of being the offspring of the reproductive female.

Some genera of golden moles are known to possess enormously hypertrophied auditory ossicles. These structures have been implicated as potentially mediating a form of inertial bone conduction, used by the golden mole to detect seismic vibrations. A simple model of ossicular inertial bone conduction, based on an existing model of the human middle ear from the literature, was used in an attempt to examine vibrational sensitivity in these animals. Those golden moles with hypertrophied ossicles are predicted to possess a sensitive inertial bone conduction response at frequencies below a few hundred hertz, whereas species lacking these middle ear adaptations are predicted to have a far less sensitive response in this ecologically important frequency range. An alternative mode of inertial bone conduction in golden moles, potentially conferring sensitivity to vertically-polarized seismic vibrations such as Rayleigh waves, is proposed. Certain behaviours of golden moles described in the literature are interpreted as augmenting seismic sensitivity.

Landmark-based geometric morphometric techniques were used to test the hypothesis that Petrochromis spp. from Lake Tanganyika are ecomorphologically equivalent to Petrotilapia spp. from Lake Malawi. Both genera are epilithic algal feeders and inhabit the rocky shores of their respective lakes. We investigated the morphological component of the ecomorphology hypothesis by investigating body shape, using landmark-based morphometric techniques. A MANOVA revealed significant differences among species and an ordination of all species along the first two CV axes showed clear separation of the two genera in the morphospace with Petrochromis fasciolatus as an intermediate. A thin-plate spline analysis revealed that Petrochromis spp. had a deeper, broader anterior body, larger gape, shorter anal fin base and narrower caudal peduncle than Petrotilapia spp. Basically, differences between lakes were found, but there were no similarities or clusters of presumptive ecomorphs. Based on such results, we reject the hypothesis of morphological equivalence between these two genera. However, considering the non-significant difference in body shape revealed between P. fasciolatus, Petrotilapia genalutea and Petrotilapia ‘mumbo blue’, we conclude that these three species represent morphological equivalence and hence display a best example of convergent evolution.

Food habits of tigers Panthera tigris in terms of prey abundance were studied in the semi-arid deciduous forests of Ranthambhore National Park, western India, between November 2000 and April 2001. Wild prey availability was assessed by line transects (n=8) and prey selection by the tigers was determined from analysis of scats (n=109). Compared to some other parts of the country, prey abundance was found to be high at 96.65 animals km−2. Chital Axis axis was the most abundant wild prey in the study area, followed by common langur Presbytis entellus, sambar Cervus unicolor, nilgai Boselaphus tragocamelus, wild pig Sus scrofa and chinkara Gazella bennetti. Chital (c. 31%) and sambar (c. 47%) constituted the bulk of the tigers' diet and were preferred prey. Nilgai and chinkara contributed minimally to the tigers' diet (c. 5–7% and <1%, respectively) and were used less than their availability. Domestic livestock made up 10–12% of the tigers' diet. The average weight of an animal consumed was between 107 and 114 kg reflecting a preference for large prey. The analysis reveals that parts of Ranthambhore have high prey abundance, thus making it important for long-term tiger conservation. Despite the high prey abundance, tigers were still considerably dependent on domestic livestock, posing challenges for the park management to resolve potential areas of conflict.

Habitat change is one of the main factors influencing the decline of the western population of European mink Mustela lutreola, but data on habitat selection are scarce. Landscape features influencing selection of habitat and resting sites of male European mink were studied using radio-tracking. None of the habitat descriptors accounted for the habitat selection of European mink during their activity periods. On the other hand, resting site selection was correlated with the presence of bramble patches. Intensive use of bramble patches is explained as a consequence of the need of mink for protection against predators. Moreover, the high availability of bramble patches provides the mink with easy resting sites.

Considerable behavioural evidence supports harassment by insects as the most important causal link between warm summer temperatures and low body condition of reindeer Rangifer tarandus, and that insect activity is influenced by weather condition. However, much less is known about the effect of insect harassment on individual performance, measured as reduced weight gain during summer, and the related consequences on both the reindeer pastoral economy and reindeer as a biological resource. Using climatic data, this paper develops a simple index for the analysis of insect harassment that takes into consideration weather variables known to significantly affect insect activity and/or the level of insect harassment. The insect harassment index, which is based on mid-day ambient temperature ≥13 °C, wind speed <6 m/s and cloud cover <40%, is further used to test the hypothesis that insect harassment has a negative effect on reindeer performance during summer in three Norwegian populations. Results show that harassment by insects negatively affects the autumn weight of reindeer calves, most probably through reduced grazing time and increased energy expenditure, but also indirectly by negatively influencing milk production of the dam. Moreover, female calves were more vulnerable to insect harassment than males. Insect harassment may have consequences on future reproductive performance, calving time, calf birth weight and hence neonatal mortality, and thus affect reindeer productivity. The presented index is easy to estimate and may be used to quantify and compare harassment levels on various reindeer summer grazing areas for management purposes. Our results also suggest that the expected temperature increase in the course of global warming may increase the insect-related stress on reindeer.

The Siberian enchytraeid Fridericia heliota Zalesskaja, 1990 has so far been known only through a paper in Russian briefly describing its luminescence and the major external and internal morphological features. The present study expands that preliminary description, amending it as regards the chaetal formula, the clitellar pattern, the form of the nephridia and the occurrence of glands at the spermathecal orifices, and providing a more comprehensive taxonomic characterization of the species. More detailed information is also given on its habitat and geographical range as well as on its unusual luminous properties. In F. heliota, the production of light is confined to the body wall with a pattern corresponding to that of the epidermal gland cells, although it does not involve discharge of luminous mucus. This contrasts with observations on other enchytraeids and on the oligochaetes in general. Records of luminosity in the Oligochaeta are reviewed with comments.

British red squirrel populations have been subject to landscape management practices resulting in large-scale fragmentation and defragmentation of habitat, as well as numerous historical introductions of closely related subspecies. This unique population history allowed us to examine: (1) the morphological changes to a rare native species probably caused by hybridization with introduced subspecies; (2) the impact of landscape management, specifically reforestation, on the spread of these morphological changes across the north of England. British red squirrels represent a peripheral population of Sciurus vulgaris, which is regarded as a separate subspecies (S. v. leucourus) to populations found on the European continent. British populations are in danger of extinction because of the introduction of the North American grey squirrel S. carolinensis. Repeated translocation of continental European S. vulgaris individuals to Britain over 150 years may be responsible for an alteration of the morphological characteristics of populations compared to the original subspecies description. The majority of examined populations possessed the coat colour characteristics associated with continental European subspecies. Only populations in the western region of Cumbria possessed coat colour characteristics similar to the described subspecies S. v. leucourus. Changes to landscape connectivity in Britain during the 1980s greatly altered dispersal patterns, resulting in an increase of gene flow from populations in the north-east of England and the Borders into Cumbria. The morphological characteristics of the Cumbrian population also changed over this period, from traits similar to the British subspecies before 1980, to traits similar to the continental European subspecies after 1980. This study demonstrates the threat landscape management practices and the introduction of closely related subspecies can present to rare peripheral populations.

Most primates typically use a diagonal-sequence footfall pattern during walking. This footfall pattern, which is unusual for mammals, is believed to have originated in ancestral primates in association with the use of grasping extremities for movement and foraging on thin, flexible branches. This theory was tested by comparing gait parameters between the grey short-tailed opossum Monodelphis domestica and the woolly opossum Caluromys philander, two didelphid marsupials that are strongly differentiated in grasping morphology of the extremities and in their reliance on foraging strategies involving thin branches. One hundred and thirty gait cycles were analysed quantitatively from videotapes of subjects moving quadrupedally on a runway and on poles of different diameters (7 and 28 mm). Duty factor (i.e. duration of the stance phase as a percentage of the stride period) for the forelimb and hindlimb, as well as diagonality (i.e. phase relationship between the forelimb and hindlimb cycles), were calculated for each of these symmetrical gait cycles. We found that the highly terrestrial Monodelphis, like most other non-primate mammals, relies primarily on lateral-sequence walking gaits on both runway and poles, and has relatively higher forelimb duty factors. Like primates, the highly arboreal Caluromys uses primarily diagonal-sequence walking gaits on the runway and pole, with relatively higher hindlimb duty factors and diagonality. The fact that the woolly opossum, a marsupial with primate-like feet that moves and forages mainly on thin branches, uses primarily diagonal-sequence gaits when walking supports the view that primate gaits evolved to meet the demands of locomotion on narrow supports. This also demonstrates the functional role of a grasping foot, in association with relatively higher hindlimb duty factors, protraction, and substrate reaction forces, in the production of such walking gaits.

Factors influencing prey selection by insectivorous bats are poorly understood but may be related to interspecific differences in echolocation call structure. The diet of Eptesicus fuscus was compared with prey abundance across a foraging season to examine prey selection by a bat species which has an echolocation call dominated by a frequency-modulated (FM) component. Guano samples were collected on 18 occasions from a single colony between May and September 2000. Abundance of aerial insects was measured using a black-light trap on nights when guano was collected. Regression analysis was used to test the hypotheses that E. fuscus: (1) does not take prey in proportion to their relative abundance; (2) forages preferentially for beetles in a manner predicted by foraging theory. Relative abundance (= proportion in relation to other prey types) of prey in trap samples was not a good predictor of prey in the diet across periods or within periods, which suggests selective foraging. Beetles (42–96% of the diet by per cent volume) were consistently eaten in higher proportions than their relative abundance. The absolute abundance of beetles (= abundance irrespective of the abundance of other prey types) explained 60% of the variation in beetle use with more beetles eaten when they were abundant. Absolute beetle abundance explained 66%, 53% and 47% of the variation in the use of three other prey types, with more of these prey eaten when beetle abundance was low. Finally, absolute beetle abundance explained 60% of the variation in diet breadth, which increased when beetle abundance was low. Our results agree with predictions derived from foraging theory and support the hypothesis that E. fuscus forages selectively for beetles, consuming a wider variety of prey when beetle abundance is low. These results are similar to those found for the greater horseshoe bat Rhinolophus ferrumequinum which has an echolocation call dominated by a constant-frequency (CF) component, and suggest that some FM bats may discriminate among prey taxa, at least at the ordinal level.

The honey badger, or ratel, Mellivora capensis has not been well studied despite its extensive distribution. As part of the first detailed study, visual observations of nine habituated free-living individuals (five females, four males) were used to investigate seasonal, annual and sexual differences in diet and foraging behaviour. Theory predicts that generalist predators ‘switch’ between alternative prey species depending on which prey species are currently most abundant, and diet breadth expands in response to decreased availability of preferred food types. There were significant seasonal differences in the consumption of eight prey categories related to changes in prey availability but no seasonal differences in food intake per kg of body mass. As predicted, the cold-dry season diet was characterized by low species richness and low foraging yield but high dietary diversity, while the reverse was true in the hot-dry and hot-wet seasons. In accordance with these predictions, results suggest that the honey badger maintains its intake level by food switching and by varying dietary breadth. Despite marked sexual size dimorphism, male and female honey badgers showed no intersexual differences in prey size, digging success, daily food intake per unit body weight or foraging behaviour. Results do not support the hypothesis that size dimorphism is primarily an adaptation to reduce intersexual competition for food.

Oligotrich ciliates are an abundant and important component of planktonic food webs. Classified together, based on morphology and the arrangement of the oral ciliature, they include sensu lato halteriids, strombidiids, strobilidiids, and tintinnids. The oral ciliature of tintinnids and strobilidiids, the subclass Choreotrichia, is composed of a closed circle of external polykinetids with internal polykinetids, leading to the cytostome. Strombidiid and halteriid species, the subclass Oligotrichia, have an open circle of adoral polykinetids and separate ventral polykinetids. Previous molecular phylogenetic analyses of spirotrichs have placed Halteria grandinella among the stichotrichs. Apart from H. grandinella, only one other small subunit ribosomal RNA (SSrRNA) gene sequence of an oligotrich ciliate (Strombidium purpureum) has been published. Fourteen additional species representative of the major oligotrich orders were collected to determine their phylogenetic relationships. These SSrRNA gene sequences include two strombidiids (Strombidium spp.), one strobilidiid (Strobilidium caudatum), and 11 tintinnid species (i.e. Codonellopsis americana, Eutintinnus pectinis, Eutintinnus sp., Favella panamensis, Metacylis angulata, Metacylis sp., Rhabdonella hebe, Tintinnidium mucicola, Tintinnopsis dadayi, Tintinnopsis fimbriata and Tintinnopsis tocatinensis). The subclasses Oligotrichia (excluding Halteria) and Choreotrichia form a monophyletic group and are sister group to the subclass Stichotrichia with high nodal support. The choreotrichs are confirmed as a monophyletic subclass whereas the halteriids do not cluster with the strombidiids, but within the subclass Stichotrichia. Therefore, the family Halteriidae is transferred to the order Sporadotrichida in the subclass Stichotrichia. The strombidiids form a sister clade to the subclass Choreotrichia. Since there are no clear morphological or genetical synapomorphies for these two groups combined, the separate subclasses Choreotrichia and Oligotrichia are retained. The genetic relationships among tintinnid genera do not correspond to relationships based on lorica morphology: species with agglomerated and hyaline loricae do not form separate branches but are interrelated.