Carotenoid-based status signalling in red-shouldered widowbirds (<Emphasis Type="Italic">Euplectes axillaris</Emphasis>): epaulet size and redness affect captive and territorial competition

In breeding plumage, the African male red-shouldered widowbirds (Euplectes axillaris) are black except for red carotenoid-based epaulets ('shoulder patches'), similar to the well-studied American red-winged blackbirds (Agelaius phoeniceus). To experimentally test the signal function of the red epaulets in male red-shouldered widowbirds, we manipulated epaulet colour and size (within natural variation), and observed the effects in two competitive contexts. First, in captivity, unfamiliar males with epaulets of different size or redness were staged in dyadic contests over food. Only epaulet manipulations significantly affected contest outcome; red epaulet males out-competed orange and control males, which in turn dominated blackened (no epaulet) males. Epaulet size manipulations had particularly strong effects; enlarged epaulet males vigorously defeated smaller epaulet males within colour treatments, whereas the independent effect of 'redness' (i.e., within size treatments) was much weaker. Second, in a field experiment, epaulets were manipulated prior to territory establishment. Males with enlarged epaulets acquired territories, whereas most males with blackened or reduced signals failed to obtain territories. Furthermore, among the territorial males, birds with enlarged epaulets defended larger territories and spent less time in boundary signalling, yet they received fewer intrusions from conspecifics. In contrast, epaulet manipulations had no effect on either female visitation rates or subsequent female settlement. These results concur with recent studies and suggest that the carotenoid-based epaulets of red-shouldered widowbirds are status signals, sexually selected through male contest competition for territories.     

Mesozooplankton community structure across the Antarctic Circumpolar Current to the north of South Georgia: Southern Ocean

During four summer seasons mesozooplankton community composition and structure in relation to water mass distribution were investigated along a 735-km transect running across the Antarctic Circumpolar Current (ACC) to the north-west of the island of South Georgia, Southern Ocean. Samples were obtained each year during December–January from the top 200 m of the water column at 22 stations spaced 35 km apart. Cluster analysis revealed four station groupings that were geographically consistent with the different water masses identified on the basis of temperature and salinity properties along the transect. A Sub-Antarctic Zone (SAZ) community characterised by low overall plankton abundance was present at the northernmost end of the transect on three of the four cruises, separated from a Polar Frontal Zone (PFZ) community by the Sub-Antarctic Front (SAF). The PFZ community lay between the SAF and the Polar Front (PF) and was characterised by highest overall abundance and little interannual variability. Two Antarctic Zone (AAZ) communities were found south of the PF that, although taxonomically similar, differed in overall abundance. Although there were significant differences in nutrients (e.g. silicate) and phytoplankton (Chl a) between the different water masses, these factors only weakly correlated with plankton community structure. Copepods were the largest contributors to total abundance within all station groupings (median percentage 83–90% of total) and pteropods were also proportionately abundant in the PFZ (11%). With the exception of pteropods (≤11%) and ostracods (≤3%) all other non-copepod taxa contributed <1% to total abundance. All station groups were characterised by varying proportions of a relatively small subset of species, many of which were present throughout the transect, for example, Oithona similis, Ctenocalanus spp., Euchaeta antarctica, and Rhincalanus gigas. Others were particularly characteristic of different station groups, thus Neocalanus tonsus, Clausocalanus ingens, and Calocalanus spp. were characteristic of the SAZ with few in the PFZ and none in the AAZ. Microcalanus pygmaeus was particularly abundant in the AAZ compared to other regions, as was the polychaete, Pelagobia longicirrata. Other taxa, although widespread, tended to be typical of particular water masses, for example, Calanus simillimus and Limacina helicina in the PFZ, Calanoides acutus, appendicularians and P. longicirrata in the AAZ, and ostracods and chaetognaths in the SAZ. The close physical and biological coupling observed across the ACC confirms the frontal zones and particularly the PF as features across which community properties change in the Atlantic sector of the Southern Ocean.     

Early reproduction and development of dominant calanoid copepods in the sea ice zone of the Barents Sea—need for a change of paradigms?

Reproduction and growth of the dominant copepods Calanus finmarchicus, C. glacialis, C. hyperboreus and Pseudocalanus minutus were studied on transects across the sea ice zone of the northern Barents Sea in May and June 1997. C. glacialis and C. finmarchicus were numerically dominant and also the largest component of the biomass. C. hyperboreus was rather rare. Moderate levels of phytoplankton and eventually high concentrations of ice algae supported maximum egg production rates of 53.6 and 48.5 eggs female^–1 day^–1 of C. glacialis in May and June, respectively. Results of incubation experiments were supported by a tremendous abundance of C. glacialis eggs in the water column ranging from 7×10³ to 4.4×10⁴ m^–2 in May and from 9.8×10³ to a maximum of 9.7×10⁴ m^–2 in June. In contrast, C. finmarchicus spawned only in the vicinity of the ice edge, at a maximum rate of 30 eggs female^–1 day^–1. Egg sacs of P. minutus were often observed in the preserved samples, but contained only few eggs, which may be due to loss during sampling. The presence of considerable concentrations of young stages in May and June indicated successful recruitment of C. glacialis and P. minutus. Back calculation using published stage duration estimates indicates March/April as the begin of the reproductive and growth period for these species under the first-year ice of the Barents Sea. Hence, secondary production in the study area starts at the same time as in open water regions and polynyas in the northern North Atlantic. Although the role of ice algae in the nutrition of copepods was not clarified here, the significant relationship between phytoplankton chlorophyll and egg production of C. glacialis suggests that high reproductive activity has already been achieved at moderate food concentrations.Communicated by O. Kinne, Oldendorf/Luhe     

In-depth studies of Magellanic penguin (<Emphasis Type="Italic">Spheniscus magellanicus</Emphasis>) foraging: can we estimate prey consumption by perturbations in the dive profile?

A new concept based on analysis of dive depth data was developed to help estimate prey consumption in ten free-ranging Magellanic penguins (Spheniscus magellanicus) that were brooding chicks. By simultaneously analysing the undulations in the dive depth profile (measured by time-depth recorders, TDRs) and beak opening (obtained from the recently developed intra-mandibular angle sensors, IMASEN), it was possible to determine the proportions of the undulations in the dive profile that resulted (or not) in prey capture. This methodology allowed the number of prey consumed to be estimated with a mean error of 10±6% using TDR data alone. If the mean mass of prey is known, then the overall mass of prey consumed per unit time can be determined. Additionally, the method allows estimation of the depth at which prey is taken and thus indicates how penguins exploit the water column. Due to its simplicity, the proposed methodology has applications for other Spheniscus penguin species and should be considered for other marine endotherm divers that show undulations in the dive depth profile.Communicated by O. Kinne, Oldendorf/Luhe     

Saving by freezing? Metabolic rates of<Emphasis Type="Italic"> Adamussium colbecki</Emphasis> in a latitudinal context

Standard metabolic rates of the endemic Antarctic scallop, Adamussium colbecki (Smith, 1902), were measured in austral summer and under simulated winter conditions. Average mass-specific metabolic rates were significantly different between "summer" (151.17±45.06 µl O ₂ g ^-1 h ^-1) and "winter" (106.52±39.65 µl O ₂ g ^-1 h ^-1) animals. The overall metabolic rates of A. colbecki are comparable to those of other Antarctic bivalve species, but well below those of temperate scallop species. Data for 24 scallop populations (13 species) from different latitudes give no evidence for elevated metabolic rates in A. colbecki as suggested by the concept of "metabolic cold adaptation". A world-wide comparison of metabolic rate and overall growth performance of scallops indicates that in the Antarctic scallop the energetic advantage of low basal metabolism does not counterbalance the disadvantage of the prolonged seasonal period of food shortage.     

Mate choice in the brown thornbill (<Emphasis Type="Italic">Acanthiza pusilla</Emphasis>): are settlement decisions,divorce and extrapair mating complementary strategies?

In socially monogamous birds, females may express mate preferences when they first select a breeding partner, through divorce and subsequent breeding dispersal to a new partner and through extrapair mating. We examined settlement patterns, divorce and breeding dispersal in a sedentary Australian passerine, the brown thornbill (Acanthiza pusilla), in relation to two traits known to influence extrapair paternity (male age and male size). Settlement decisions, divorce and territory switching behaviour were all female strategies that reduced their likelihood of breeding with 1-year-old males. Females preferred to settle in territories with 2+ -year-old males, were more likely to divorce 1-year-old males, and only switched territories if they had an opportunity to form a new pair bond with an old male. In contrast, female settlement and divorce decisions were not influenced by male size. Female thornbills obtain a direct benefit from preferring older males as social mates because breeding success improves with male age in brown thornbills. Nevertheless, divorce rates in this species were low (14% of pair bonds were terminated by divorce), and individuals rarely switched territories following the death of a mate. Both of these mating strategies appeared to be primarily constrained by the distance adults moved to initiate a new pair bond (1–2 territories) and by the limited availability of unpaired older males in the immediate neighbourhood.Communicated by M. Webster     

Shell-size selection by intertidal sympatric hermit crabs

This study evaluated selection for shell size by three species of tropical intertidal hermit crabs, Clibanarius antillensis, C. sclopetarius, and C. vittatus, from species of shells which are frequently used in nature. Crab size and weight were strongly and significantly related to all measured parameters of the selected shells. The strength of these relationships (r² values) depended neither on the crab nor on the shell variables taken into account. The relationships between crab size and the dimensions of the selected shells showed higher r² values than the corresponding relationships with the shells that the crabs had occupied when they were collected (0.482–0.903 in comparison to 0.091–0.652, respectively), indicating that the crabs were occupying sub-optimal shells in nature. Negative allometry was frequently found in the relationships between crab and shell variables, indicating that large crabs select and use proportionally lighter shells than do small crabs. This negative allometry was stronger for the shells used in nature (except for C. antillensis), i.e. larger crabs tended to select heavier shells in the laboratory than in nature. Different allometric relationships were also recorded among the dimensions of shells used in nature and those selected by the hermit crabs in free-access experiments: as shell length increased, the selected shells were heavier and had larger apertures than the shells used in nature. The relationships between crab size and the length and weight of the selected shells did not depend on the species of crab or species of shell, but only on crab size. Therefore, analyses using these variables can be performed without taking the species of crab or shell into account, i.e. data from different crab or shell species can be pooled. The influence of crab and/or shell species was recorded only in the models fitted for aperture length and width, variables which were more related to shell architecture than did shell length or weight. In contrast, if crab weight is used as an independent variable, different crab or shell species can be analyzed together independently of the particular shell parameter. This indicates that crab weight may be less susceptible than crab shield length to shell morphological constraints. Finally, the results indicate that the preferred size of a given shell type chosen by a given hermit crab will depend more on crab size or weight, than on the crab or shell species under consideration, i.e. crab shell-size relationships are not species specific.Communicated by P.W. Sammarco, Chauvin     

Population genetic structure of the milkfish,<Emphasis Type="Italic"> Chanos chanos</Emphasis>, based on PCR-RFLP analysis of the mitochondrial control region

The milkfish, Chanos chanos (Forsskål, 1775) is a pelagic, monotypic gonorhynchiform widely distributed in the tropical Indo-Pacific. This study evaluates temporal variability of milkfish samples from the Philippine archipelago, and spatial variability at two geographic scales based on restriction fragment length polymorphism (RFLP) analysis of a portion of the mitochondrial control region. High levels of genetic diversity characterize the milkfish control region (mean h=0.908, =1.59%), with 74 haplotypes detected among the 367 fish analyzed. For temporal analysis of Philippine samples, milkfish were collected over 2 years from three sites (inter-annual variation), and sampled twice within a year during different seasons at four sites (intra-annual variation). No significant temporal variability was detected between or within years. Significant spatial differentiation among the Philippine samples was observed (F_ST=0.006, P<0.05), with two northeastern samples, Claveria and Dingalan, found to be genetically distinct. However, an hierarchical analysis of molecular variance (AMOVA), where samples were grouped into four geographic regions, revealed very low levels of genetic partitioning, with less than 1% of the total variation attributed to between-region differences, and lack of genetic structure. Nonetheless, the existence of putative northeastern Philippine populations is not discounted. Strong genetic structure across broad geographical scales was revealed by AMOVA, with 11% of the molecular variance based on haplotype frequencies allocated between three distinct groups: Indian Ocean, west Pacific (Philippines) and north central Pacific (Hawaii) The broad-scale genetic structure points to limited gene flow among disjunct Indo-Pacific populations.Communicated by T. Ikeda, Hakodate     

Male residence and the patterning of serum testosterone in vervet monkeys (<Emphasis Type="Italic">Cercopithecus aethiops</Emphasis>)

This study investigated hormonal and demographic processes underlying unimale and multimale mating systems in primates. Reproductive skew and challenge models of male competition provide conflicting predictions of the relationship of male residence to group composition and androgen regulation. These predictions were tested using endocrine and socioecological data from Kenyan vervet monkeys (Cercopithecus aethiops). Serum samples from 57 adult male monkeys, drawn from 19 separate groups and 4 populations, were assayed for testosterone by radioimmunoassay. Male ability to respond to conspecific challenge was assessed by their testosterone response to the capture protocol.Analyses showed that reproductive skew models were useful predictors of intergroup and interpopulation variation in male residence and T profiles. The Limited Control model of male residence was supported by positive correlations of the number of males per group with the number of females without dependent offspring, demonstrating that monopolization potential was a key determinant of male residence. Testosterone concentrations under conditions that elevated serum levels were positively correlated with infanticide risk, supporting the Concession model. Population comparisons provided evidence for increased T responsiveness where groups were predominantly unimale. Unimale populations were from sites with higher rainfall, suggesting that ecological factors contributed to population differences in male residence and T regulation.For species name, we follow the 2000 taxonomy of the IUCN/SSC Primate Specialist Groups workshop (Grubb et al. 2003)     

Microgeographic heterogeneity in spatial distribution and mtDNA variability of gray mouse lemurs (<Emphasis Type="Italic">Microcebus murinus</Emphasis>, Primates: Cheirogaleidae)

The objective of our study was to investigate the spatial distribution and genetic structure of a solitary primate at the microgeographical scale of adjacent local populations. We obtained spatial data and tissue samples for mtDNA analysis from 205 gray mouse lemurs (Microcebus murinus) captured along transects and within 3 grid systems within a 12.3 km² area in Kirindy Forest, western Madagascar. Our capture data revealed that, even though the forest was continuous, gray mouse lemurs were not evenly distributed, and that daily and maximum dispersal distances were significantly greater in males. The frequency distribution of 22 mtDNA D-loop haplotypes was highly skewed. Nine haplotypes were unique to males, indicating male-mediated gene flow from surrounding areas. The geographic distribution of haplotypes revealed that males were also more dispersed than females. Females with the same haplotype showed a tendency towards spatial aggregation, and the correlation between genetic and geographic distances was higher in females. In several areas of the forest, however, spatially clustered females were not of the same haplotype, and females were not always found in clusters. Hence, in contrast to suggestions from previous studies, matrilineal clustering is not the only way females are socially organized. In addition, our study revealed heterogeneity and patterns in population structure that were not evident at smaller spatial scales, some of which may be relevant for designing conservation strategies.Communicated by C. Nunn