The mating behaviour of two species of spiders in their natural habitat is described. A male Leucauge nigrovittata from Sumatra mated repeatedly with a newly-moulted adult female which he had been guarding. Each copulation was in response to a body-jerking ‘invitation’ display by the female. In Tylorida ventralis from Sulawesi, males and females copulated almost continuously from dawn until dusk over several days. Females switched from diurnal to nocturnal web construction during this time. After temporary separations, females were almost as likely as males to solicit the re-initiation of copulation. In both species the sexes are of approximately equal size. The possible adaptive reasons for the extended mating times in T. ventralis are discussed.

Applications published in the Bulletin of Zoological Nomenclature

The following Applications were published on 30 September 1999 in Vol. 56, Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on any of these applications is invited for publication in the Bulletin and should be sent to the Executive Secretary (I.C.Z.N.), c/o The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: [email protected]).

Bite strength is of great importance to carnivores, as their jaws must produce forces of sufficient magnitude to kill and consume their prey. Spotted hyenas, well known for crushing and consuming bones, were studied to determine how tooth and jaw growth affect bite strength and feeding behaviour. Nine captive individuals, aged 6 months to 2 years of age, were sampled as they grew. At 8- to12-week intervals, morphological measurements that estimated jaw muscle mass, tooth size and skull size were taken. Using a force transducer, bite force was measured directly for these juveniles as well as other captive individuals of different ages. In addition, feeding behaviour and performance were quantified periodically by bone tests in which individuals were offered a sheep femur for 15 min. Behaviour and performance were expected to change with the shift from juvenile to adult dentition. Results were not entirely as expected. Morphological measurements of growth reached a plateau at about 20 months, whereas bite strength increased in a linear fashion up to 5 years of age. A fundamental change in tooth use during bone cracking followed the replacement of deciduous teeth with permanent teeth; the primary area of tooth use moved from anterior to rearmost premolars, increasing the mechanical advantage of the jaw adductors. The timing of this shift seemed to be a function of a decrease in gape limitation as a result of growth as well as caudal movement of the premolars. Our data demonstrated that juvenile hyenas had not achieved adult feeding performance levels at 12 months of age, when they are typically weaned in the wild. This suggests that recently weaned cubs may be at an increased risk of starvation and that selection might favour later weaning times.

Sleepy lizards Tiliqua rugosa are large, long-lived, Australian skinks that live in stable home ranges. Previous studies showed that the lizards have a good sense of orientation and navigation towards home. However, since they are active only 4 months in every year, there is a question about which environmental cues enable them to locate their home boundaries. In the present study we determined whether surrounding olfactory and chemical cues are important for the sleepy lizards for home-range maintenance. In the first part of the study we showed in the laboratory that zinc sulphate (ZnSO4) is a chemical that induces anosmia and prevents detection of chemical cues in sleepy lizards for up to 30 days, with no evident long-term harm to the animals. We then monitored the location of 50 radio-tagged sleepy lizards. After determining their home ranges they were divided into three groups: an untreated control, a saline-treated (sham-operated) group and a ZnSO4-treated group. Starting 2 days after the treatment, the home ranges of all lizards were re-determined within the period that ZnSO4 blocked olfaction. Three variables were calculated comparing home ranges before and after treatments: the change in home-range size, the distance between the centre of activities of the lizards, and the overlap between the individual home ranges before and after treatments. Lizard home ranges varied from 1.9 ± 0.6 ha to 4.6 ± 1.4 ha before treatment, and 1.3 ± 0.3 ha to 3.1 ± 0.7 ha after treatment. There was no significant effect of treatments or of sex on the home-range size or on the distances between the centre of activities, or on the overlap between the individual home ranges. Our data indicate that olfaction and vomerolfaction are not a major cue for home-range maintenance in the sleepy lizards.

The behaviour of free-ranging groups of Callithrix aurita and Callithrix flaviceps, small-bodied omnivorous marmosets, was monitored at two sites (Cunha and Caratinga, respectively) in south-eastern Brazil during equivalent periods in consecutive years. Quantitative behavioural data were collected using a standard scan-sampling protocol, with scans being taken at 5-min intervals throughout the daily activity period. Data were collected during the wet season months of January and February at Cunha in 1993 and 1994, and during the dry season months of July and August at Caratinga in 1985 and 1986. In both cases, wet-season precipitation levels were considerably higher during the first year. Activity budgets, daily ranging and the foraging behaviour of both species were compared between years. Significant between-year differences were observed in several behaviour categories in both species, and were consistent with seasonal and longitudinal variation in resource abundance. In C. aurita, apparent changes in fruiting patterns resulted in a significant increase in exudate feeding in the second year, both in relative and absolute terms, but little change in prey foraging. By contrast, a marked reduction in arthropod prey in 1986 resulted in major changes in most aspects of C. flaviceps behaviour, with the exception of plant feeding. While consistent with behaviour patterns in other marmoset species, the numerous significant differences observed in the present study further highlight the problems associated with the evaluation of intertaxon differences in the behaviour and ecology of primates, in particular marmosets, on the basis of relatively short-term studies of a few individuals, often at only a single study site.

Although there has been extensive research on bottlenose dolphins (genus Tursiops), much controversy over the classification within this genus still exists. Even though many morphological variants occur, the prevailing view is for a single species, Tursiops truncatus (Montagu, 1821). To evaluate this view, two sympatric forms of bottlenose dolphins inhabiting Chinese waters were examined using multivariate statistical analyses of osteological data for two meristic and 18 cranial morphometric characters from 71 specimens. Cluster and principal components analyses revealed clear osteological separation of the two forms. Furthermore, the distributions of the total number of vertebrae and several proportions of cranial characters were non-overlapping between the two forms. These results provided strong evidence that the two sympatric forms of Tursiops in Chinese waters are isolated reproductively and do not support the current view of a monotypic genus. Classification functions of a discriminant analysis and a key of several characters were developed to help researchers identify unassigned specimens. Provisional names (T. truncatus and T. aduncus) were suggested, but a formal taxonomic revision of this genus is still required. However, regardless of the nomenclature, immediate amendments to all present wildlife conservation legislation are strongly urged.

A recent survey of the high-mountain zone of the Madagascar Parc National (PN) d'Andringitra revealed the presence of an apparently isolated troop of the ring-tailed lemur Lemur catta. These animals display phenotypic and ecological characteristics that are unusual for the monotypic genus Lemur, thus raising the possibility that they are members of a different undescribed species. We present analyses of two mitochondrial genes to test the hypothesis that L. catta from Andringitra should be considered a distinct species. The results indicate that taxonomic revision is not warranted under the expectations of the phylogenetic, coalescent, or biological species concepts. Rather, the genetic patterns observed among the Andringitra and lowland mitochondrial haplotypes are consistent with those expected for a single species.

Three groups of factors which might affect the blood biochemistry of red deer (Cervus elaphus) were examined. These were the wounding site (head/neck versus chest), the stalker who collected the blood (coupled with the geographical area where each deer was shot), and the sex and nutritional status of the deer. The activities of muscle enzymes, creatine kinase, aspartate amino transferase, and lactate dehydrogenase, were markedly higher in the plasma of deer shot in the chest as compared with those shot in the head or neck; the plasma also looked different, being deep cherry red in colour in the chest-shot deer. Other biochemical measures were unaffected by the wounding site. Blood collected by one stalker from the chest cavity had higher activity of the liver enzyme, glutamate dehydrogenase, higher concentration of potassium and lower concentrations of sodium and chloride than blood collected by another stalker from a knife wound in the base of the neck. Otherwise the method of collecting blood had very little effect on blood biochemistry. Cortisol was unaffected by the wound site nor, probably, by the method of collecting blood, but was highest in the stags living in a more mountainous region and in those deer that had to be shot twice before they died. Stags, which were shot in the rutting season and were probably fasting, had significantly higher concentrations of free fatty acids than hinds. Lactating hinds had significantly less fat on their kidneys than non-lactating hinds and stags. All hinds, which were shot in winter, had lower concentrations of urea than stags, which were shot in the autumn.

A type of apparent photoreceptor, the nuchal organ, is described in the sepiolid squid Euprymna scolopes. These vesicles are identified in a variety of other cephalopod species. The nuchal organs have specific structural similarities to the already well-described photosensitive vesicles. Both organs have a distinct vesiculate structure and possess a rind consisting of several types of cells. Certain rind cells from both organs send processes with microvilli into the lumen of the vesicle forming a core area. Within the cores of both organs numerous microvilli occupy much of the core volume. These histological similarities provide strong evidence that the nuchal organs, like the photosensitive vesicles, are photoreceptors. Three distinct differences also exist between the nuchal organs and photosensitive vesicles: (1) the location of the nuchal organs differs from that of all known photosensitive vesicles in cephalopods; (2) the location of the nuchal organs is highly conservative in decapod cephalopods compared to the varying locations of the photosensitive vesicles; (3) the degree of structural order generally found within photosensitive vesicles is not apparent in the nuchal organs. The function of the nuchal organ is probably different from previously described photoreceptors in cephalopods.

Following deglaciation of the Cook Inlet region of Alaska approximately 16 000 years ago, anadromous threespine stickleback (Gasterosteus aculeatus) rapidly colonized emerging lakes and rivers forming resident, freshwater populations. Although the precise body shape of the ancestral marine population is unknown, marine sticklebacks sampled from both Pacific and Atlantic sites present remarkably little body shape variation among populations, which suggests that the morphology of any of the marine populations could be used to represent the ancestral phenotype. To infer the net evolutionary trajectories of body shape change in the Cook Inlet radiation, derived body shapes of lacustrine samples were compared to the presumptive, primitive body shape, represented by the mean shape of two anadromous samples from Cook Inlet. In general, some derived body shape traits are shared by all freshwater populations but many traits evolved in opposite directions. The principal axes of shape variation among freshwater sample means were computed using Principal Components Analysis. The strong correlation between the direction of the principal component axes and lake habitat variables suggest that populations evolved toward selection peaks that are biased along the component axes due to biotic and abiotic features of the lakes.

Daily and seasonal activity of the mountain tapir Tapirus pinchaque was measured along trails and at two salt licks in mature and secondary montane forests of the Central Andes of Colombia using active and passive infrared monitors. Daily activity of the mountain tapir showed a clear bimodal pattern, with maxima during the early hours of the day (5:00–7:00) and early hours of the evening (18:00–20:00). This bimodal activity pattern has also been found in radio-tracking studies of T. terrestris and T. bairdii. Overall, there was more activity in mature than in secondary forest and at lower (3100 m) than higher altitudes (3600 m). Night-time activity was higher during full moon than during quarter and new moons. There was no correlation between monthly rainfall and seasonal activity. The time the mountain tapir was at the salt licks varied between 1 and 190 min. Daily activity at Quebrada Paraiso salt lick (3400 m) showed a clear bimodal pattern with maxima around midnight (0:00–2:00) and midday (11:00–14:00). These patterns of diurnal and nocturnal activity of the mountain tapir could be used in combination with future radio-tracking studies to better understand habitat use by this species.

Brown long-eared bats Plecotus auritus occupying 30 summer roosts in north-east Scotland were studied over 15 years. During this time 1365 bats were ringed, and a further 720 recaptures were made. Individual bats showed a high degree of roost fidelity, returning to one main roost site; < 1% of recaptured bats had moved among roost sites, and all recorded movements (n = 5) were < 300 m. Adults of both sexes were loyal to the roost sites at which they were first captured, indicating long-term use of roosts. At least some juveniles (n = 32) of both sexes returned to the natal roost. Mark–recapture estimates indicated that colonies of this species were substantially larger (c. 30–50 individuals) than assumed in previous studies. Plecotus auritus differs from most other temperate zone, vespertilionid species in that there was no evidence of sexual segregation during summer, with males present in all colonies throughout the period of occupancy. Population structure in summer seems to be consistent with a metapopulation model, with discrete sub-populations showing minimal interchange. The group size, colony composition and population structure described in this species may be associated with the wing shape (particularly aspect ratio) and foraging behaviour of P. auritus. It is postulated that relative motility, linked to wing structure, may affect the distribution of individuals, and may have implications for the genetic structure of this species. Correlations between aspect ratio and both colony size and migratory behaviour, across British bat species, indicate that wing shape could be an important factor contributing to patterns of social behaviour and genetic structuring in bats.

The energetics and burrowing behaviour of the semifossorial Octodon degus (Rodentia: Octodontidae) were investigated and compared with that of more specialized fossorial rodents. An open-flow respirometry system was used to record energy expenditure of single degus inside respirometers partially filled with soft (moist) or hard (dry) soil. In addition, digging behaviour was recorded in groups of three animals inside a large terrarium under controlled conditions of food, photoperiod and temperature. In the field, the digging activity of degus was monitored, along with seasonal variations in rainfall, content of soil moisture and soil hardness. Mass-specific metabolic rate during digging was found to be higher in animals burrowing in soft soils compared to hard soil. However, animals burrowing in soft soil removed more soil per min than animals in hard soil. Thus, gram per gram, excavating in hard soil was energetically more expensive. The digging cost of semifossorial degus tends to be either similar to or above those of similarly sized, but more fossorial, rodents. In the field, heightened digging activity coincided with the occurrence of rainfall, greater content of soil moisture and relatively soft soil conditions. Degus generally use their front feet and teeth to shear the soil; disposal of accumulated debris being carried out by moving their front and hind feet backwards. We also observed the establishment of digging chains when two or three individuals burrowed at the same site. As far as digging is concerned, the behaviour of degus is similar to that of other fossorial rodents, such as African bathyergids and the more closely related South American ctenomyids.

This study examines the ecology of a population of Geoffroy's side-necked turtle Phrynops geoffroanus inhabiting a polluted urban river in Ribeirão Preto city, São Paulo state, south-eastern Brazil. Adult turtles fed mainly on Chironomidae larvae and pupae (Chironomus cf. plumosus, 100% of occurrence frequency) and domestic waste, but they also consumed terrestrial items (cockroach, snails) and carrion. Juvenile turtles showed more feeding diversity than the adults and exhibited a trend for predation on Chironomidae pupae, but this is not reflected in resource partitioning. The elevated number of turtles (170–230 turtles/ha of river) and biomass (255–345 kg/ha of river) inhabiting this urban river is probably the result of the abundance of sewage and organic waste produced by humans, the absence of predators, and increased availability of nesting areas. Such factors convert this area into an environment highly advantageous for the survival of Geoffroy's side-necked turtle.

Recent characterization of nuclear ribosomal small subunit (SSU) genes has shown that variant nucleotides within this region could be useful for species and species group identification within the genus Lymnaea (Gastropoda: Lymnaeidae). This study aimed to characterize a range of populations of Lymnaea natalensis Krauss, 1848 on Madagascar, and addressed two related questions. First, is there any evidence of intraspecific variation of the SSU and, if so, what might be its significance? Secondly, might this variation jeopardize the use of SSU for lymnaeid taxonomy and phylogeny? Lymnaea natalensis (n = 212) was collected from 17 sampling localities, spanning the northern and southern ends of the island. Variation within a selected region of the SSU known to vary between species, the V1 and V2, was assayed by polymerase chain reaction (PCR) linked restriction fragment length polymorphism (RFLP) and denaturing gradient gel electrophoresis (DGGE) analysis. The PCR-RFLP profiles indicated a striking dimorphism across populations at two restriction site loci (CfoI & MspI) within the E10-1 helix of the V2 region. The observed RFLP variation was confirmed by direct sequencing and by genomic digestion with subsequent hybridization. Putative heterozygotes were also encountered and in these individuals the SSU arrays composed of two distinct types approximately 1% divergent. A severe departure from Hardy–Weinberg equilibrium with a highly statistically significant (P < 10-5) heterozygote deficiency was found and genetic variation among populations was highly structured (Fst = 0.53). The geographic distribution of the variants was mapped, revealing that one variant was restricted to higher, predominately colder environments and was thought to be an adaptation. The molecular basis of the SSU variation was caused by single nucleotide polymorphisms (SNPs). To test for the possibility of cryptic taxa, an analysis of individuals representative of the SSU variant types with isoenzyme analysis (ISA), randomly amplified polymorphic DNA (RAPDs) and PCR-RFLP analysis of the ribosomal Internal Transcribed Spacer (ITS) was performed. Little variation was revealed and none that correlated to the groups suggested by SSU, confirming that the SSU variation was intraspecific. The levels of intraspecific divergence of the V1 and V2 within Lymnaea were not appreciably different (1%) from interspecific and would therefore question the validity of these data for lymnaeid taxonomy and phylogeny.

A total of 104 Damaraland mole-rats was collected in 1988, 1990, 1991 and 1993 from an open water canal in the Otjiwarongo district of Namibia. The mole-rats were apparently dispersing above ground when they fell in and drowned; they had travelled at least 20 m in distance from their natal burrow systems. The dispersing mole-rats of both sexes were significantly larger than 73 animals trapped from a nearby control population. Sequential tooth wear and eruption patterns showed that dispersers were adult animals. There was a bias towards male mole-rats dispersing. Furthermore, in years with lower rainfall (< 300 mm) it was found that dispersal became even more male biased. Eighty per cent of males and 40% of female dispersers showed signs of reproductive readiness, with more males than females being ready to breed. Thus it appears that, in these otherwise totally subterranean mammals, at least one form of dispersal occurs, above ground, and that adult mole-rats are the principal dispersers.

Mesenchytraeus antaeus sp. n., an unpigmented giant enchytraeid inhabiting Carmanah valley (48°37′N, 124°44′W), Vancouver Island, is described. With its 102–127 segments and the size of fixed specimens reaching 61×2.9 mm (diameter maximal at midbody), this species qualifies as one of the largest members of the Clitellata family. The body wall comprises: a surprisingly thin cuticle (0.002 mm), the epidermis (0.02–0.04 mm), circular muscles (0.02 mm), plus thick longitudinal muscles (eight-layered, totally 0.16–0.18 mm) divided into seven unequal fields. The chaetae are grouped in bundles of two to seven but show some unusual traits which suggest interesting convergences with the lumbricids. For instance, the most lateral chaeta in a bundle is always the largest; the larger chaetae exceed in length one-third of a millimetre; the ectal tips of the chaetae are oriented in opposite directions in the anterior and posterior halves of the body. The spermathecal pores are prominent and open laterally in the middle of V, an unprecedented location among enchytraeids. The nature of the spermathecal ampullae (free, diverticulate and greatly elongate) indicates a close relationship to a group of species that seems to be confined to the Pacific coast of North America and the Arctic coast of Siberia. This amphi-Beringean lineage includes other gigantic forms such as M. harrimani Eisen, 1904 and M. maculatus Eisen, 1904, from which, however, M. antaeus is clearly differentiated by the lack of pigmentation, the different size distribution of the chaetae within bundles, and the unique arrangement of the male apparatus. The latter includes unusually long sperm funnels (extending over 10 segments) and two groups of large accessory glands opening through two circular papillae at the base of each penial bulb, independently of the penial pores. All other congeners known to possess this type of glands (e.g. M. franciscanus Eisen, 1904) are at most only half as large as M. antaeus. The chaetal muscles of M. antaeus are heavily infested by encysted nematodes.

The giant mole-rat Cryptomys mechowi exhibits an extreme reproductive division of labour. Reproduction in the colony is restricted to a single breeding pair. Non-reproductive individuals do not breed unless removed from the colony and paired with unfamiliar conspecifics. Circulating basal concentrations of luteinizing hormone (LH) as well as LH levels measured in response to a single exogenous gonadotropin hormone (GnRH) challenge are not significantly different between the reproductive and non-reproductive groups of either sex. Neither non-reproductive males nor females are physiologically suppressed at the level of the pituitary, as they do not exhibit a reduced pituitary secretion of LH or decreased sensitivity to hypothalamic GnRH. Rather, reproductive skew seems to result from an inhibition to mate with familiar conspecifics. Familiar individuals housed in the absence of the breeding pair did not breed, whereas pairing unfamiliar individuals of the opposite sex resulted in sexual activity and recruitment. The results from the giant mole-rat suggest that reproductive skew may result from a predominantly behavioural repression in non-reproductive females arising from an incest avoidance mechanism.

The rare Australian venomous elapid snake ‘Echiopsis’ atriceps has been the subject of considerable taxonomic instability with the five known specimens assigned to four genera by various authorities. Phylogenetic affinities of the rare Elapognathus minor also are poorly understood and have been the subject of some disagreement. To examine the phylogenetic affinities of these two rare taxa, a molecular data set comprising 1680 base pairs of mtDNA was assembled from a representative of each of the terrestrial Australian viviparous elapid genera and two species of Drysdalia, a genus about which there also has been phylogenetic controversy. A total of 936 base pairs of 12S rRNA, 454 base pairs of 16S rRNA and 290 base pairs of cytochrome b mtDNA were sequenced for 15 species. The Asian elapid Naja naja was used as the outgroup. These mtDNA regions provided 195, 38 and 72 parsimony informative sites, respectively, for a total of 315 parsimony informative characters. Unweighted phylogenetic analyses were performed under both parsimony and neighbour-joining criteria. Parsimony analyses of the unweighted, combined data set resulted in a single fully resolved most parsimonious tree 1225 steps long. The neighbour-joining tree differed by only a single weakly supported branch. These data strongly support a sister group relationship between ‘Echiopsis’ atriceps and the Australian broadheaded snakes of the genus Hoplocephalus with a bootstrap value of 99%. Templeton tests soundly reject all previous taxonomic arrangements for this species. Our data also strongly support a sister group relationship between Elapognathus minor and Drysdalia coronata with a bootstrap value of 98%. Importantly, Drysdalia coronata and Drysdalia coronoides do not form a monophyletic group, supporting some previous studies. Based on our results, we allocate ‘Echiopsis’ atriceps to a new monotypic genus and re-describe Elapognathus to include ‘Drysdalia’ coronata.

Differences in behaviour, such as the mate-locating tactic, may favour the evolution of interspecific and intersexual differences in the structure of the visual system. This notion was tested by examining the relationships between eye size, body size, sex and mate-locating tactic in 16 species of butterflies. The 16 species were grouped in eight phylogenetically close pairs that differed in the mate-locating tactic displayed by the males. Eye size was characterized by eye surface area, which was estimated from a set of linear eye measurements made on 10 individuals of each sex in each species. The major findings were that eye surface area was positively correlated with body size both among and within species, males have larger eyes relative to their body size than females, and after controlling for body size and phylogeny, interspecific differences in the mate-locating tactic of males does not explain interspecific variation in the size of male eyes or in the magnitude of sexual difference in eye size. The optical and behavioural consequences of variation in eye size with body size and sex will await a better understanding of how eye structure varies with size within and between species.