Do male barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>) experience a trade-off between the expression of multiple sexual signals?

Some sexual selection models envisage exaggerated male secondary sexual characters to be costly and therefore reliable indicators of the quality of potential mates to choosy females. If male secondary sexual characters have a natural selection cost, they may be linked to each other by reciprocally constraining relationships that would prevent individual males from increasing their level of multiple signaling. Barn swallow (Hirundo rustica) males have at least two costly signals relevant to socio-sexual interactions: tail length and song. Under the hypothesis that a trade-off exists between male signals, we manipulated the maintenance cost of tail ornaments to test whether this reduced the quantity and quality of song, a condition-dependent, phenotypically plastic signal. Contrary to our expectation, tail elongation had no effect on singing activity and song complexity. However, tail-elongated males produced songs with longer terminal parts ('rattles'). Long rattles are associated with highly competitive social contexts and high circulating levels of testosterone, suggesting that tail-elongated males were more frequently involved in either male-male aggressive or inter-sexual interactions. Therefore, this study shows that males are not displaying multiple signals at the maximum possible level, implying that this system is open to unreliable communication. However, long-term trade-offs between signal expression and viability may prevent males from displaying both signals at higher rates.Communicated by: M. Webster     

Female courtship in the Banggai cardinalfish: honest signals of egg maturity and reproductive output?

Despite the vast literature on male courtship behaviour, little is known about the function and information content of female courtship behaviour. Female courtship behaviour may be important in many species, particularly where both sexes invest heavily in the offspring, and if such behaviours contain honest information regarding a females potential reproductive investment, they may be particularly important in male mate choice. Using observations of two female courtship behaviours (the rush and the twitch) from experimental pairings in the Banggai cardinalfish (Pterapogon kauderni), I addressed the question of whether these courtship behaviours contained information on female reproductive output (clutch weight) and egg maturity (proximity to spawning), traits commonly associated with male mate choice. I especially focused on the importance of these courtship behaviours in relation to other female characters, such as size and condition, using multiple regression. I found that one of these behaviours, the rush, was strongly associated with fecundity, whereas size, condition and the twitch were not. Further, I found that the twitch behaviour was associated with how close to actual spawning a female was. The results suggest that female courtship behaviour may convey highly important information in a mate choice context. I discuss the adaptive value of honest information in female courtship behaviour in light of these results.Communicated by K. Lindström     

Multiple color signals may reveal multiple messages in male Schreiber’s green lizards, <Emphasis Type="Italic">Lacerta schreiberi</Emphasis>

Multiple traits may either signal different characteristics of a male or be redundant. These multiple signals may convey different messages if they are intended for different receivers (e.g., male or females) that have different interests. We examined the functions of multiple colorful visual traits of male Schreiber’s green lizard (Lacerta schreiberi). Results showed that interindividual variation in the characteristics of coloration of males can be related to variation in morphology, health state, dominance status, and pairing status, but that different relationships were found for each color signal. For example, dominant males had brighter “blue” throat and with higher values of ultraviolet (UV) and bluish coloration and darker and greenish dorsal coloration than subordinate males. Health state was also reflected in coloration; males with a higher immune response had “blue” throats with lower amounts of UV coloration, but had “yellow” chests with higher amounts of UV coloration. Males found guarding females also differed in coloration from males found alone. These data suggest that characteristics of coloration of the different multiple signals may reveal different messages for different receivers, either male or female conspecifics. The development of the different signals, based on different morphological and physiological mechanisms and trade-offs, may allow signal reliability of multiple colorful traits in different social contexts.     

Does the presence of non-breeders enhance the fitness of breeders? An experimental analysis in the clown anemonefish<Emphasis Type="Italic"> Amphiprion percula</Emphasis>

The stability of animal societies depends on individuals decisions about whether to tolerate or evict others and about whether to stay or leave. These decisions, in turn, depend on individuals costs and benefits of living in the group. The clown anemonefish, Amphiprion percula, lives in groups composed of a breeding pair and zero to four non-breeders. To determine why breeders accept the presence of non-breeders in this species I investigated the effect of non-breeders on multiple components of the breeders fitness. Non-breeders did not assist breeders in any obvious way. Experimental removal of non-breeders had no significant effect on the survival, growth, or reproductive success of breeders. Experimental removal of one of the breeding pair showed that non-breeders had little effect on the time taken for a widowed breeder to recommence breeding. The results indicate that the presence of non-breeders neither enhances, nor reduces, the fitness of breeders in A. percula. I suggest that non-breeders might modulate their effect on the fitness of breeders, either by reducing the costs they inflict or by increasing the benefits they provide, such that it just pays breeders to tolerate, rather than to evict, them. This study illustrates that animal societies can be stable even when some individuals gain nothing from the association.Communicated by M. Abrahams     

How does burrowing by the isopod <Emphasis Type="Italic">Limnoria agrostisa</Emphasis> (Crustacea: Limnoriidae) affect the leaf canopy of the southern Australian seagrass <Emphasis Type="Italic">Amphibolis griffithii</Emphasis>?

In south-western Australia, the isopod Limnoria agrostisa commonly burrows into leaf clusters and immature shoots of Amphibolis griffithii. The isopod also burrows into the sheath and rhizomes of Posidonia species. In A. griffithii, the isopod consumes new tissue within the sheath, damaging or destroying the meristem. This results in malformation of new leaves or destruction of whole leaf clusters with the potential to reduce the photosynthetic area of a shoot. The isopod has been found in all but one meadow of A. griffithii examined over 1,000 km of the Western Australian coastline. It was present throughout the year and showed little variation in abundance. Young were produced year round, but were more abundant in the summer months. Females, 3.5 mm in length or larger, produced 2–5 young that were brooded within the leaf cluster or base of an immature shoot. Within a meadow, 40–70% of shoots and 10–20% of leaf clusters were damaged by isopods. Seasonal trends were not consistent, but damage appeared to be higher in summer when isopod abundance was higher. Approximately 40% of clusters were destroyed by isopod damage. Isopods attack shoots of all ages, but damage was often located on apical clusters. There was no evidence that isopod damage initiated branching or leaf cluster formation. Estimations of clusters damaged or destroyed may be conservative, as only those clusters remaining on a shoot could be counted, and cluster loss could not be quantified. Examination of defoliated stems on upright shoots and horizontal rhizomes indicated that many were destroyed by isopods. The effect of L. agrostisa although substantial appears to be a feature of healthy seagrasses throughout southern Australia.     

Function of the major cheliped in male–male competition in the hermit crab <Emphasis Type="Italic">Pagurus nigrofascia</Emphasis>

Functions of the major cheliped in pagurid hermit crabs have been studied in fights for shells. The major cheliped often shows sexual size dimorphism, suggesting that sexual selection favors the development of the male major cheliped. The function of the major cheliped in male–male competition was examined in Pagurus nigrofascia collected from April to June 2009 on the intertidal rocky shore in southern Hokkaido, Japan (41°N, 140°E). Sexual size dimorphism of the major cheliped was observed, and precopulatory guarding males had larger major chelipeds than solitary ones. Guarding males used the major cheliped to deter intruders during competitive interactions. Males without a major cheliped were disadvantaged even if they were larger than opponents and had ownership. Cheliped size affected the outcomes of contests between similar sized males. This suggests that the male major cheliped in P. nigrofascia protects mates from competitors and, consequently, enhances male mating success. Sexual selection may favor the development of the major cheliped in male pagurids.     

Dance precision of <Emphasis Type="Italic">Apis florea</Emphasis>—clues to the evolution of the honeybee dance language?

All honeybee species make use of the waggle dance to communicate the direction and distance to both food sources and potential new nest sites. When foraging, all species face an identical problem: conveying information about profitable floral patches. However, profound differences in nesting biology (some nest in cavities while others nest in the open, often on a branch or a cliff face) may mean that species have different requirements when dancing to advertise new nest sites. In cavity nesting species, nest sites are a precise location in the landscape: usually a small opening leading to a cavity in a hollow tree. Dances for cavities therefore need to be as precise as possible. In contrast, when the potential nest site comprises a tree or perhaps seven a patch of trees, precision is less necessary. Similarly, when a food patch is advertised, dances need not be very precise, as floral patches are often large, unless they are so far away that recruits need more precise information to be able to locate them. In this paper, we study the dance precision of the open-nesting red dwarf bee Apis florea. By comparing the precision of dances for food sources and nest sites, we show that A. florea workers dance with the same imprecision irrespective of context. This is in sharp contrast with the cavity-nesting Apis mellifera that increases the precision of its dance when advertising a potential new home. We suggest that our results are in accordance with the hypothesis that the honeybees’ dance communication initially evolved to convey information about new nest sites and was only later adapted for the context of foraging.     

Does the antidiabetic drug metformin affect embryo development and the health of brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis> f. <Emphasis Type="Italic">fario</Emphasis>)?

Background

Due to the rising number of type 2 diabetes patients, the antidiabetic drug, metformin is currently among those pharmaceuticals with the highest consumption rates worldwide. Via sewage-treatment plants, metformin enters surface waters where it is frequently detected in low concentrations (µg/L). Since possible adverse effects of this substance in aquatic organisms have been insufficiently explored to date, the aim of this study was to investigate the impact of metformin on health and development in brown trout (Salmo trutta f. fario) and its microbiome.

Results

Brown trout embryos were exposed to 0, 1, 10, 100 and 1000 µg/L metformin over a period from 48 days post fertilisation (dpf) until 8 weeks post-yolk sac consumption at 7 °C (156 dpf) and 11 °C (143 dpf). Chemical analyses in tissues of exposed fish showed the concentration-dependent presence of metformin in the larvae. Mortality, embryonic development, body length, liver tissue integrity, stress protein levels and swimming behaviour were not influenced. However, compared to the controls, the amount of hepatic glycogen was higher in larvae exposed to metformin, especially in fish exposed to the lowest metformin concentration of 1 µg/L, which is environmentally relevant. At higher metformin concentrations, the glycogen content in the liver showed a high variability, especially for larvae exposed to 1000 µg/L metformin. Furthermore, the body weight of fish exposed to 10 and 100 µg/L metformin at 7 °C and to 1 µg/L metformin at 11 °C was decreased compared with the respective controls. The results of the microbiome analyses indicated a shift in the bacteria distribution in fish exposed to 1 and 10 µg/L metformin at 7 °C and to 100 µg/L metformin at 11 °C, leading to an increase of Proteobacteria and a reduction of Firmicutes and Actinobacteria.

Conclusions

Overall, weight reduction and the increased glycogen content belong to the described pharmaceutical effects of the drug in humans, but this study showed that they also occur in brown trout larvae. The impact of a shift in the intestinal microbiome caused by metformin on the immune system and vitality of the host organism should be the subject of further research before assessing the environmental relevance of the pharmaceutical.

     

Where are larvae of the vermetid gastropod <Emphasis Type="Italic">Dendropoma maximum</Emphasis> on the continuum of larval nutritional strategies?

This study evaluated whether larvae of the Indo-Pacific vermetid gastropod Dendropoma maximum are obligate planktotrophs, or whether they exhibit an intermediate feeding strategy. Experiments were conducted in Moorea, French Polynesia (149°50′W, 17°30′S), Sep–Oct 2009, to examine D. maximum larval growth and metamorphic responses to different diets and amounts of food. Dendropoma maximum larvae required particulate food to undergo metamorphosis, but were able to survive and grow in the absence of food for up to 20 days. Larvae in Low and Unfed food treatments exhibited phenotypic plasticity by growing a larger velum (the larval feeding structure) compared with those in high food. Unfed D. maximum larvae had a slower initial growth rate; however, by 11-day post-hatch fed and unfed larvae had converged on the same mean shell height (553 μm), which was only 10% larger than the initial size at hatching. Therefore, although the nutritional strategy of D. maximum larvae is best described as obligate planktotrophy, it appears to approach an intermediate feeding strategy.     

Why are larvae of the social parasite wasp <Emphasis Type="Italic">Polistes sulcifer</Emphasis> not removed from the host nest?

A challenge for parasites is how to evade the sophisticated detection and rejection abilities of potential hosts. Many studies have shown how insect social parasites overcome host recognition systems and successfully enter host colonies. However, once a social parasite has successfully usurped an alien nest, its brood still face the challenge of avoiding host recognition. How immature stages of parasites fool the hosts has been little studied in social insects, though this has been deeply investigated in birds. We look at how larvae of the paper wasp obligate social parasite Polistes sulcifer fool their hosts. We focus on cuticular hydrocarbons (CHCs), which are keys for adult recognition, and use behavioral recognition assays. Parasite larvae might camouflage themselves either by underproducing CHCs (odorless hypothesis) or by acquiring a chemical profile that matches that of their hosts. GC/MS analyses show that parasite larvae do not have lower levels of CHCs and that their CHCs profile is similar to the host larval profile but shows a reduced colony specificity. Behavioral tests show that the hosts discriminate against alien conspecific larvae from different colonies but are more tolerant towards parasite larvae. Our results demonstrate that parasite larvae have evolved a host larval profile, which overcomes the host colony recognition system probably because of the lower proportion of branched compounds compared to host larvae. In some ways, this is a similar hypothesis to the odorless hypothesis, but it assumes that the parasite larvae are covered by a chemical blend that is not meaningful to the host.     

Optimal maternal investment in the dung beetle<Emphasis Type="Italic"> Onthophagus taurus</Emphasis>?

Optimal investment theory is based on the assumption that the proximate constraint acting on parental investment is resource based. A trade-off between per offspring investment and total investment seems intuitive. Consequently, a parents investment strategy is expected to represent a trade-off between the benefits of investment for current offspring and the costs to future reproduction for parents. In this study, we provide clear evidence that the costs and benefits of maternal provisioning in the dung beetle Onthophagus taurus influence the amount of provisions provided by the mother. Horse dung is typically of a higher nutritional value than cow dung and females were shown to provide 20% less dung to offspring when provisioning with horse dung. By reducing their investment per offspring and exhibiting a clear preference to provision offspring with horse dung, females were able to produce significantly more offspring. Females provisioning with horse dung received greater fitness returns per unit of investment and experienced lower provisioning costs, in terms of the minimum amount of dung required to produce a surviving offspring, than females provisioning with cow dung. Females provisioning in soil of low moisture content were found to have higher tunneling costs than those provisioning in soil of high moisture content, while the fitness returns per unit of investment did not differ. We adopted a marginal value theorem (MVT) approach to calculate the theoretical optimal level of investment for each dung type and for each soil moisture. Predicted levels of provisioning were lower for horse dung than for cow dung and for moist soil than for dry soil. Therefore, the results of this study are in qualitative agreement with MVT predictions and provide empirical support for the proposal that females can adaptively adjust their level of investment in response to resource and/or habitat quality. However, the theoretically predicted optimal investment yielded a poor quantitative fit with our observed levels of investment, with females providing over twice the investment predicted by the MVT approach. We suggest that this difference may reflect either our inability in directly quantifying all the necessary costs and benefits of investment in O. taurus and/or the applicability of the underlying assumptions of MVT.Communicated by D. Gwynne     

Sibling species or poecilogony in the polychaete<Emphasis Type="Italic"> Scoloplos armiger</Emphasis>?

In marine invertebrates multiple modes of development, or poecilogony, may occur in a single species. However, after close examination, many of such putative cases turned out to be sibling species. A case in point may be the cosmopolitan orbiniid polychaete Scoloplos armiger, which inhabits marine shallow sediments. In addition to the well-known direct, holobenthic development from egg cocoons, pelagic larvae have also been described. Our culture experiments revealed a spatially segregated source of the two developmental modes. All females of an intertidal population produced egg cocoons and no pelagic larvae. All but 2 out of 15 females of an adjacent subtidal population produced pelagic larvae and no egg cocoons. Based on these results we performed a molecular genetic analysis (RAPD-PCR) on three intertidal and four subtidal populations in the North Sea. Selected samples from all sites were analysed also by the AFLP method. We found significantly higher genetic diversity within subtidal than within intertidal populations. This is consistent with a wider dispersal by pelagic larvae and a smaller effective population size when development is holobenthic. Total genetic divergence is not related to distance but to the intertidal/subtidal division. We suggest that S. armiger actually represents two sibling species.     

Strategies of a parasite of the ant–<Emphasis Type="Italic">Acacia</Emphasis> mutualism

Mutualisms can be exploited by parasites—species that obtain resources from a partner but provide no services. Though the stability of mutualisms in the presence of such parasites is under intensive investigation, we have little information on life history traits that allow a species to be a successful mutualist or rather a parasite, particularly in cases where both are closely related. We studied the exploitation of Acacia myrmecophytes by the ant, Pseudomyrmex gracilis, contrasting with the mutualistic ant Pseudomyrmex ferrugineus. P. gracilis showed no host-defending behavior and had a negative effect on plant growth. By preventing the mutualist from colonization, P. gracilis imposes opportunity costs on the host plant. P. gracilis produced smaller colonies with a higher proportion of alates than did the mutualist and thus showed an “r-like” strategy. This appears to be possible because P. gracilis relies less on host-derived food resources than does the mutualist, as shown by behavioral and stable isotope studies. We discuss how this system allows the identification of strategies that characterize parasites of mutualisms.     

What did Lotka really say? A critical reassessment of the “maximum power principle”

This paper presents a critical discussion of the so-called “maximum power principle”, often quoted in studies about the energy balance of living systems and also known in the emergy literature as “maximum em-power principle”. Several authors consider this principle highly relevant and some even proposed it as a “fourth law of thermodynamics”. A critical analysis of the original source, namely Alfred Lotka's 1921-22 papers, conducted both in an historical perspective (the connection between Lotka's writings and the ongoing debate at his time) and in a more modern context, leads to a more detailed and less biased assessment. It turns out that in spite of Lotka's very anticipatory and incredibly sharp vision of the possible interconnections between the second law of thermodynamics and evolutionism, doubts arise about the general applicability of his “maximum power principle”. From an accurate analysis of his writings, it can be concluded that: (a) Lotka explicitly and consistently addressed the “optimal use” of the flow of exergy (available energy), and therefore the quantity defined as “em-power” is an incorrect interpretation of Lotka's constrained maximum power principle; (b) “Lotka's principle” can be reformulated within Ziegler's “maximum entropy production” or Prigogine “minimum entropy generation” paradigm only under two different respective sets of rather stringent additional conditions which Lotka was probably already aware of but never explicitly stated.     

Respiration and lipid content of the Arctic copepod<Emphasis Type="Italic"> Calanus hyperboreus</Emphasis> overwintering 1 m above the seafloor at 2,300 m water depth in the Fram Strait

In August 2000 high concentrations of the dominant herbivorous copepod Calanus hyperboreus were detected in the Arctic Fram Strait, west of Spitsbergen, 1 m above the seafloor at 2,290 m water depth. Individuals from that layer were sampled by a hyper-benthic net attached to the frame of an epi-benthic sledge. For comparison, the vertical distribution of C. hyperboreus in the water column was studied simultaneously by a multiple opening/closing net haul from 2,250 m depth to the surface. Maximum abundance was found close to the surface with 6.6 and 10.0 ind. m^?3 at 0–50 m and 50–100 m depth, respectively. However, the major fraction of the population (>40%) occurred between 1,000 and 1,500 m depth. In the deepest layer (2,000–2,250 m) abundance measured 2.2 ind. m^?3 and was twice as high as between 100 and 1,000 m depth. In comparison to individuals from surface waters, copepods from the hyper-benthic layer were torpid and did not react to mechanical stimuli. Stage CV copepodids and females from the deep sample contained 4–10% less lipid and showed significantly reduced respiration rates of 0.24 and 0.26 ml O₂ h^?1 g^?1 dry mass (DM) as compared to surface samples (0.49 and 0.43 ml O₂ h^?1 g^?1 DM). All these observations indicate that the hyper-benthic part of the population had already started a dormant overwintering phase at great depth. Based on the lipid deposits and energy demands, the potential maximum duration of the non-feeding dormant phase was estimated at 76–110 days for females and at 98–137 days for CV copepodids, depending on what indispensable minimum lipid content was assumed. In any case, the estimated times could not meet the necessary requirements for a starvation period of >6 months until the next phytoplankton bloom in the following spring. The ecological implications of these results are discussed with respect to the life cycle and eco-physiological adaptations of C. hyperboreus to its high-Arctic habitat.     

TiO<Subscript><Emphasis Type="Bold">2</Emphasis></Subscript> nanoparticles – Relationship between dispersion preparation method and ecotoxicity in the algal growth test

PurposeLittle is known about the ecotoxicity of nanomaterials and there are no specific guidelines for sample preparation and testing. We set out to establish whether the method used to prepare TiO₂ dispersions had a significant impact on aquatic ecotoxicity. We also followed the formation of agglomerates during the incubation period.MethodsWe applied the algal growth inhibition test (OECD test guideline no. 201). Dispersions were prepared by stirring and/or ultrasonication for different durations, and by filtration according to an OECD procedure recommended for testing difficult substances.ResultsSamples stirred for 7?d were not toxic, but EC₂₀ values could be calculated for all the other treatments. Shorter treatments generated EC₂₀ values in the range 1–27?mg/L. Only the shortest treatment (1 min stirring, 1 min ultrasonication) produced an unusually high EC₂₀ value, indicating low toxicity. Development of agglomerate size and of toxicity depends on the nanoparticles. We found that ecotoxicity was predominantly caused by a fraction of nanoparticles and agglomerates obtained by passing dispersions through a 0.22-µm filter.ConclusionsWe propose a short treatment regime to generate the most relevant ecotoxicity data for TiO₂, for example stirring for 1?min followed by 3 min ultrasonication. Until more data concerning the ecotoxicity of different fractions are available, we recommend the testing of unfiltered dispersions rather than filtrates. Relating ecotoxicity to the total hydrodynamic surface of the nanomaterials rather than concentration does not seem to improve the accuracy of ecotoxicity assessments using the algal growth inhibition test.     

Multiple lineages and absence of panmixia in the “circumpolar” crinoid <Emphasis Type="Italic">Promachocrinus kerguelensis</Emphasis> from the Atlantic sector of Antarctica

Despite considerable interest in physiology, evolution and life history of Antarctic marine invertebrates, only a limited number of studies have examined the genetic variability and diversity patterns of these organisms. Moreover, understanding and characterizing patterns of Antarctic biodiversity has taken on a degree of urgency because of potential impacts of global warming. To expand an understanding of the evolutionary history of Antarctic marine invertebrates, the genetic diversity of the crinoid Promachocrinuskerguelensis Carpenter, 1888 was investigated, which is documented to have a circumpolar distribution extending to subantarctic islands. Specimens of P. kerguelensis were collected from the western side of the Antarctic Peninsula, and the subantarctic islands South Georgia, South Sandwich and Bouvetøya Island from 2001 to 2004. P. kerguelensis was previously subject to morphological review that confirmed the taxonomic recognition of only one species. The wide distribution and reported high dispersal capability for P. kerguelensis predicts one large panmictic population. In contrast, nucleotide sequence data from mitochondrial cytochrome oxidase subunit I and cytochrome b genes, collected herein, reveal distinct genetic structure and cryptic speciation within P.kerguelensis. In the Antarctic Atlantic sector alone, there were at least five “species-level” clades. Some of these clades are geographically limited, and most exist in sympatry. The largest and most widespread of these clades was examined to help elucidate connectivity along the subantarctic islands of the Scotia Arc and the Antarctic Peninsula. Within this clade, most genetic diversity was contained within populations, but significant differences were present between regions (Antarctic Peninsula, South Sandwich Is., South Georgia, Bouvetøya Is.), suggesting a corresponding lack of gene flow. Given that P. “kerguelensis” is a well-studied taxon, the finding of considerable genetic diversity within the Atlantic sector alone suggests that the recognized diversity of Antarctica’s benthic marine life may be underestimated, and will rise dramatically with phylogeographic analyses of putative widespread species.     

Rearing<Emphasis Type="Italic"> Pandalus borealis</Emphasis> (Krøyer) larvae in the laboratory

Northern shrimp Pandalus borealis (Krøyer) larvae hatch in the northern Gulf of St. Lawrence from early May to the end of June, and larval development occurs over a range of relatively cold water temperatures. Because of the long duration of the pelagic phase and the difficulty of sampling all successive larval stages at sea, we used laboratory experiments to assess the effects of water temperature on larval development and growth. In spring 2000, P. borealis larvae were reared from hatching to the first juvenile stages (i.e., stage VI and VII) at three temperatures (3, 5, and 8°C) representing conditions similar to those in spring in the northern Gulf of St. Lawrence. Larval development and growth were dependent on temperature, with longer duration and smaller size (cephalothorax length, CL, and dry mass, DM) at 3°C relative to the 5 and 8°C treatments. There were no significant differences in the morphological characters of the different stages among treatments, indicating that regular moults occurred at each temperature. The results suggest a negative impact of cold temperatures (lower intra-moult growth rates and smaller size) and, possibly, higher cumulative mortality due to longer development time that could affect the success of cohorts at sea. However, CL and DM for stage III and later larvae were smaller than those of larvae identified at the same developmental stage in field locations. It is possible that the diet offered to larvae in this experiment (Artemia nauplii, either newly hatched nauplii or live adults, depending on the developmental stage) was not optimal for growth, even though it is known to support successful P. borealis larval development. In the field, there is the possibility that phytoplankton contributes to the larval diet during the first stages and stimulates development of the digestive glands. Furthermore, the nutritional quality of the natural plankton diet (e.g., high protein content, fatty acid composition) might be superior and favourable to higher growth rates even at lower temperatures.Communicated by R.J. Thompson, St. Johns     

Kin discriminators in the eusocial sweat bee <Emphasis Type="Italic">Lasioglossum malachurum</Emphasis>: the reliability of cuticular and Dufour’s gland odours

The ability to discriminate degrees of relatedness may be expected to evolve if it allows unreciprocated altruism to be preferentially directed towards kin (Hamilton in J Theor Biol 7:1–16, 1964). We explored the possibility of kin recognition in the primitively eusocial halictid bee Lasioglossum malachurum by investigating the reliability of worker odour cues that can be perceived by workers to act as indicators of either nest membership or kinship. Cuticular and Dufour’s gland compounds varied significantly among colonies of L. malachurum, providing the potential for nestmate discrimination. A significant, though weak, negative correlation between chemical distance and genetic relatedness (r = −0.055, p < 0.001) suggests a genetic component to variation in cuticular bouquet, but odour cues were not informative enough to discriminate between different degrees of relatedness within nests. This pattern of variation was similar for Dufour’s gland bouquets. The presence of unrelated individuals within nests that are not chemically different from their nestmates suggests that the discrimination system of L. malachurum is prone to acceptance errors. Compounds produced by colony members are likely combined to generate a gestalt colony chemical signature such that all nestmates have a similar smell. The correlation between odour cues and nest membership was greater for perceived compounds than for non-perceived compounds, suggesting that variability in perceived compounds is a result of positive selection for nestmate recognition despite potentially stabilising selection to reduce variability in odour differences and thereby to reduce costs derived from excessive intracolony nepotistic behaviour.