The effects of an extraordinary El Niño / La Niña event on the size and growth of the squid<Emphasis Type="Italic"> Loligo opalescens</Emphasis> off Southern California

The population structure of the California market squid Loligo opalescens was studied for the Channel Islands region off Southern California between June 1998 and March 2000. During this time Californian waters were exposed to an extraordinary El Niño event that was possibly the most dramatic change in oceanographic conditions that occurred last century. There was then a rapid transition to record cool La Niña conditions. Statolith increments were used to determine age parameters and increment periodicity was validated for the first 54 days of life. Based on statolith increment counts, the oldest males and females were 257 and 225 days respectively and individuals matured as young as 129 and 137 days respectively. No distinct hatching period was detected. There was a general trend of increasing body size throughout the study period. Squid that hatched and grew through the El Niño were strikingly smaller and had slower growth rates compared to squid that grew through the La Niña. This was related to oceanography and associated productivity. There was a positive correlation between squid mantle length and upwelling index and a negative correlation between mantle length and sea temperature. The 'live-fast die-young' life history strategy of squid makes them ideal candidates for following the effects of the dramatic changes in oceanographic conditions off California. We propose that squid can serve as ecosystem recorders and productivity integrators over time and space and are useful organisms to tie oceanography to biology.     

El Niño periods increase growth of juvenile white seabass (<Emphasis Type="Italic">Atractoscion nobilis</Emphasis>) in the Southern California Bight

Studies of the impact of El Niño periods on marine species have usually focused on negative, highly visible effects, e.g., decreasing growth rates or increasing mortality due to a decline in primary productivity in typically nutrient rich upwelling zones; but positive effects related to elevated water temperature are also known. This study examined how the growth rate of juvenile white seabass, Atractoscion nobilis, responded to changes in ocean temperature in an El Niño period (1997–1998) in the northern portion of the Southern California Bight, USA. Growth rates of juvenile white seabass during their first 4 years of life were estimated as the slopes of linear relationships between body mass and age (from otoliths) of 800 fish collected at 11 stations throughout the bight. Growth rates differed significantly among cohorts hatched in 1996–2001. Specifically, white seabass that hatched in 1996 and 1997 grew significantly faster than those that hatched in 1998, 1999, and 2001. These differences in growth rates of cohorts appeared to be driven by variation in sea-surface temperature (SST). Growth rates averaged over the first three or 4 years of life were significantly positively correlated to average daily SST during the first 1–4 years of life. Increased growth of juvenile white seabass during the warm El Niño period likely provided a number of benefits to this warm-temperate species. This study demonstrated that some species will benefit from these warm-water periods despite reduced system-wide primary production.     

Covering behaviour in <Emphasis Type="Italic">Paracentrotus</Emphasis><Emphasis Type="Italic">lividus</Emphasis>: is light important?

"Covering" or "heaping" behaviour is common to a number of regular echinoids living in a variety of different habitats. Many theories have been proposed to explain this behaviour, among which are several that link covering to light, including ultraviolet (UV) light. However, previous investigations of this light theory have been largely qualitative. In the present study, we used a systematic laboratory protocol to examine quantitatively the covering behaviour of the shallow water echinoid, Paracentrotus lividus, under four different light regimes: white light (400-700 nm), UV-A+B (315-400 nm), UV-A (320-400 nm), and darkness. These experiments demonstrated that light, in particular UV light, influences the covering behaviour of P. lividus. Under the UV regimes, significantly more individuals were found to display covering behaviour, and individuals spent more time at the base of aquaria, farthest from the light source. Moreover, covering items were retained for the longest period of time under the UV-A+B regime. We propose that protection against the harmful effects of UV radiation may be one of the functions of covering behaviour in P. lividus.     

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.     

Simulation of exchange flow between open water and floating vegetation using a modified RNG <Emphasis Type="Italic">k</Emphasis>-<Emphasis Type="Italic">ε</Emphasis> turbulence model

The paper focuses on the numerical simulation of the exchange flow between open water and floating vegetation, which plays an important role in maintaining the ecological balance by transporting nutrient matter. The simulation was conducted using a new solver developed upon OpenFOAM. A modified RNG k-ε turbulence model, which is expected to model both the high- and low-Reynolds number flows correctly, was used to determine the eddy viscosity. Several particular terms were added into the momentum equations and turbulence model equations to model the effects of vegetation and buoyancy. Among these terms, the term for the effect of vegetation in the ε-equation was re-modelled. The model was validated by properly predicting the profiles of mean velocity and turbulent kinetic energy for flows through suspended canopies. The density flow between open and vegetated water was simulated with the same conditions as those of the experiment conducted by Zhang and Nepf. The predicted results agreed well with the experimental data and provided more detailed information of such exchange flow. The convection between the root layer and the layer beneath the roots, which was not observed in the experiment, was observed in the numerical simulation.     

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.     

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     

Laboratory investigation of <Emphasis Type="Italic">Fallopia</Emphasis> × <Emphasis Type="Italic">bohemica</Emphasis> fruits dispersal by watercourses

Seed and fruit dispersal along watercourses favours the long-distance migration of invasive species, not only for aquatic or wetland species, but also for terrestrial wind-dispersed plants, like the Japanese knotweed. The present paper aims at investigating the role of watercourses in the dispersal of the knotweed due to its frequent occurrence on riverbanks and production of fertile achenes (type of fruit of the Japanese knotweed). This dispersal occurs along two steps after the fruits deposit on the water surface: floatation first and then sinking towards the bottom of the watercourse. Regarding the first step, the effects of agitation of the water, temperature, surface tension and luminosity on the achenes floatability are experimentally studied. While no influence of luminosity is observed, an increase of temperature greatly decreases the floating time. Floating time also decreases as the contact between water and the fruit is enhanced (through submersion of achenes, agitation of the water or lower surface tension). Regarding the second step, the fall velocity of the fruits in water at rest is measured and appears to be independent of the seed history (floating time). 3D helical motions are systematically observed with constant tangential velocity with respect to the falling velocity. The trajectory of the fruits in a shear flow is then measured and the evolution of their velocity components along the sinking process is discussed. Finally, the contribution of both steps to the long-distance migration of the seeds is estimated.     

Genetic variation among life-history stages of mussels in a<Emphasis Type="Italic"> Mytilus edulis–M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis> hybrid zone

The mussel species Mytilus edulis L. and M. trossulus Gould coexist and hybridize throughout a large area that includes the north coast of Maine and Atlantic Canada. Previous studies provided genetic evidence for limited hybridization between the two species for mussels >15 mm. The present study used two genetic markers (ITS, Glu-5) to examine the genetic composition of early life-history stages by sampling veliger and pediveliger larvae, juveniles (<2.0 to 15.0 mm shell length) and adults (>15 mm shell length) in Trinity Bay, Newfoundland, during three consecutive years (1995–1997) to determine if differential mortality limits the relative abundance of hybrids. The relative frequency of the two species and the different hybrid genotypes was similar among the larvae, juveniles and small adult mussels. The double hybrid genotype (F₁-like) was the rarest genotype observed. There was no evidence for differential mortality during the early life-history stages, and factors limiting production of hybrids appear to operate before the late larval stage. The observed frequency of hybrids is probably due to a combination of pre- and postzygotic reproductive isolating mechanisms operating early in the mussels life history. M. trossulus dominated the early life-history stages, possibly due to a higher population density and a greater reproductive output than M. edulis. Differential mortality may explain the observed decrease in frequency of M. trossulus and increase in frequency of M. edulis with increasing shell length. A similar frequency of hybrid mussels from larvae to the size class of 55 mm shell length may indicate a rate of mortality intermediate between the two parental species. The M. edulis–M. trossulus hybrid zone appears to be maintained by reproductive isolating mechanisms limiting the production of hybrids and life-history differences that allow the two species to coexist.Communicated by J.P. Grassle, New Brunswick     

Variation in colony geometry modulates internal light levels in branching corals, <Emphasis Type="Italic">Acropora humilis</Emphasis> and <Emphasis Type="Italic">Stylophora pistillata</Emphasis>

Colonial photosynthetic marine organisms often exhibit morphological phenotypic plasticity. Where such plasticity leads to an improved balance between rates of photosynthesis and maintenance costs, it is likely to have adaptive significance. To explore whether such phenotypic plasticity leads to more favourable within-colony irradiance for reef-building branching corals, this relationship was investigated for two coral species Acropora humilis and Stylophora pistillata, along a depth gradient representing light habitats ranging from 500 to 25 μmol photons m⁻² s⁻¹, during 2006 at Heron Island, Great Barrier Reef (23.44°S, 151.91°E). In the present study changes in flow-modulated mass transfer co-varied with light as a function of depth. In low-light (deep) habitats, branch spacing (colony openness) in A. humilis and S. pistillata was 40–50% greater than for conspecifics in high-light environments. Also, branches of A. humilis in deep water were 40–60% shorter than in shallow water. Phenotypic changes in these two variables lead to steeper within-colony light attenuation resulting in 38% higher mean internal irradiance (at the tissue surface) in deep colonies compared to shallow colonies. The pattern of branch spacing was similar for S. pistillata, but this species displayed an alternate strategy with respect to branch length: shade adapted deep and cave colonies developed longer and thinner branches, allowing access to higher mass transfer and irradiance. Corals in cave habitats allowed 20% more irradiance compared to colonies found in the deep, and had a 47% greater proportion of irradiance compared to colonies in the shallow high-light environment. Such phenotypic regulation of internal light levels on branch surfaces partly explains the broad light niches of many branching coral species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Life cycle of <Emphasis Type="Italic">Oithona</Emphasis> <Emphasis Type="Italic">similis</Emphasis> (Copepoda: Cyclopoida) in Kola Bay (Barents Sea)

The annual population dynamics (nauplii, old copepodites CIV–CV and adults) and seasonal variations in reproductive parameters of the cyclopoid copepod Oithona similis were investigated on the basis of the data 1999–2006 in Kola Bay, a large subarctic fjord in the Barents Sea. Population density of O. similis ranged from 110 to 9,630 ind m⁻³ and averaged 1,020 ± 336 ind m⁻³. The relative abundance of adults was high during winter (~60%). At the end of winter (mid-March), the population included a large percentage of later-stage copepodites (stage CIV 23% and stage CV 57%). There were two periods of mass spawning, in late June and September. Autumn and summer generations strongly differed in abundance, average prosome length (PL), clutch size (CS), egg diameter (D), egg production rates (EPR and SEPR) and female secondary production. Average PL decreased with increasing water temperature, while D and CS were strongly correlated with PL but unaffected by temperature. Annual average EPR and SEPR were 0.55 ± 0.18 eggs female⁻¹ day⁻¹ and 0.0011 ± 0.003 day⁻¹, respectively. Female secondary production averaged 0.8 ± 0.3 μg C m⁻³ day⁻¹ (range 0.001–3.58). There were positive relationships between abundance, EPR, SEPR, production and water temperatures. Reproductive parameters appeared to be controlled by hydrological factors and food conditions.     

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     

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.

     

Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Differential interests between the sexes regarding the number of copulations can result in sexual harassment. Hence, females may have less time available for foraging. Male sexual harassment often leads to fitness reduction in females. We used the mating complex of the bisexual fish Poecilia mexicana and the co-occurring all-female Poecilia formosa to study sexual harassment and its incurred cost on female feeding efficiency. P. formosa is a sperm-dependent parthenogen that requires mating with host males to induce embryogenesis, but the male genes are not used. We therefore predicted P. mexicana males to prefer conspecific females. Hence, costs of male sexual harassment should not occur in unisexuals. While P. formosa are at a disadvantage compared to P. mexicana females due to male mate choice (leading to sperm limitation), this could be traded-off by suffering less from sexual harassment. In our experiment, we found males to direct significantly more pre-copulatory mating behaviour towards conspecific females, whereas actual mating attempts did not differ between species. Contrary to our prediction, both types of females started feeding later and spent less time feeding in the presence of a male partner compared to the time spent feeding with another female, suggesting that females of both species suffer from male harassment. The focal females' feeding time declined with increasing body size of the female competitor, and the same pattern was found when a male was present. We discuss that—besides sexual harassment—other factors such as food competition and female mate choice may affect female feeding efficiency.     

Do stress hormones suppress helper reproduction in the cooperatively breeding red-cockaded woodpecker (<Emphasis Type="Italic">Picoides borealis</Emphasis>)?

In the cooperatively breeding red-cockaded woodpecker (Picoides borealis), male helpers are subordinate to male breeders and do not mate with females, even when unrelated to the breeding female within their group or through extra-group matings, yet exhibit reproductive hormone profiles similar to those of breeders. We investigated whether reproduction might be suppressed in helper males via high levels of the stress hormone corticosterone. We also examined effects of group size and season on corticosterone levels by comparing baseline and maximal plasma levels of corticosterone between helper males and breeding males, and among helper males and breeders of both sexes living in groups of different sizes throughout the reproductive cycle. We also measured plasma levels of estrogen, progesterone, and testosterone to examine other potential hormonal differences between helpers and breeders. Male status did not explain variation in any hormones; therefore, our data do not support the hypothesis that helper males are reproductively suppressed via corticosterone or the other hormones investigated. However, the presence of two or more helper males in a group tended to reduce baseline corticosterone in breeding and helper males, but not breeding females, suggesting that helper males reduce parental effort of other male group members. Seasonally, maximal corticosterone peaked during the nestling provisioning phase for breeding and helper males, but not breeder females, suggesting that males show an increased response to stressors posing a potential threat to survival of offspring.     

Extrapair paternity in the swamp sparrow, <Emphasis Type="Italic">Melospiza georgiana</Emphasis>: male access or female preference?

Over the past two decades, the combination of molecular and field methods has revealed considerable variation in the level of extrapair fertilizations among socially monogamous birds. Models predicting extrapair young range in scale from a single population to multiple Orders, and there is no single, unifying theory for these reproductive tactics. We investigated proximate explanations of extrapair fertilizations in two subspecies of the swamp sparrow, Melospiza georgiana georgiana and Melospiza georgiana nigrescens, across a range of social and environmental conditions. The presence of extrapair young was best predicted by the size of two male plumage badges (one correlated with parental care and one with territorial aggression) relative to the badge size of their immediate neighbors, the interaction of these two measures, mean territory size, and the maximum size of the aggression badge among neighbors. The size of the male’s parental care badge (relative to neighbors) was negatively correlated with the probability of lost paternity. The relative size of the aggression badge was positively correlated with the presence of extrapair young when the parental care badge was small and negatively correlated when the badge was large. Controlling for these crown measures, males with larger territories were less likely to suffer losses in paternity. There was no effect of breeding density, breeding synchrony, their interaction, subspecies, or weather during the fertile period on the presence of extrapair young. These results suggest that female preference for males that provide more parental care (or preference for genes that convey this trait) plays a dominant role in extrapair interactions among swamp sparrows. Models based on female assessments of relative mate quality offer a promising explanation of patterns in extrapair fertilizations among bird species.     

Mating behavior of <Emphasis Type="Italic">Abdopus aculeatus</Emphasis> (d’Orbigny 1834) (Cephalopoda: Octopodidae) in the wild

The mating system of Abdopus aculeatus incorporates sneaker matings, mate guarding, sex-specific body patterns, frequent copulations, and male–male competition for mates, making it more similar to that of aggregating decapod cephalopods than any previously known octopus social system. Large male–female A. aculeatus occupy ‘Adjacent’ (G_A) dens and copulate frequently in mate-guarding situations over successive days. Nearby individuals copulate in ‘Temporary guarding’ (G_T) and ‘Transient’ (T; non-guarding) situations, the latter of which can involve ‘Sneaker’ (S) mating. In a focal animal study of these octopuses in the wild (Sulawesi, Indonesia) we addressed the hypotheses that they demonstrate: (1) precopulatory mate choice, (2) differential copulation rates by individuals employing different mating tactics, and (3) distant sex identification. We quantified daily copulation rates of A. aculeatus of reproductive size as well as aspects of copulation duration, display, mate-competition, and mate rejection. Mating tactic correlated with daily copulation rates. ♂G_A spent significantly more time copulating than did ♂T, while ♀G_A spent more than twice as much time per day in copula than did other females. Sneaker copulations lasted longer than those by males adopting other tactics. Mate-guarding was an effective and important tactic used by males to temporarily monopolize mating with apparently non-selective females. Males demonstrated clear pre-copulatory mate choice by guarding and mating repeatedly with large females (typically ♀G_A). While foraging alone away from the den, ♂G procured ‘Transient’ copulations with unguarded females. However, mate-guarding reduced the amount of time ♂G were alone and may impede their ability to seek out new mates. Low-copulation rates by ♀T, the smallest female tactic on average, may reflect this trade-off between mate preference and mate-searching by males, or non-receptivity of some females. A male-typical body pattern (black and white stripes) appeared to facilitate distant sex identification. Although mating and aggression were often initiated before contact between individuals, same-sex copulations and intense male–female aggression were rare. By contrast frequent male–female copulations and intense male–male aggression were consistent behavioral components of mating in A. aculeatus at these sites. Because the behavioral and ecological characters conducive to this complex system are not exclusive to A. aculeatus, it is possible that other octopuses exhibit some or all of these behaviors. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Reproduction in the deepwater squid<Emphasis Type="Italic"> Moroteuthis ingens</Emphasis>, what does it cost?

The cost of reproduction for the terminal spawning onychoteuthid squid, Moroteuthis ingens, was analysed using measures of condition and tissue biochemistry. Both males and females showed a dramatic drop in the weight of the gonad in stage 6 (spent) individuals. The mantle weight and nidamental gland weight of females also decreased during the maturation process. Males, however, had a marked increase in both the penis and spermatophoric complex weight in spent individuals, while female oviducal gland weight and nidamental gland length also increased in stage 6 individuals. Residual analysis indicated that testis growth was not developing at the expense of mantle growth, although there was a suggestion of cost to the fins. Females showed that the development of the ovary occurred at a cost to both the mantle and fins. Overall body condition also declined with maturity stage for both males and females, with stage 6 individuals of both sexes in poor condition. Very few females had eggs in the oviducts, suggesting that the oviducts are used as ducts instead of storage organs. Proximal analysis revealed a loss of constituents within the mantle during maturation, with an associated increase in water, indicating the remobilization of energy from the mantle to fuel reproduction. This study suggests that the digestive gland is not used as an energy store in this species.Communicated by M.S. Johnson, Crawley     

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.