Physioecology of zooplankton. I. Effects of phytoplankton concentration,temperature, and body size on the growth rate of Calanus pacificus and Pseudocalanus sp.

Changes in dry weight and in weight-specific growth rates were measured for copepodite stages of Calanus pacificus Brodsky and Pseudocalanus sp. cultured under various combinations of phytoplankton concentration and temperature. Mean dry weight of early copepodites was relatively unaffected by either food concentration or temperature, but mean dry weight of late stages increased hyperbolically with food concentration and was inversely related to temperature. The food concentration at which maximum body weight was attained increased with increasing temperature and body size, and it was considerably higher for C. pacificus than for Pseudocalanus sp. This suggests that final body size of small species of copepods may be determined primarily by temperature, whereas final body size of large species may be more dependent on food concentration than on temperature. Individual body weight increased sigmoidally with age. The weight-specific growth rate increased hyperbolically with food concentration. The maximum growth rate decreased logarithmically with a linear increase in body weight, and the slope of the lines was proportional to temperature. The critical food concentration for growth increased with body size proportionally more at high than at low temperature, and it was considerably higher for C. pacificus than for Pseudocalanus sp. Because of these interactions, early copepodites optimized growth at high temperature, even at low food concentrations, but under similar food conditions late stages attained higher growth at low temperature. The same growth patterns were found for both species, but the rates were significantly higher for the larger species, C. pacificus, than for the smaller one, Pseudocalanus sp. On the basis of findings in this study and of analyses of relationships between the maximum growth rate, body size, and temperature from other studies it is postulated (1) that the extrapolation of growth rates from one species to another on the basis of similarity in body size is not justified, even for taxonomically related species; (2) that the allometric model is inadequate for describing the relationship between the maximum weight-specific growth rate and body size at the intraspecific level; (3) that the body-size dependence of this rate is strongly influenced by temperature; and (4) that species of zooplankton seem to be geographically and vertically distributed, in relation to body size and food availability, to optimize growth rates at various stages of their life cycles.Contribution No. 1127 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

不同溶剂对提取原煤中PAHs种类、含量与分布的影响

以淮北煤田的煤样为研究对象,选取二硫化碳（CS2）、二氯甲烷（CH2Cl2）、正己烷（C6H14）三种溶剂,采用索氏提取法对原煤中的多环芳烃进行提取．利用气相色谱-质谱联用仪对提取液中的多环芳烃进行定量分析,研究了不同溶剂对原煤中多环芳烃种类、含量和分布的影响,通过研究发现,C6H14的提取能力显著低于CS2和CH2Cl2,CS2容易提取低环数的PAHs,CH2Cl2更容易提取高环数的PAHs．     

Bright turquoise as an intraspecific signal in the chameleon grasshopper (Kosciuscola tristis)

Bright colours often communicate important information between conspecifics. In sexually dichromatic species where males exhibit bright colours, two hypotheses are often invoked to explain the function of the colour. First, if a male’s bright colour contains information about his quality, females may prefer brighter males. Equally, male colour may reliably provide other males with information about fighting ability or resource holding potential. In such circumstances, brighter males may win altercations and/or males may use rival colour to assess their likelihood of winning an interaction. In the chameleon grasshopper (Kosciuscola tristis), males but not females turn bright turquoise when their body temperature exceeds 25 °C. In this study, we tested whether the turquoise phase of colour change has a signaling role in inter- and intrasexual contexts. We predicted that females would prefer bright turquoise males over dull males, but found no evidence from several choice experiments to support this hypothesis. We also predicted that brighter males would win more fights than duller males. Whilst we did not find that brighter males won more fights in staged experiments, we found that the brightness of males who chose to enter fights was significantly correlated with their opponents’ brightness. Our results suggest that the brightness of males’ turquoise phase may provide competitors with important information about their rival’s fighting ability.     

Temperature relationships of some tropical Pacific intertidal gastropods

Tissue and environmental-temperature measurements were made using thermistors in Hawaii, Guam, Palau and Singapore in the summer of 1970. Limpets have a very limited ability to regulate body temperature, and several species possessed tissue temperatures greater than those of the substratum. Some neritids and the trochid Monodonta labio possessed tissue temperatures as much as 2.2° to 2.8°C below substratum temperature. Littorinids attached by a mucus film to the substratum were found to have relatively lower body temperatures than those attached by the foot. Among neritids there is a tendency for high intertidal species to be better temperature regulators than low intertidal species. A very important factor in the temperature regulation of intertidal snails appears to be the extent of the area of contact between the animal and the substratum, which is greatest in limpets and least (negligible) in littorinids. Evaporative cooling and light coloration of the shell play a role in certain species and groups.     

Parasites determine a predator's optimal feeding strategy

Summary The diet selected by three-spined sticklebacks (Gasterosteus aculeatus) depends on the degree of parasitization by one or both of two parasite species (Schistocephalus solidus, Glugea anomala). Uninfested fish prefer the more profitable of two different size classes of prey (Daphinia magna). Fish parasitized by Glugea or Schistocephalus attack both prey types equally often, whereas sticklebacks infested by both parasite species prefer to attack the less profitable prey. The diet selected is optimal under the condition that parasites decrease their host's competitive ability.     

UV wavelengths make female three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>) more attractive for males

Numerous fishes possess UV vision and show UV patterns, which often play a role in social communication, especially during courtship. UV light is scattered strongly in water and thus might be used for intra-specific close-range communication without being detected by predators. In addition to the red-belly colouration and blue-coloured iris, male three-spined sticklebacks possess UV reflectance on their skin, and females prefer males presented with UV light rather than without. To investigate whether UV also influences male visual preference for females in this species, we used a dichotomous mate choice design in which one female could be viewed by a male in two visual conditions. Courting male sticklebacks preferred females that were presented in full-spectrum conditions including UV to the same females presented in conditions lacking this light component. Furthermore, control trials with neutral-density filters indicated that male preference in the UV treatment was not affected by a difference in achromatic brightness between the UV+ and UV− stimuli. Reflectance measurements of gravid females suggest an enhanced UV contrast between the dark bar pattern, which is characteristic of gravid females, and silvery body parts.     

Foraminiferan (Protozoa) epizoites on Arctic isopods (Crustacea) as indicators of isopod behaviour?

A total of 38 219 specimens representing 63 species of marine isopods (Crustacea) from deep and shallow Arctic waters were studied in a search for epizoic foraminifers (Protozoa). Foraminifers occurred on 21 species, and their frequency was generally low. A total of 290 foraminifer individuals were found, of which 289 belonged to Cibicides wuellerstorfi, C. refulgens and Cibicides spp. (juveniles) (Cibicidae), while only a single individual belonged to Cornuspira sp. (Cornuspiridae). The foraminifers were most frequent on species of the families Munnidae, Ischnomesidae (suborder Asellota) and on Gnathia stygia (suborder Gnathiidea), but were totally absent from the asellote families Janiridae, Haploniscidae, Nannoniscidae and from the suborder Epicaridea. The foraminifers were mainly located on the legs (Munna acanthifera), the anterior part of the body (Haplomesus quadrispinosus, heteromesus frigidus), or on the head (G. stygia adults). The epizoic foraminifers occur mainly on epibenthic isopods, which do not or only rarely clean themselves. The foraminifers are known to prefer elevated substrata, and in this the habitat of the isopods and the foraminifers coincide. The size of individual isopods was not related to the presence or absence of foraminifers.     

Effects of environmental factors and body size on rates of oxygen consumption in Archaeomysis grebnitzkii and Neomysis awatschensis (Crustacea: Mysidae)

The rates of oxygen consumption in relation to oxygen tension, temperature, salinity and body size were determined for Archaeomysis grebnitzkii Czerniavsky and Neomysis awatschensis (Brandt). Oxygen uptake was regulated by both species down to an oxygen tension of about 50 mm Hg (30% of saturation value); below this level, it was related to the oxygen tension in the medium. Oxygen tension below 20 mm Hg was lethal. A statistically significant interaction was noted for the effects of species, temperature, and salinity. In general, oxygen-uptake rate decreased with decreasing salinity in both species. The effect of size is expressed as a power function of body weight. The regression coefficients of oxygen consumption on body weight, 0.70 for A. grebnitzkii and 0.62 for N. awatschensis, were found to be significantly different from each other.Contribution No. 726 from the Department of Oceanography, University of Washington, Seattle, USA.     

Male lagoon gobies, <Emphasis Type="Italic">Knipowitschia panizzae</Emphasis>, prefer more ornamented to larger females

Female ornamentation may be directly sexually selected, by male choice or female competition, or occurs as the result of a genetic correlation, arising from sexual selection on males. However, increasing evidence supports the former hypothesis, suggesting that males actively choose their partner preferring traits indicative of female quality. In the lagoon goby, Knipowitschia panizzae, a polygynous species whose males perform parental care to eggs, body length and the size of a sex-specific yellow patch on the belly are known to be reliable indicators of female fecundity. In this paper, we tested, using dummies, the male’s mating preferences for female body and yellow belly patch sizes. The two experimental trials in which a single female trait was variable showed that males prefer a larger belly patch and a larger body size, indicating that both these characters are selected by male mate choice. However, when faced with dummies exhibiting an inverse combination of body and belly patch sizes (experiment 3), males significantly preferred the smaller ones with larger yellow belly patches. A calculation of dummy theoretical fecundity reveals that in the first two experiments, males would have received an immediate benefit from their choice in terms of egg number, whereas in the third one, males chose partners that would have provided them with fewer eggs. The male lagoon goby preference for females with larger belly patches, regardless of their size, suggests that this trait, in addition to indicating fecundity, conveys information about other aspects of female and/or egg quality.     

Respiration and ammonia excretion by marine metazooplankton taxa: synthesis toward a global-bathymetric model

For thirteen representative taxa of metazooplankton from various depth horizons (<4,200 m) of the world’s oceans, respiration rate (681 datasets on 390 species) and ammonia excretion rate (266 datasets on 190 species) are compiled and analyzed as a function of body mass (dry mass, carbon or nitrogen), habitat temperature, habitat depth and taxon. Stepwise-regression analyses reveal that body mass is the most important parameter, followed by habitat temperature and habitat depth, whereas taxon is of lesser importance for both rates. The resultant multiple regression equations show that both respiration rate and ammonia excretion rate (per individual) increase with increase in body mass and habitat temperature, but decrease with habitat depth. Some taxa are characterized by significantly higher or lower rates of respiration or ammonia excretion than the others. Overall, the global-bathymetric models explain 93.4–94.2 % of the variance of respiration data and 80.8–89.7 % of the variance of ammonia excretion data. The atomic O:N ratios (respiration/ammonia excretion) are largely independent of body mass, habitat temperature, habitat depth and taxon, with a median of 17.8. The present results are discussed in light of the methodological constraints and the standing hypotheses for the relationship between metabolic rate and temperature. Perspectives for model improvement and possible application of it to plankton-imaging systems for rapid assessment of the role of metazooplankton in C or N cycles in the pelagic ecosystem are briefly discussed.     

The presence of eggs in the nest and female choice in common freshwater gobies (Rhinogobius brunneus)

In many species of fishes with paternal care, females prefer to spawn with males who are already guarding eggs. We studied the effect of egg presence on female mate choice in common freshwater gobies, Rhinogobius brunneus sp. OR. In our tests, females did not prefer males with eggs, suggesting egg presence per se may not act as a cue to attract females. We also examined the effects of brood size on paternal care and offspring survival to look for possible benefits females could obtain when choosing males with eggs. Both fanning by egg-guarding males and egg survival increased with brood size. The presence of neighboring males did have a significantly negative effect on males' parental activity, which subsequently results in a lower level of egg survival than in solitary egg-guarding males. This result provides a partial explanation for the result of eggs not attracting females to mate.     

Sexual selection in darkness? Female mating preferences in surface- and cave-dwelling Atlantic mollies,<Emphasis Type="Italic"> Poecilia mexicana</Emphasis> (Poeciliidae,Teleostei)

In many species, females prefer large males as mating partners. Male size can easily be determined visually. We examined how such mate choice evolves when individuals are deprived of sensory information previously used to determine male quality. Using a cave-dwelling population of Atlantic mollies, Poecilia mexicana, as a model, we asked whether the female mating preference for large male body size still occurs in this cave fish, or whether it is lacking due to the absence of visual communication in the cave environment. In simultaneous choice tests we compared the time females spent with a large or a small male. In the river-dwelling and cave-dwelling fish, as well as the population originating from the cave entrance, we found a significant preference for large males in light. Hence, the ancestral mating preference of surface-dwelling mollies has been maintained during the evolution of the cave form. However, only the cave population exhibited a mating preference in darkness. Determination of male body size appears to have been taken over by a non-visual sensory system, probably the lateral line.Communicated by K. Lindström     

Metabolic rates of epipelagic marine copepods as a function of body mass and temperature

Metabolic rates (oxygen consumption, ammonia excretion, phosphate excretion) have been calculated as a function of body mass (dry, carbon, nitrogen and phosphorus weights) and habitat temperature, using multiple regression. The metabolic data used for this analysis were species structured, collected from Arctic to Antarctic seas (temperature range: -1.7°C to 29.0°C). The data were further divided into geographical and/or seasonal groups (35 species and 43 data sets for oxygen consumption; 38 species and 58 data sets for ammonia excretion; 22 species and 31 data sets for phosphate excretion). The results revealed that the variance attributed to body mass and temperature was highest (93-96%) for oxygen consumption rates, followed by ammonia excretion rates (74-80%) and phosphate excretion rates (46-56%). Among the various body mass units, the best correlation was provided by the nitrogen unit, followed by the dry weight unit. The calculated Q₁₀ values varied slightly according to the choice of body mass units; overall ranges were 1.8-2.1 for oxygen consumption rates, 1.8-2.0 for ammonia excretion rates and 1.6-1.9 for phosphate excretion rates. The effects of body mass and temperature on the metabolic quotients (O:N, N:P, O:P) were insignificant in most cases. Although the copepod metabolic data used in the present analysis were for adult and pre-adult stages, possible applications of the resultant regression equations to predict the metabolic rates of naupliar and early copepodite stages are discussed. Finally, global patterns of net growth efficiency [growth (growth+metabolism)^-1] of copepods were deduced by combining the present metabolic equation with Hirst and Lampitt's global growth equation for epipelagic marine copepods.     

A test of male mating and hunting success in the kestrel: the advantages of smallness?

We tested female choice for male wing and tarsus length and body mass in the kestrel (Falco tinnunculus), a species in which males average about 10% smaller than females. We also studied how male characters are related to their hunting success. In the laboratory, females preferred lighter males with shorter tarsi as mates, if the difference in those characters between competing males was larger than average. Lighter and shorter-winged males seemed to be better hunters than heavier and longer-winged males. Field observations in a year in which voles were scarce suggested that shorter-winged males were also better food providers in courtship feeding than longer-winged males,although in good vole years such a relationship was not found. We argue that females may prefer to pair with smaller males, because they have higher flight performance and better hunting success than heavier males. By doing so, females may gain direct breeding advantages. We conclude that both female choosiness and the hunting efficiency of males well contribute to reversed sexual size dimorphism (RSD, females larger than males) in the kestrel. Received: 18 July 1995/Accepted after revision: 17 August 1996     

Physioecology of zooplankton. IV. Effects of phytoplankton concentration,temperature, and body size on the net production efficiency ofCalanus pacificus

Weight-specific rates of individual production, total metabolic expenditure and assimilation, and net production efficiencies were estimated forCalanus pacificus Brodsky of selected body weights cultured at various phytoplankton concentrations and temperatures. The weight-specific rate of individual production increased hyperbolically with food concentration, and the maximum rate of individual production decreased logarithmically with a linear increase in body weight propotionally more at high than at low temperature. The weight-specific rate of total metabolic expenditure decreased logarithmically with increasing body weight and was unaffected by changes in food concentration. The effects of food concentration and temperature on the weight-specific rate of assimilation were similar to those on the rate of individual production, but the effect of body size differed considerably. The diversity in the temperature and body-size dependence of the maximum weight-specific rates of various physiological processes suggest (1) that, except for the metabolic rate, the allometric model (log-log relation) is inadequate for describing relationships between maximum rates of physiological processes and body size within species, and (2) that the common assumption that temperature affects the rates of various physiological processes in similar ways is not justified. Net production efficiency increased hyperbolically with food concentration, and the maximum production efficiency first increased slightly and then decreased gradually with increasing body weight. Small copepods attained higher efficiency at high temperature but larger ones did so at low temperature. The critical food concentrations for production efficiency and for the rate of individual production increased with increasing temperature and body size. Because of the effects of interactions among critical food concentration, temperature, and body size on the rates of growth and individual production and on net production efficiency, early development stages ofC. pacificus optimized growth and food conversion efficiency at high temperature, but late stages, particularly at low food concentrations, grew best and transformed food more efficiently at low temperature.Contribution No. 1130 From the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Behavioral adjustments of a pipefish to bacterial Vibrio challenge

Animals can profit from increasing temperatures by prolonged breeding seasons and faster growth rates. However, these fitness benefits are traded off against higher parasite load and increased virulence of temperature-sensitive pathogens. In thermally stratified habitats, behavioral plasticity can allow hosts to choose the optimal temperature to enhance individual fitness and to escape parasite pressure. To test this idea, we performed a temperature choice experiment with the host–parasite system of the sex-role reversed broad-nosed pipefish (Syngnathus typhle) and its bacterial pathogen Vibrio spp. In this species, pregnant males are expected to face a trade-off between shortening their brooding period in warm water and decreasing the effect of the infection in cold water. We found that exposure to Vibrio changed the temperature preference for both pregnant and nonpregnant males, as well as females compared to nonchallenged fish that tended to prefer warm water. This study shows that behavioral plasticity is one option for avoidance of higher bacterial prevalence, as expected due to rising ocean temperatures.     

Mate choice,fecundity and sexual dimorphism in two pipefish species (Syngnathidae)

Summary In order to understand the causes of sexual dimorphism, mate choice and size-related fecundity were studied in two pipefish species, Syngnathus typhle and Nerophis ophidion. Sexual dimorphism is more pronounced in N. ophidion; females are larger, have sexual colourings, and are more active during courtship. In S. typhle the sexes are alike in all these respects. Males brood their offspring in both species. In N. ophidion fecundity was positively correlated with both body size and the amount of sexual colouring in females. In males no correlation between body size and fecundity, or between body size and embryo size existed. Predictably, in mate choice experiments with equal-sized females, males chose females with more extensive sexual colourings. We explain sexual dimorphism in this species as a consequence of both natural selection (fecundity increases with size in females but not in males) and sexual selection (males prefer larger females). We argue that sexual size dimorphism did not evolve by selection minimizing overlap in food niches between the sexes, because food production is high in the Zostera beds where the fishes live, and no size dimorphism was found in the sympatrically occurring S. typhle. Furthermore, in N. ophidion dimorphism is not greater in a particular mouth character than in overall body size. In S. typhle egg size and the average number of eggs transferred per spawning were positively correlated with female body size. Apparently more energy per offspring was provided by larger males than by smaller males, and larger males also carried more offspring. As predicted, large mates were preferred by both sexes in mate choice experiments. This is explicable in terms of both natural selection (fecundity increases with size in both sexes) and sexual selection (both sexes prefer large mates). As a consequence of selection acting in the same direction in both sexes, sexual dimorphism is absent in S. typhle.     

Metabolic rates of epipelagic marine zooplankton as a function of body mass and temperature

The metabolic rates (oxygen uptake, ammonia excretion, phosphate excretion) of epipelagic marine zooplankton have been expressed as a function of body mass (dry, carbon, nitrogen and phosphorus weights) and habitat temperature, using the multiple-regression method. Zooplankton data used for this analysis are from phylogenetically mixed groups (56 to 143 species, representing 7 to 8 phyla, body mass range: 6 orders of magnitude) from various latitudes (habitat temperature range:-1.4° to 30°C). The results revealed that 84 to 96% of variation in metabolic rates is due to body mass and habitat temperature. Among the various body-mass units, the best correlation was provided by carbon and nitrogen units for all three metabolic rates. Oxygen uptake, ammonia excretion and phosphate excretion are all similar in terms of body-mass effect, but differ in terms of temperature effect. With carbon or nitrogen body-mass units, calculated Q₁₀ values are 1.82 to 1.89 for oxygen uptake, 1.91 to 1.93 for ammonia excretion and 1.55 for phosphate excretion. The effects of body mass and habitat temperature on the metabolic quotients (O:N, N:P, O:P) are insignificant. The present results for oxygen-uptake rate vs body mass do not differ significantly from those reported for general poikilotherms by Hemmingsen and for crustaceans by Ivleva at a comparable temperature (20°C). The importance of a body-mass measure for meaningful comparison is suggested by the evaluation of the habitat-temperature effect between mixed taxonomic groups and selected ones. Considering the dominant effects of body mass and temperature on zooplankton metabolic rates, the latitudinal gradient of community metabolic rate for net zooplankton in the ocean is estimated, emphasizing the non-parallelism between community metabolic rates and the standing stock of net zooplankton.     

Forced copulation as a conditional alternative strategy in camel crickets

In the animal kingdom, males typically acquire reproductive access to females through courtship, nuptial gifts, or combat. In many species, small or poor-condition males that are unable to produce an attractive display, proffer an adequate mating gift, or dominate in male–male competition may resort to forced copulation. Forced copulation has been previously demonstrated in the camel cricket Pristoceuthophilus marmoratus and is uniquely facilitated by male hind leg weaponry that functions in both intrasexual competition and sexual coercion. Because male leg armaments are positively allometric, the largest males would seem best equipped to force-copulate, although they might be the most attractive to females, and so least likely to adopt a forced copulation strategy. In order to resolve this potential paradox and determine which males are in fact most likely to force-copulate, we manipulated male body size using diet and performed mating trials. Results showed that (1) male size increased with diet quality, (2) females appear to prefer larger males, and (3) smaller males were more likely to attempt forced copulation. Our results thus suggest that forced copulation is a condition-dependent alternative mating tactic used by small males unattractive to females.     

Global effects of extreme temperatures on wild bumblebees

Climate plays a key role in shaping population trends and determining the geographic distribution of species because of limits in species’ thermal tolerance. An evaluation of species tolerance to temperature change can therefore help predict their potential spatial shifts and population trends triggered by ongoing global warming. We assessed inter- and intraspecific variations in heat resistance in relation to body mass, local mean temperatures, and evolutionary relationships in 39 bumblebee species, a major group of pollinators in temperate and cold ecosystems, across 3 continents, 6 biomes, and 20 regions (2386 male specimens). Based on experimental bioassays, we measured the time before heat stupor of bumblebee males at a heatwave temperature of 40 °C. Interspecific variability was significant, in contrast to interpopulational variability, which was consistent with heat resistance being a species-specific trait. Moreover, cold-adapted species are much more sensitive to heat stress than temperate and Mediterranean species. Relative to their sensitivity to extreme temperatures, our results help explain recent population declines and range shifts in bumblebees following climate change.