Growth and longevity of the mud snail Batillaria attramentaria

Crab pots were used to sample a population of Scylla serrata (Forskal) in an estuarine area in Queensland, Australia. Pots were laid 100 m apart at fixed positions for 4 d each month for 1 yr (April 1980–June 1981). Data from recapture of tagged crabs showed that males larger than 140 mm carapace width and females larger than 150 mm had a higher capture probability than did smaller crabs; thus size-frequency distributions based on crab-pot captures are biased. Spacing trials showed that pots positioned 50 m apart fished competitively but that there was no difference in catch between those placed 100 and 200 m apart. Catch distribution indicated that the presence of a crab in a pot reduced the probability of further captures. Temperature and incidence of recently moulted crabs accounted for 66% of variation in monthly catches. Catch-per-unit-effort (CPUE) data can be used as a measure of relative abundance of adults if allowance is made for temperature and the incidence of moulting. The capture-mark-release-recapture method for making population estimates was tested but it is concluded that, because of cost and bias in collecting techniques, it cannot be recommended as a technique for studying S. serrata.     

Matching spatial distributions of the sea star Echinaster sepositus and crustose coralline algae in shallow rocky Mediterranean communities

Understanding why a species is present in a particular location and the consequences of its presence is complex but necessary to identify the mechanisms that generate and maintain ecological diversity. The common sea star Echinaster sepositus can be either very abundant or non-existing in nearby localities of the western Mediterranean. Yet, the factors that shape its distribution and the impact of the sea star on natural communities remain uninvestigated. Here, we quantified multiple biotic and abiotic factors that may affect the distribution of E. sepositus and tested whether this sea star can shape the organization of the community it inhabits. Our results showed that the distribution of this sea star was highly contagious and positively correlated with the abundance and distribution of crustose coralline algae from tens of meters to tens of kilometers. Despite significant differences in community composition between localities with high or low abundance of the sea star, experimental addition of E. sepositus to natural communities failed to shift the composition of the algal community in 4 months. Overall, our results suggest that within habitat variability in the abundance of crustose coralline algae may explain the abundance of E. sepositus at multiple geographic scales, emphasizing the need to investigate small-scale processes at larger geographic scales.     

Early reproductive stages in the crustose coralline alga Phymatolithon lenormandii are strongly affected by mild ocean acidification

Coralline algae (Corallinales, Rhodophyta) are predicted to be negatively impacted by near-future ocean acidification. The effect of low pH/high pCO₂ on early life stages of Phymatolithon lenormandii (Areschoug) Adey was studied in a perturbation experiment. Several parameters including mortality, calcification (calcein staining) and development (growth and abnormalities) have been monitored for a month under experimental conditions ranging from pH_T = 8.00 (pCO₂ = 398 μatm) and pH_T = 7.55 (pCO₂ = 1,261 μatm). Our results demonstrate that survival and development of P. lenormandii early life stages can be impacted by small pH changes (ΔpH < ?0.1 pH unit). A negative impact of decreasing pH was observed including an increased mortality and a higher rate of abnormalities. Growth and calcification were still observed at the lowest pH (ΔpH = ?0.45). Growth rate was similar at all tested pH, but the maintenance of the skeleton under low pH was only possible through a persistent dynamic dissolution/calcification process, an energetically costly mechanism potentially draining resources from other vital processes.     

Bacteria on the surface of crustose coralline algae induce metamorphosis of the crown-of-thorns starfish Acanthaster planci

The crustose coralline alga Lithothamnium pseudosorum induces high rates of settlement and metamorphosis of larvae of the coral-eating crown-of-thorns starfish (Acanthaster planci). In cases where crustose coralline algae (CCA) induce metamorphosis of marine invertebrate larvae it is normally assumed that the inductive molecules are produced by the alga, but an alternative is that they originate from bacteria on the plant surface. Bioassays using shards of L. pseudosorum treated with several antibiotics, whereby some shards were reinfected with bacteria from the alga, showed that if bacteria populations are depleted then settlement and metamorphosis of larvae of A. planci are inhibited. This demonstrates that bacteria are necessary for induction and suggests that morphogenic substances are produced by bacteria on the surface of the alga and not directly by the alga itself. However, surface bacteria are not inductive if they are isolated from soluble algal compounds, suggesting either that they require a substrate from the alga to produce the inductive agents or, alternatively but less likely, that compounds from both the alga and bacteria are required. There is no evidence that inductive compounds derive from the alga, since algal cell debris and soluble extracts prepared from the alga do not induce metamorphosis of A. planci. This is the first time that induction of metamorphosis in a marine invertebrate by CCA has been shown to be mediated by bacteria associated with the alga.     

Growth and production of the red alga Gelidium sesquipedale off the Basque coast (northern Spain)

Several aspects of the growth of the agarophyte Gelidium sesquipedale Clemente (Thuret) were studied in sublittoral populations of the Basque coast (northern Spain) from 1981 to 1986. The standing stock of the populations varied greatly with season. During two consecutive years, 60 to 70% of summer biomass had been lost by the end of the unfavourable period of year to growth (autumn and winter). The decline in biomass was mainly due to partial losses of branches, with more branches being lost from larger fronds. Yearly variations in the standing stock were also recorded. Production estimates obtained by a destructive sampling method were not satisfactory because of the spatial heterogeneity of the populations at the three locations studied. Indirect estimation of production by measuring the growth of marked fronds in a control quadrat gave more reliable estimates. Production values of 622±124, 735±147 and 615±123 g dry wt m^-2 yr^-1 were obtained at the three locations, respectively. Frond elongation rates of 7.5, 6.1, 7.5 and 7.9 cm yr^-1, respectively, were recorded during four consecutive years. In winter elongation rates of fronds were rather low. Elongation did not seem to be affected by frond length, the self-shading effect of the canopy, or changes in depth. Cuts resulting from grazing were frequent on fronds through the year, decreasing significantly with increasing depth. Sheared fronds regenerated promptly, independent of season or height at which the cut was made. For commercial purposes the best season for harvesting is summer, which is the period of maximum standing stock, infertility, and calm warm waters that provide easier working conditions for divers.     

Sperm morphology, swimming velocity, and longevity in the house sparrow Passer domesticus

Sperm competition exerts strong selection on males to produce spermatozoa with an optimal morphology that maximizes their fertilization success. Long sperm were first suggested to be favored because they should swim faster. However, studies that investigated the relationship between sperm length and sperm competitive ability or sperm swimming velocity yielded contradictory results. More recently, ratios of the different sections of a spermatozoon (the head, midpiece, and flagellum) were suggested to be more crucial in determining swimming velocity. Additionally, sperm ability to remain and survive in the female storage organs may also influence fertilization success, so that optimal sperm morphology may rather maximize sperm longevity than velocity. In this study, we investigated how sperm morphology is related to sperm velocity and sperm longevity in the house sparrow Passer domesticus. Sperm velocity was found to be correlated with head/flagellum ratio. Sperm with small heads relative to their flagellum showed higher swimming velocity. Additionally, shorter sperm were found to live longer. Finally, we found sperm morphological traits to vary substantially within males and the head/flagellum ratio to be unrelated to total sperm length. We discuss the hypothesis that the substantial within-male variation in sperm morphology reflects a male strategy to produce a diversity of sperm from long, fast-swimming to short, long-living sperm to maximize their fertilization success in a context of sperm competition.     

Space use,longevity, and reproductive success in meadow voles

Summary We addressed the question of how reproductive success (RS) was limited in the shortlived but highly fecund meadow vole, Microtus pennsylvanicus. In so doing, we asked how differential space use patterns could affect longevity and hence RS in each sex. The sample comprised all voles achieving sexual competency over the course of a 40-week breeding season in a live-trapped population in Manomet, MA USA. Matrilineal families were determined using a radionuclide labelling technique; paternity was estimated using a maximum likelihood model. Individual RS was defined as the number of offspring successfully recruited into the trappable population per adult. We found that the variance in RS among female meadow voles was greater than the variance among males. In an attempt to explain this pattern, reproductively successful individuals were compared to reproductively unsuccessful individuals with regard to survivorship, maximum body weight achieved, and spatial mobility. The only difference between fathers and reproductively unsuccessful males was that fathers were heavier. In contrast, mothers differed from unsuccessful females in every measurement. Females lived longer than males, and mothers lived longer than either fathers or reproductively unsuccessful females. The observed differences in longevity may have been largely the result of differences in levels of mobility, assuming more mobile voles were more susceptible to predation. Mothers were significantly more site tenacious than were either males or unsuccessful females. These patterns explain the distribution of RS in our population if predation differentially affects male and female meadow voles. The meadow vole is the only non-polyandrous vertebrate reported to date in which the variance in RS among females exceeds the variance in RS among males.     

Fecundity,egg weight and longevity in relation to multiple matings in females of the monarch butterfly

Summary In many insects nutrients transferred by the male at mating are later incorporated into both the eggs and soma of the mated females. Accordingly, it has been suggested that insect females can use these male-derived nutrients for somatic maintenance and enhancement of their fecundity and fitness of their offspring. I this paper we tested the validity of these predictions by studying the longevity and reproductive output of 1, 2, 3, 4 or 5 times mated female monarchs Danaus plexippus, a nectar-feeding butterfly that is long-lived and strongly polyandrous and emerges as an adult without mature eggs. Females mated five times received ejaculates that corresponded to an average of 38% of their body weight at eclosion. However, we found that the number of times females had mated had no effect on their longevity, life-time fecundity, or egg weight. Although negative evidence should always be interpreted with caution, our study suggests that male-derived nutrients are less important for female longevity and reproductive output than are larval and adult food.     

Sperm motility and longevity in the giant cuttlefish, Sepia apama (Mollusca: Cephalopoda)

The sperm kinetics and fertilisation literature in marine invertebrates is heavily biased toward free-spawning species. Nonetheless, many species (e.g. cephalopods) transfer and/or fertilise gametes in confined external spaces or internally, creating very different selective pressures on sperm storage, sperm longevity and hence sperm competition. Here we report the results of an investigation into the effects of sperm age, water temperature and sperm concentration on sperm motility in the giant cuttlefish (Sepia apama). Significant positive correlations were found between percent motility and sperm concentration, and between sperm motile speed and sperm concentration. Mean percent motility of cuttlefish sperm suspension was still 9% eight hours after being released from the spermatophore and diluted into filtered seawater at 12°C (ambient field temperature during the spawning season). Sperm resuspended from spermatangia taken from (mated) females in the field were motile for up to 100 hours. When spermatophores were stored at 4°C motility was still observed in resuspended sperm after two months. Our results show that spermatangia and spermatophores can retain and release live sperm for long periods. The observed longevity of sperm in S. apama greatly increases the potential for sperm competition in this species.     

Queen longevity,queen adoption,and posthumous indirect fitness in the facultatively polygynous ant Myrmica tahoensis

In many polygynous ant species, established colonies adopt new queens secondarily. Conflicts over queen adoption might arise between queens and workers of established colonies and the newly mated females seeking adoption into nests. Colony members are predicted to base adoption decisions on their relatednesses to other participants, on competition between queens for colony resources, and on the effects that adopted queens have on colony survivorship and productivity. To provide a better understanding of queen-adoption dynamics in a facultatively polygynous ant, colonies of Myrmica tahoensis were observed in the field for 4 consecutive years and analyzed genetically using highly polymorphic microsatellite DNA markers. The extreme rarity of newly founded colonies suggests that most newly mated queens that succeed do so by entering established nests. Queens are closely related on average (rˉ = 0.58), although a sizable minority of queen pairs (29%) are not close relatives. An experiment involving transfers of queens among nests showed that queens are often accepted by workers to which they are completely unrelated. Average queen numbers estimated from nest excavations (harmonic mean = 1.4) are broadly similar to effective queen numbers inferred from the genetic relatedness of colony members, suggesting that reproductive skew is low in this species. Queens appear to have reproductive lifespans of only 1 or 2 years. As a result, queens transmit a substantial fraction of their genes posthumously (through the reproduction of related nestmates), in comparison to direct and indirect reproduction while they are alive. Thus queens and other colony members should often accept new queens when doing so will increase colony survivorship, in some cases even when the adopted queens are not close relatives. Received: 20 February 1996/Accepted after revision: 25 May 1996     

Growth,mortality and yield-per-recruit of deep-water royal red prawns (Haliporoides sibogae) off eastern Australia,using the length-based MULTIFAN method

The MULTIFAN method of Fournier et al. was used to analyse a series of monthly length-frequency distributions of deep-water royal red prawns (Haliporoides sibogae de Man, 1907, Solenoceridae) for the estimation of growth, mortality and yield-per-recruit. Length data were collected from commercial catches made between Latitude 33° and 35°S from November 1988 to May 1990. MULTIFAN distinguished six cohorts in the royal red prawn population, aged at 6 mo intervals from about 1.5 to 4 yr. Most prawns were between 2 and 3 yr of age. Females and males were fully recruited at 23.1 and 21.5 mm carapace length (CL) respectively, when about 2 yr old. Females grew faster and reached larger sizes at age than males (growth coefficient, K=0.37 yr^-1 and asymptotic length, L =48.3 mm CL for females, and K=0.49 yr^-1 and L =33.5 mm CL for males). Total mortality (Z) was estimated at 1.6 and 1.2 yr^-1 for females and males, respectively. Approximate values for natural mortality (M) ranged from 0.6 to 0.7 yr^-1 for females, and from 0.4 to 0.8 yr^-1 for males. Analysis of yield-per-recruit suggested that while the current yield could possibly be increased by decreasing the length at first capture (L _c ), there seems to be little potential for an increase in yield from increased fishing effort in the area currently fished.     

A note on the willingness to pay approach to the valuation of longevity

A modification of Freeman's model of willingness to pay for reduction of environmental risk to life is presented. Rather than treating work-related, consumption-related, and exogenous hazards as additive components of an overall level of safety, the revised model acknowledges the interdependence of these components. The results derived from this approach imply that (a) willingness to pay for a marginal reduction in risk depends upon the individual's initial level of utility; and (b) willingness to pay for a reduction in a given component of safety is related to the level of each component risk. The results of the revised model become more dramatic at higher levels of risk.     

Immigration pattern and success in red squirrels

Summary We studied the characteristics of immigrants and the effects of immigration on reproductive activity and spacing behaviour in red squirrels living in high-quality woodlands. Male immigration peaked in spring, female immigration in autumn. There was no sex bias in dispersal distance of local recruits or in the proportion of male/female immigrants, but more subadults than adults immigrated on the study plots. Hence, hypotheses explaining sex-biased dispersal were irrelevant in explaining immigration patterns in our study populations. Immigrant females were not in breeding condition, nor had they produced a litter prior to immigration. Hence breeding dispersal did not occur. Red squirrels are promiscuous, and females defend intrasexual territories while males have overlapping home ranges with a dominance hierarchy (Wauters et al. 1990; Wauters and Dhondt 1992). Site fidelity is very important to reproductive success and most parents still have a high residual reproductive value after having produced a litter. Under such circumstances, the resident fitness hypothesis (RFH; Anderson 1989) predicts that parents can benefit by forcing emigration of offspring if the latter are likely to find nearby vacancies. The settlement pattern of successful immigrants, which had a higher probability of becoming established when they had high body mass and when they were settling in plots with reduced intrasexual competition, agreed with the RFH and with the proximate dispersal mechanism suggested by Gliwicz (1992), that dispersal tendency in both sexes depends on the degree of intrasexual competition under local conditions. The fact that close inbreeding was never observed could indicate that random immigration of both sexes, within the social environment of a partly territorial, relatively long-lived species, has evolved not only to reduce competition for resources between parents and offspring but also as an inbreeding avoidance mechanism. Correspondence to: L. Wauters     

Influence of age polyethism on longevity of workers in social insects

Age polyethism is widespread among social insects and, as a rule, safe tasks are performed by workers earlier in life than are risky ones. Mathematical models were used to compare expected longevity of workers in colonies with and without age polyethism. The results of the models suggest that if aging does not depend on activity then age polyethism is profitable when safer tasks are performed earlier in life. If, however, aging depends on activity, age polyethism is profitable when safer tasks are performed earlier in life and if they are associated with higher aging-related mortality. On the other hand, age polyethism is not profitable if safer tasks are performed later in life, and if they are associated with lower aging-related mortality. Furthermore, if there is no aging, then age polyethism does not bring any benefits. Electronic supplementary material to this paper can be obtained by using the Springer Link server located at http://dx.doi.org/10.1007/s00265-001-0429-z     

Intra-clonal variation in the red seaweed Gracilaria chilensis

The phenotypic plasticity often found in seaweed populations has been explained only from the perspective of inter-population or inter-individual differences. However, many seaweeds grow and propagate by fragmentation of genetically identical units, each with the capacity to function on its own. If significant differences in performance exist among these supposedly identical units, such differences should be expressed upon the release and growth of these units. In this study we document two such types of variation in the red seaweed Gracilaria chilensis. Populations of sporelings, each grown under similar culture conditions and derived from carpospores shed by the same cystocarp exhibit significant differences in growth. In this species, each cystocarp develops from a simple gametic fusion, and cystocarp fusions occur too infrequently to account for the growth differences observed among recruits. In adult thalli, branches (ramets) derived from the same thallus (genet) and grown under similar conditions exhibit significant variation in growth rates and morphology. These findings have several implications. They suggest that carpospore production is not only an example of zygote amplification but that it also could increase variability among mitotically replicated units. Intra-clonal variability followed by fragmentation and re-attachment may increase intra-population variation which, in species of Gracilaria, is often larger than inter-population variation. In addition, the existence of intra-clonal variability suggests that strain selection in commercially important species may require a more continuous screening of highquality strains because of frequent genotypic or phenotypic changes in the various cultivars.     

Photosynthetic carbon acquisition in the red algaGracilaria conferta

Photosynthetic properties of the common red algaGracilaria conferta, collected from the eastern Mediterranean Sea were investigated in 1989, in order to begin evaluating its adaptative strategies with regard to the inorganic carbon composition of seawater, and to test whether the alleged C₄ photosynthesis of anotherGracilaria species is common within the genus. Net photosynthetic rates ofG. conferta were, under ambient conditions of inorganic carbon (ca. 10µM, CO₂ and 2.2 mM HCO ₃ ^- ), not sensitive to O₂ over the range 10 to 300µM, and the CO₂ compensation point was low (ca. 0.005µM). Ribulose-1,5-bisphosphate carboxylase/oxygenase was the major carboxylating enzyme, with a crude extract activity of 175µmol CO₂ g^–1 fresh wt h^–1 while phosphoenolpyruvate carboxylase and phosphoenolpyruvate carboxykinase were present at 70 and 20%, respectively, of that activity. No activities of the decarboxylases NAD-and NADP-malic enzyme could be detected. The¹⁴C pulse-chase incorporation pattern showed thatG. conferta fixes inorganic carbon via the photosynthetic carbon reduction cycle only, with no evidence for photosynthetic C₄ acid metabolism. Photosynthesis at the natural seawater pH of 8.2 was, at 25°C and saturating light, saturated at the ambient inorganic carbon concentration of 2.5 mM. It is proposed that, under ambient inorganic carbon conditions, a CO₂ concentrating system other than C₄ metabolism provides an internal CO₂ concentration sufficient to suppress the O₂ effect on ribulose-1,5-bisphosphate carboxylase/oxygenase and, thus, on photorespiration, in a medium where the external free CO₂ concentration is lower than theK _m(CO₂) of the carboxylating enzyme. Since inorganic carbon, under natural saturating light conditions, seems not to be a limiting factor for photosynthesis ofG. conferta, it likely follows that other nutrients limit the growth of this alga in nature.     

Skeleton and sclerite formation in the precious red coral Corallium rubrum

The carbonate skeleton of the gorgonian coral Corallium rubrum (L.) is composed of both a skeletal axis and numerous sclerites scattered in the mesoglea. Studies carried out on these skeletal elements and their associated tisues using microscopy and X-ray microanalysis, suggest a close relationship between the process of sclerite formation and skeletogenesis. The skeleton is surrounded by an axial epithelium composed of a single cell type. These cells associate intimately with mesogleal sclerites and scleroblasts, incorporating them into a nascent skeleton at the branch tip. Subsequent (sub-apical) growth appears to occur solely through the agency of the axis epithelial cells that serve to physically separate mesogleal sclerites and scleroblasts from contact with the axis. The epithelium is associated with the production of layered calcite crystals and irregular protuberances that constitute the mature, calcareous skeleton. Free sclerites in the mesoglea appear to be the product of multiple cells that are cytologically indistinguishable from those in the axis epithelium. Like the axis, sclerites are produced as layers of calcite crystals with irregular protuberances. The protuberances differ only slightly from those of the axis, and the skeleton is mineralogically indistinguishable from the sclerites. Thus, the skeleton of red coral is not primarily the product of fused sclerites. Instead, we suggest that the axis epithelium treats the incipient skeleton as if it were the core of a single sclerite, and conversely, that the mesogleal scleroblasts of C. rubrum constitute a fragmented axis epithelium.     

Variation in the food hoarding behaviour of red squirrels

Summary To investigate variation in food storing behaviour within a population of red squirrels (Tamiasciurus hudsonicus) we intensively sampled the size and spatial dispersion of hoards. Single-item and large hoards were far more prevalent than expected by chance (Figs. 1 and 2) and appeared to be distinct components of the hoarding tactic employed in this population. Scattered single-item hoards represented more than half of all hoards constructed and approximately one-fifth of all cones stored. Large hoards (> 10 cones) comprised less than 15% of all hoards, and about 50% of all cones stored. In response to experimental manipulations, squirrels altered their hoarding behaviour in order to concentrate cones in the litter and brush heaps we provided, but still scattered a large proportion of their stores. We conclude that hoarding patterns in red squirrels are more variable and plastic than has previously been recognized, and discuss this variability in terms of adaptive explanations. Offprint requests to: R.J. Robertson