The hypothesis that the periods of dormancy previously described in the millipede Polydesmus angustus may be photoperiodically induced diapauses was tested experimentally. In this species, biennial individuals exhibit two successive periods of dormancy: aestivation in the penultimate stadium (stadium VII) and reproductive dormancy in the adults, which emerge in autumn. It was first established that the reproductive dormancy is not a thermally controlled state of quiescence. When adults emerging in autumn were kept at 16 °C under natural photoperiod, their reproduction was delayed for several months in comparison with adults emerging in spring at similar temperatures. This indicates that the reproductive dormancy begins with a period of diapause. Further experiments provided evidence of a photoperiodic induction of the adult diapause. When millipedes were reared under short day length (L:D 12:12 h) throughout their development, they required more time to reproduce than millipedes reared under long day length (L:D 16:8 h) at the same temperatures. Photoperiod influenced reproduction in females, but no significant effects were detected in adult males. On the other hand, stadium VII was markedly longer at L:D 16:8 h than at L:D 12:12 h in both sexes, which strongly suggests that aestivation is also induced by photoperiod. However, the effects on the duration of stadium VII varied among individuals, some of which showed no response to long days. This study is the first to document photoperiodic regulation of the life cycle in the class Diplopoda, a trait common in other classes of terrestrial arthropods.

From naked mole rats to elephants, research in wildlife biology increasingly entails interference with individual animals, and there are innumerable reasons to justify this interference for the proper management of populations. As Boyd (2002) has emphasized, however, there is a central question about the extent to which it is reasonable to intervene in a population when the species is endangered. This comment arose as part of a debate published last year in this journal on the possible harmful effects of immobilizing black rhinos and fitting them with radio-collars. Alibhai, Jewell & Towindo (2001) had prompted the debate with a discussion on the possible reduction of fertility in black rhinos caused by immobilization followed by an account of the wounding of a rhino by an ill-fitting radio-collar (Alibhai & Jewell, 2001).

We are writing to respond to an article published in the Journal of Zoology257: 37–42 by Tuyttens, Macdonald and Roddam. The article describes analyses to investigate the potential effects on European badgers of the fitting and monitoring of radio-collars. The results suggest that there is a transient detrimental effect on body condition for up to 100 days post fitting.

The underlying aim of our paper (Tuyttens et al., 2002) was to draw attention to the desirability – for reasons of science and animal welfare – of applying the scientific method to test for effects of techniques in field biology. Therefore, we are grateful to Delahay et al. (2002) for their response which not only advances that aim, but provides us with the opportunity to elaborate on the importance of the issue. In this reply we focus first on the questions they raise about our analysis and then, second, and more importantly, we set the answers in a broader context of what we foresee as a new era of animal welfare science as a component of conservation biology.

Flat lizards Platysaurus intermedius wilhelmi occur on small discrete rock outcrops in Mpumalanga Province, South Africa. These rock outcrops are structurally simple and this, combined with the lizard's behaviour (ambush foraging in the open), make them ideal for field studies of anti-predatory behaviour. Lizards were approached in the field and how escape behaviour was influenced by habitat and age-sex class was recorded. Juveniles (c. 4 months of age) responded quite differently to an approaching human ‘predator’ compared with adult males and females (which responded similarly). Compared to adults, juveniles allowed a closer approach by the investigator; took longer to find a refuge and therefore fled further; were more likely to remain visible in the open and maintain visual contact with the investigator; and more likely to flee into vegetation when given the opportunity to take refuge in a crevice. We suggest that because a greater suite of predators (including arthropods living in rock crevices) feed on small juvenile lizards, this may affect their choice of refuge and result in the avoidance of crevices when chased. Finally, because juveniles were frequently found on small rock outcrops, the influence of rock outcrop area on anti-predatory behaviour was tested. Escape behaviour (time to refuge) was independent of rock outcrop area.

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.

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.

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.

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.

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 results are reported of a survey of rodents on 10 forested land-bridge islands ranging in size from 0.2 to 350 ha in the state of Bolívar, Venezuela. The islands were contained within a lake formed c. 12 years before the study by the damming of the Caroni River for hydroelectric power. Rodents were sampled on each island by live-trapping along transects that sampled all available habitat types on each island, and microhabitat structure was measured at each trap station. A total of 674 captures of 359 individuals of six species of rodents was recorded. Species composition changed from the largest to the smallest islands, and small and medium islands (0.2–11 ha) displayed the typical effects of insularity, with fewer species and increased abundances and biomass. The largest island (350 ha) seemed to function more like a mainland. Most species were associated with a suite of microhabitat variables. It is suggested that release from top-down control by predators was responsible for higher abundances and biomass on the smaller islands and that predators moving between large islands and other nearby landmasses help maintain a mainland community structure on large islands. However, changes in species composition on smaller islands may be the result of patchy occurrences of some species before isolation, changes in microhabitat structure following isolation, and species-specific microhabitat requirements.

Data are presented on a bat assemblage captured among 10 water holes in Colorado over 5 years. The assemblage consists of Myotis ciliolabrum, M. evotis, M. lucifugus, M. thysanodes, M. volans, Eptesicus fuscus, Lasiurus cinereus, Lasionycteris noctivagans and Corynorhinus townsendii. Results show that reproductive females and juveniles are captured in higher frequencies at water holes containing higher water hardness and that water hardness correlates highly significantly with dissolved calcium content. Also presented are laboratory test data on the stomach volume of Eptesicus fuscus that provide a model for understanding the effect of dissolved calcium content in water as a significant resource. These data indicate that water holes provide supplemental sources of calcium for bats not provided by diet.

The kinematics and hydrodynamics of swimming chironomid larvae were investigated with the aid of videography and dye streamers used to visualize near-body flow. Chironomids employ a characteristic ‘figure-of-eight’ swimming technique based on high-amplitude side-to-side bending of the body. These scissor-like movements produce relatively slow (two body lengths (BL) s−1) forward motion but also serve to support the weight of the insect against its own negative buoyancy. The main wake element identified by the present technique consisted of a discrete ring vortex with an external diameter of c. 0.3 BL which was shed to the rear of the body towards the end of each half-stroke. During level swimming, the jet of the vortex was directed 10° below the horizontal plane indicating that it was mainly providing thrust. An additional, but poorly defined, flow was associated with the rapid downwards motion of the head at the start of each half-stroke and it is proposed that this contributes to the vertical force needed to support the weight of the body during swimming.

The maned wolf Chrysocyon brachyurus is the largest canid inhabiting South America. Its geographic distribution includes the open fields of Brazil's central area, which is currently undergoing agricultural expansion. The diet of the maned wolf and its seasonal variation was determined on a dairy cattle ranch (São Luís farm, 566 ha) in the State of Minas Gerais, Brazil. From January to December faeces of the maned wolf were collected monthly (n = 150 scats; 397 food item occurrences). Twenty-nine taxa were identified from scats, 18 of animal origin (46% or 183 occurrences) and 11 of plants (54% or 214 occurrences). The fruits of Solanum lycocarpum were the dominant food item in our study (29%). Mammals contributed 13%, arthropods 12%, birds 11% and reptiles 2% of the food items. Arthropods and fruits were prevalent in the rainy season and mammals in the dry season. As expected for a heavily farmed region, frugivory results were at the lower end of the diversity scale (9–33 species) and included four old garden species. No previous study of the diet of maned wolf has registered as many species of Solanaceae as this one. Although dietary richness was lower, the main food items (wolf fruit, armadillos, rodents, birds) were the same as study sites in ‘cerrado’ and upland meadows. In this region, the open habitats occupied by the maned wolf were previously covered by Atlantic forest, suggesting that landscape modification such as cattle ranching has opened new frontiers for distribution expansion of the maned wolf. The impact of loss of dietary richness and the increase in Solanaceae on the survival of the maned wolf need to be evaluated.

This paper examines the relationship between the number of occupied setts in a badger social group territory and badger group size, breeding success, and status of infection with Mycobacterium bovis (TB). The data used were from a long-term epidemiological and ecological study of a high-density population of badgers Meles meles in south-west England. The number of occupied setts in a social group was significantly and positively related to the number of badgers caught in the social group, so that as a social group increases in size, badgers occupy more of the available setts. This relationship remained significant when numbers of adults, adult males and adult females were examined. The number of breeding females, number of cubs and sex ratio was not related to the number of occupied setts in a social group. It is possible that the advantages to breeding females of a larger number of setts available to breed in might be outweighed by the increased aggression found in larger groups. The TB score for prevalence and for incidence of social groups was significantly and positively related to the number of occupied setts in a social group, such that the more occupied setts there were in a territory, the higher the TB index of the group. Possibly the setts themselves contribute to the persistence of TB within social groups, or badgers infected with TB might show a difference in behaviour from uninfected badgers resulting in their increased use of outlying setts.

Journal of Zoology Volume 258 Part 1, September 2002 (pages 115–129)

‘Temperature, genetic and hydroperiod effects on metamorphosis of brown frogsRana arvalisandR. temporariain the field’, by J. Loman