Growth of Antarctic krill Euphausia superba at South Georgia

 Antarctic krill Euphausia superba has a central role in the ecosystem of the Southern Ocean and knowledge of its growth rate is central to determining the factors influencing population dynamics. The length of Antarctic krill in the diet of Antarctic fur seals Arctocephalus gazella at South Georgia revealed a consistent increase in size between ca. 42 and ca. 54 mm over the period October–March, indicating growth rates much higher than predicted by existing models. Geographical variation in growth rate may result in 2-year-old krill at South Georgia attaining the same size as 3-year-old krill in the Antarctic Peninsula region. The effect of geographical variation in growth rate on the population structure of krill has important implications for comparing the fate of individual cohorts over large scales and in the interpretation of krill life-cycles. Received: 20 May 2000 / Accepted: 11 August 2000     

Synchronous recruitment and growth pattern of planktonic larvae of the amphioxus Branchiostoma belcheri in the Seto Inland Sea, Japan

Planktonic larvae of the amphioxus Branchiostoma belcheri were collected in the western part of the Seto Inland Sea, Japan, in order to document their seasonal occurrence, recruitment, and growth patterns. The larvae appeared from mid-July to early October and their size ranged from 0.6 to 5.5 mm. Three distinct cohorts were observed during the summers, indicating multiple, synchronous spawning within the population. The length increment pattern of a cohort demonstrates slow growth for at least a week during the early larval stage. July 10, 20, and 30 are the inferred dates when spawnings started to produce the three cohorts in 2001. Favorable temperature range for spawning was 21–23°C. Bias in population structure due to advection can be assumed to be negligible; therefore the length increment of the cohort, about 0.05 mm day⁻¹, is regarded as the larval growth rate after the initial slow-growth stage. The planktonic stage of the first cohort in 2001 is estimated to have lasted around 55 days. The relative proportions of the second and third cohorts in samples from surface and mid-depth waters suggest downward migration of the planktonic larvae, as they age.     

Biodiversity in Littorina species (Mollusca: Gastropoda): a physiological approach using heat-coma

 The upper critical thermal limits were measured as heat-coma temperatures in Littorina species as a tool for examining physiological diversity. Thermal tolerance was found to be stable within a population, but extensive variation occurred between populations and species. All species examined displayed heat-coma values of ∼30 °C and did not show a positive correlation with shore height. Eulittoral-fringe species tended to have higher coma temperatures than eulittoral species. Coma temperatures varied both seasonally and geographically. Lethal thermal limits were also investigated; these were ∼10 C° higher than coma temperatures. Received: 16 August 1999 / Accepted: 13 June 2000     

Importance of food quantity to structural growth rate and neutral lipid reserves accumulated in Calanus finmarchicus

 Growth and developmental rates were determined for copepodids of Calanus finmarchicus (Gunnerus) from experimental seawater mesocosms in a western Norwegian fjord. The instantaneous growth rates (g) from copepodid stage I (CI) to adult ranged from 0.08 to 0.10 d⁻¹. Daily per capita mortality rate of the cohorts was as low as 0.012 d⁻¹ (1.2% d⁻¹). At local increasing temperatures (5.1 to 8.3 °C), development was equiproportional, and the cumulative median development time from egg to CV was approximately 65 d. CV moulted to males and females, and egg production was initiated. Enhancement of food resources by nutrient addition caused a 23.4% increase in growth rates from CI to adult. Additionally, copepodid stages showed a generally larger body size, carbon and nitrogen content and total storage lipid content (wax esters + triacylglycerols) in response to enhanced resources. Our data support an elsewhere proposed exponential-growth hypothesis; growth of the structural compartments and store lipids (mostly wax esters) was exponential during the copepodid stages. However, a sigmoidal pattern of growth best described growth of adult stages if reared at high resources, and depot lipid accumulation in late CVs and adults at high resources. Body nitrogen growth increased exponentially, however, no significant changes in nitrogen specific growth rates were found between individuals from low and high resources. CV and adults seem to have reached near-maximal weights at high resources, whereas structural weight continued to increase at low resources. Despite the differences in structural growth dynamics, cohort development was similar until the end of CV. During the onset of sexual differentiation, the male:female ratio and the adult:CV ratio were highest at high food resources, suggesting that the time used for the final moult depends on the feeding history of the copepods in relation to food quality and quantity. It appears that relatively small changes in food availability strongly influence the biochemical composition of C. finmarchicus copepodids. Received: 28 May 1999 / Accepted: 10 February 2000     

Life history and production of the mysid <Emphasis Type="Italic">Orientomysis robusta</Emphasis>: high <Emphasis Type="Italic">P/B</Emphasis> ratio in a shallow warm-temperate habitat of the Sea of Japan

Although mysids play important roles in marine food chains, studies on their production are scarce, especially for warm-water species. We investigated life history and production of Orientomysis robusta in a shallow warm-temperate habitat of the Sea of Japan. Its spawning and recruitment occurred throughout the year; 19 overlapping cohorts were recognizable over an annual cycle. The summer cohorts recruited in July–September exhibited rapid growth, early maturity, small brood size, and small body size. A converse set of life history traits characterized the autumn–winter cohorts recruited in October–March. The spring cohorts recruited in April–June had intermediate characteristics of both cohorts. Life spans were 19–33, 21–48, and 69–138 days for summer, spring, and autumn–winter cohorts, respectively, and mortality rates were high for spring and summer cohorts, especially during June–August but were low for autumn–winter cohorts. Production calculated from the summation of growth increments was 488.8 mg DW m⁻² year⁻¹ with an annual P/B ratio of 21.26. The short life span seems to be responsible for such an extremely high P/B ratio. A method not requiring recognition and tracking cohorts gave similar values (534.0 mg DW m⁻² year⁻¹ and 20.49). The close agreement in production values between the two methods indicates our estimates are valid.     

Effects of temperature,size, and food on the growth of <Emphasis Type="Italic">Membranipora membranacea</Emphasis> in laboratory and field studies

In the rocky subtidal ecosystem of the western North Atlantic outbreaks of the introduced epiphytic bryozoan Membranipora membranacea cause defoliation of kelp beds and facilitate the introduction of other non-native benthic species. We quantified size- and temperature-dependent growth rates of M. membranacea colonies in the field and the laboratory for durations of 8–23 days. Also, we examined the interaction between food abundance and temperature on growth rates of newly settled colonies in the laboratory. Growth rates were positively related to temperature and increased exponentially with size of colonies over the ranges examined (5.7–16.2°C and 0.5–192 mm, respectively), and were significantly higher in the field than in the laboratory. There was an interactive effect between food and temperature on the size and growth rates of colonies, with the most pronounced effects of food limitation on colonies grown at the warmest temperatures, and no effect of food on colonies grown at the coldest temperatures. Quantifying the growth rates of introduced species is essential to understanding their population dynamics, particularly when outbreaks can have severe impacts on the native community.     

Questions on the mechanism of temperature adaptation in marine phytoplankton

The rate of light-saturated photosynthesis in 3 marine algae [Phaeodactylum tricornutum Bohlin, Nitzschia closterium (Ehrenberg) Smith and Dunaliella tertiolecta Butcher] varies during growth in batch culture. The photosynthetic rare declines most rapidly during growth at the higher temperatures. Because of these changes in photosynthesis rate, the previously reported enhanced photosynthetic abilities caused by growth at lower temperatures (generally interpreted as evidence for higher enzyme levels) can only be observed when measurements are made late in the exponential phase or after the onset of the stationary phase of growth. When allowance is made for the earlier peak of photosynthetic ability in cultures growing at higher temperatures, there is no evidence for adaptation to lower temperatures being caused by increased levels of the enzymes required for carbon-dioxide fixation. When the changes due to growth in batch culture are taken into account, certain effects of temperature can be recognized. the dry weight: chlorophyll ratio of all 3 algae increases with decreasing growth temperatures. For P. tricornutum and N. closterium, growth at lower temperatures reduces the cellular content of chlorophyll a, but has little effect on the chlorophyll content of D. tertiolecta. The dry weight: cell-number ratio of D. tertiolecta and P. tricornutum increases with lower growth temperatures, but growth temperature has little effect on the cell mass of N. closterium. Growth of the 3 algae at lower temperatures does not increase their ability to photosynthesize at these lower temperatures. Rather, it reduces their ability to assimilate carbon dioxide at the higher temperatures.     

The nature of heat coma in Littorina littorea (Mollusca: Gastropoda)

 The nature of heat coma was examined in the edible periwinkle Littorina littorea (L.). Duration of acclimation did not influence heat-coma temperature at 12 °C, although other acclimation temperatures were important in influencing thermal tolerance, with positive shifts in coma temperature observed in response to elevated temperatures. Previous thermal history also influenced heat-coma temperatures. Individuals subjected to repeat heat-coma events on a daily basis showed significant declines (P < 0.05) in coma-temperature; in contrast individuals exposed to repeat heat-coma events on a weekly basis showed no decline in thermal tolerance. Size-effects occurred at selected sites, where decreased heat-coma temperatures were recorded in large individuals. Received: 16 August 1999 / Accepted: 13 June 2000     

Generalists and specialists along a latitudinal transect: patterns of thermal adaptation in six species of damselflies

Tropical organisms colonizing temperate environments face reduced average temperatures and dramatic thermal fluctuations. Theoretical models postulate that thermal specialization should be favored either when little environmental variation is experienced within generations or when among-generation variation is small relative to within-generation variation. To test these predictions, we studied six temperate species of damselflies differing in latitudinal distribution. We developed a computer model simulating how organisms experience environmental variation (accounting for diapause and voltinism) and performed a laboratory experiment assaying thermal sensitivities of growth rates. The computer model showed opposing latitudinal trends in among- and within-generation thermal variability: within-generation thermal variability decreased toward higher latitudes, whereas relative levels of among-generation thermal variability peaked at midlatitudes (where a shift in voltinism occurred). The growth experiment showed that low-latitude species were more thermally generalized than mid- and high-latitude species, supporting the prediction that generalists are favored under high levels of within-generation variation. Northern species had steeper, near-exponential reaction norms suggestive of thermal specialization. However, they had strikingly high thermal optima and grew very slowly over most of the thermal range they are expected to experience in the field. This observation is at present difficult to explain. These results highlight the importance of considering interactions between life history and environmental variation when deriving expectations of thermal adaptation.     

Impact of temperature on juvenile growth and age at first egg-laying of the Pacific reef squid Sepioteuthis lessoniana reared in captivity

Cephalopod mollusks exhibit highly plastic life cycle traits influenced primarily by the interactive effects of food availability, light cycle and temperature, with the latter perhaps the most influential. Hatchlings of the tropical reef squid Sepioteuthis lessoniana were hatched from field-collected eggs in the laboratory and cultured at different temperatures to evaluate the effect of temperature on growth rates. All groups showed rapid, sustained growth rates from hatching to a size of 10–25 g. Beyond this size range, growth was slower and not clearly exponential in form. Growth rate was closely linked to temperature. Squids grown at approximately 27 °C attained a size of 10 g in as little as 45 days at sustained growth rates of 12.2% body weight day⁻¹ (%bw day⁻¹), while squids cultured at 20 °C required almost 100 days to attain the same size at rates of 5.7%bw day⁻¹. At an age of 55 days and approximately 1 g body weight, juvenile squids cultured at 20 °C were able to accelerate growth rates from 5.7%bw day⁻¹ to over 12%bw day⁻¹ when temperature was raised to 27 °C. They maintained this growth rate to a size of about 10 g and an age of at least 75 days post-hatching, indicating that body size and not age is the limiting factor for this rapid post-hatching growth. By comparison, conspecifics cultured near 27 °C from hatching had shifted out of the rapid post-hatching growth phase by day 50 at sizes between 10 and 50 g. The hatchlings from temperate to subtropical Japan had consistently higher growth rates at comparable temperatures than hatchlings from tropical Okinawa. When plotted as growth rate versus temperature, the Japanese group had a clearly higher slope to the relationship than the tropical populations, equivalent to a 2%bw day⁻¹ difference in growth rate at 25 °C. Age at first egg-laying was decreased at higher culture temperatures; however, overall life span was not. Received: 21 February 2000 / Accepted: 6 September 2000     

Familiarity with adults,but not relatedness,affects the growth of juvenile red-backed salamanders (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>)

The relative roles of kinship and familiarity in affecting an individual’s growth and fitness are not easy to disentangle. Not only is an individual more likely to have prior behavioral interactions with conspecifics in close proximity, it may also be related (in terms of kinship) to those nearby conspecifics. While some studies have inferred that kin discrimination affects fitness correlates, other studies found that familiarity alone can reduce aggressive interactions, thus increasing fitness. These studies have all focused on intra-age class pairs or groups. However, many animals interact with conspecifics from different cohorts. In many populations, adults of Plethodon cinereus territorially defend rocks and logs that retain moisture and food resources. We investigated whether juveniles of P. cinereus grew more in the presence of adults that were relatives or familiar. We collected pairs of juveniles and adults found under the same cover objects in the forest (familiar) and pairs of juveniles and adults found under different cover objects, approximately 10 m apart (unfamiliar). We determined parentage and relatedness of the adult–juvenile pairs and then placed these pairs in semi-natural mesocosms for 17 days. We found that juveniles housed with familiar adults had significantly greater increases in mass and snout–vent length than juveniles housed with unfamiliar adults in 2006 but not in 2007. Relatedness had no effect on growth. In addition, juveniles cohabitating with adults were not more likely to be their offspring. At least under certain environmental conditions, familiarity with adults, independent of relatedness or parentage, increased the growth of juvenile salamanders.     

The effects of temperature on the growth of juvenile scleractinian corals

Tropical reef corals are well known for their sensitivity to rising temperature, yet surprisingly little is known of the mechanisms through which temperature acts on intact coral colonies. One such mechanism recently has been suggested by the association between the growth of juvenile corals and seawater temperature in the Caribbean, which suggests that temperature causes a transition between isometric and allometric growth scaling in warmer versus cooler years, respectively (Edmunds in Proc R Soc B 273:2275–2281, 2006). Here, this correlative association is tested experimentally for a cause-and-effect relationship. During April and May 2006, juvenile colonies (8–35 mm diameter) of massive Porites spp. from Moorea, French Polynesia, were incubated at warm (27.8°C) and cool (25.7°C) temperatures for 15 days, and their response assessed through the scaling of growth (change in weight) with colony size. The results reveal that the scaling of colony-specific growth (mg colony⁻¹ day⁻¹) was unaffected by temperature, although growth absolutely was greater at the cool compared to the warm temperature, regardless of colony size. This outcome was caused by contrasting scaling relationships for area-specific growth (mg cm⁻² day⁻¹) that were negatively allometric under warm conditions, but independent of size under cool conditions. In April 2007, a 22 days field experiment confirmed that the scaling of area-specific growth in juvenile Porites spp. is negatively allometric at a warm temperature of 29.5°C. Based on strong allometry for tissue thickness, biomass, and Symbiodinium density in freshly collected Porites spp., it is hypothesized that the temperature-dependency of growth scaling in these small corals is mediated by the interaction of temperature with biomass.     

Bioenergetics of ephyra larvae of the scyphozoan jellyfish Aurelia aurita in relation to temperature and salinity

The body mass of Aurelia aurita ephyrae was better correlated with the diameter of the central disc than with the distance between opposite rhopaliae or distance between opposite lappet tips. Body dry weight (y, in μg) related to the disc diameter (x, in mm) through the equation y = 22.33 x ^1.99. The exponent 1.99 was significantly lower than that for the medusa stage, indicating a tendency to grow in diameter rather than in weight through the ephyra stage. The average ash-free dry weight (AFDW) of ephyrae was 38.0% of the dry weight. The AFDW/diameter relationship was used to convert measured diameters to body AFDW and calculate succession in body mass, daily ration, daily growth rate and gross growth efficiency. Effects of temperature (6, 9.5, 12, 15 and 18 °C) and salinity (17.5, 22, 26, 30.5 and 35 PSU) on these parameters and feeding were studied at saturated prey concentration (222 Artemia nauplii l⁻¹ initial concentration) by daily measurements over 10 d. There was a strong effect of temperature for total ingestion, growth rate, growth efficiency and final body mass of individual ephyrae, whereas the daily ration was not significantly different between the different temperatures. The experimental group kept at the highest temperature (18 °C) diverged the most, and ephyrae at this temperature ingested 2.7 times more and increased in weight 5.4 times more than at 6 °C. The average daily growth rate and gross growth efficiency of these ephyrae were 34.5% and 25.1%, respectively, significantly higher than at 6, 9.5 and 15 °C. Significant effects of salinity were shown for total ingestion, daily ration, daily growth rate and final weight, although only total ingestion and daily ration diverged sufficiently to show effects in a post-hoc test. This test showed that total ingestion was significantly different for all salinities except between 22 and 35 PSU and between 17.5 and 26 PSU. The daily ration for 35 PSU diverged from all other salinities, whereas none of the other salinities showed any significant differences. Thus, provided food in excess A. aurita can double its weight every 2 to 4 d, dependent on temperature and can therefore develop to the medusa stage in short time. Differences in environmental salinity in the range 17.5 to 35 PSU have little or no effect on growth rate and growth efficiency, whereas our results indicate that the full seawater salinity (35 PSU) causes significantly higher ingestion rate compared to lower salinities. Received: 11 January 1999 / Accepted: 11 May 1999     

Parameter estimation for distributed delay based population models from laboratory data: egg hatching of Oulema duftschmidi Redthenbacher (Coleoptera,Chrysomelidae) as an example

The cohort development of poikilotherms under favorable temperature conditions may be described by the time-invariant distributed delay models that require three parameters, i.e. the thermal threshold T₀, the thermal constant W and the variability parameter H representing distributed maturation times, also known as time distributed or stochastic development. Here, a parsimonious method is developed that uses stage-frequency matrices obtained under controlled conditions. The analysis of these matrices permits the estimation of the three parameters, while the incorporation of both developmental threshold and thermal constant into the time-invariant distributed delay model permits the representation of stochastic cohort development under constant temperatures. The evaluated parameters can also be used in time-varying distributed delay models that are particularly useful for a wide range of fluctuating temperature conditions.We consider stage-frequency matrices obtained from cohorts of eggs and larvae of Oulema duftschmidi Redthenbacher at different constant temperatures. A visual inspection of the matrices clearly shows the temperature dependent as well as the time distributed passage from the egg to the larval stage. Under the current range of temperatures, a linear model for developmental rate satisfactorily describes egg development, and estimates the thermal threshold (T₀=11.2 °C), the thermal constant (W=81.3 day-degrees) and the delay order parameter (H=70). The resulting model can be used to represent the development of cohorts of O. duftschmidi eggs in a favorable temperature range.     

Asymptotic growth trajectories of larval sardine (Sardinops melanostictus ) in the coastal waters off western Japan

Growth trajectories of individual larvae of Japanese sardine, Sardinops melanostictus, caught in the coastal waters off western Japan were back-calculated from the first feeding stage up to date of capture (approximate size of 20 to 35 mm total length; TL) based on individually determined allometric relationships between otolith daily ring radii and fish total lengths. The larvae in January-, February-, and March-hatched cohorts in the coastal waters grew faster and more uniformly than those in the oceanic waters offshore of the Kuroshio current. Growth trajectories of the three hatch-month cohorts were similar and could be expressed by the Gompertz model. The inflection points of the growth curves were reached at 9 to 11 d after hatching, when larvae were 10.8 to 11.8 mm TL. Maximum growth rates at these points were 0.80 to 0.85 mm d⁻¹. Growth rates gradually declined after the inflection points, and larval TLs converged into the infinite length of 29 to 32 mm, the sizes at which metamorphosis from larvae to juveniles is initiated. This asymptotic growth pattern in the larval stage resulted in the narrow ranges in TLs in spite of the wide range of ages of the larvae caught by boat seiners in the coastal waters. Slow growth and therefore long duration of the metamorphosing stage could be influential in determining the cumulative total mortality in the early life stages of the Japanese sardine. Received: 14 July 1996 / Accepted: 20 August 1996     

Can marine mammals be used to monitor oceanographic conditions?

The breeding performance of higher predators has often been used to monitor fluctuations in the abundance of important prey stocks in marine ecosystems. The development of electronic data-loggers in recent years has also provided the opportunity of using wide-ranging marine animals to measure physical oceanographic conditions. In this study, time–depth recorders (TDRs) programmed to record temperature were deployed on female Antarctic fur seals (Arctocephalus gazella) at Bird Island, South Georgia (54°00′S; 38°02′W) during the breeding seasons 1994 to 1998. Temperature sensors had relatively slow response times, and thermal radiation errors occurred during the day when seals spent a large proportion of their time at the surface. Nevertheless, measurements provided temperature–depth profiles which were typical of the vertical stratification of the ocean. During the early stages of a foraging trip temperature increased, suggesting that fur seals travelled northwards from South Georgia towards the warmer waters of the Polar Front. In addition, higher temperatures were recorded by females that remained at sea for longer, implying that these individuals also travelled further. Mean sea-surface temperature (SST) increased from ∼1 to 4 °C from December to March and agreed with SSTs from ship, buoy and satellite. Future studies on marine mammals which combine satellite tracking with oceanographic measurements are likely to provide valuable information on biophysical aspects of the ocean. Received: 16 June 1998 / Accepted: 13 February 1999     

The effects of temperature and salinity on reproductive success of <Emphasis Type="Italic">Temora longicornis</Emphasis> in the Baltic Sea: a copepod coping with a tough situation

At specific locations within the Baltic Sea, thermoclines and haloclines can create rapid spatial and temporal changes in temperature (T) and salinity (S) exceeding 10°C and 9 psu with seasonal ranges in temperature exceeding 20°C. These wide ranges in abiotic factors affect the distribution and abundance of Baltic Sea copepods via species-specific, physiological-based impacts on vital rates. In this laboratory study, we characterized the influence of T and S on aspects of reproductive success and naupliar survival of a southwestern Baltic population of Temora longicornis (Copepoda: Calanoida). First, using ad libitum feeding conditions, we measured egg production (EP, no. of eggs female⁻¹ day⁻¹) at 12 different temperatures between 2.5 and 24°C, observing the highest mean EP at 16.9°C (12 eggs female⁻¹ day⁻¹). Next, the effect of S on EP and hatching success (HS, %) was quantified at 12°C for cohorts that had been acclimated to either 8, 14, 20 or 26 psu and tested at each of five salinities (8, 14, 20, 26 and 32 psu). The mean EP was highest for (and maximum EP similar among) 14, 20 and 26 psu cohorts when tested at their acclimation salinity whereas EP was lower at other salinities. For adults reared at 8 psu, a commonly encountered salinity in Baltic surface waters, EP was relatively low at all test salinities—a pattern indicative of osmotic stress. When incubated at 12°C and 15 different salinities between 0 and 34 psu, HS increased asymptotically with increasing S and was maximal (82.6–84.3%) between 24 and 26 psu. However, HS did depend upon the adult acclimation salinity. Finally, the 48-h survival of nauplii hatched and reared at 14 psu at one of six different temperatures (10, 12, 14, 16, 18 and 20°C) was measured after exposure to a novel salinity (either 7 or 20 psu). Upon exposure to 7 psu, 48-h naupliar mortality increased with increasing temperature, ranging from 26.7% at 10°C to 63.2% at 20°C. In contrast, after exposure to 20 psu, mortality was relatively low at all temperatures (1.7% at 10°C and ≤26.7% for all other temperatures). An intra-specific comparison of EP for three different T. longicornis populations revealed markedly different temperature optima and clearly demonstrated the negative impact of brackish (Baltic) salinities. Our results provide estimates of reproductive success and early survival of T. longicornis to the wide ranges of temperatures and salinities that will aid ongoing biophysical modeling examining climate impacts on this species within the Baltic Sea.     

Physiological ecology of the clonal corallimorpharian <Emphasis Type="Italic">Corynactis californica</Emphasis>

For clonal taxa, the reduced genetic variability associated with clonal proliferation is hypothesized to reduce the ability to respond to variable conditions, unless a general-purpose genotype (GPG) confers success in multiple environments. In this study, Corynactis californica (Carlgren 1936) from the subtidal of California was used as a model system to test the hypothesis that clones dampen fluctuations in fitness through a GPG that facilitates phenotypic plasticity. To achieve this goal, tissue composition, respiration, excretion, and growth were compared among clones of C. californica at one site, and a reciprocal transplant experiment was used to test the response of clones to differing conditions at two sites. All experiments were completed at Santa Catalina Island (N 33°25′, W 118°30′) between April and September 1991. Clones at a single site differed significantly in multiple traits, varying as much as 1.6-fold in protein content, 3.4-fold in respiration, and 3.5-fold in excretion. Interestingly, while tissue growth was the most labile trait (differing up to 35.4-fold among clones), polyp fission rates were not significantly different among clones, in part because fission continued even though tissue growth was unable to restore polyp size in between divisions. Partial energy budgets revealed that the majority (47–90%) of the daily energy expenditure was accounted for by respiration, 13–47% by growth, and 0.3–14% by excretion. In the transplant experiment, reaction norms revealed strong effects of the environment on some traits but not others, notably with growth differing between sites in a pattern that differed among clones, and excretion differing between sites; neither respiration nor fission were affected by transplantation. Partial energy budgets revealed that the energy allocation to respiration varied between sites in a pattern that differed among clones, and a similar trend was evident for tissue growth. Together, these results demonstrate that clones of C. californica have markedly different phenotypes and exploit phenotypic plasticity to maintain relatively constant fission rates, even though tissue growth varies greatly among clones and between environments. While these findings support the GPG hypothesis for clones of C. californica—at least based on relative fitness achieved through asexual proliferation—this conclusion depends on the extent to which polyps are successful when they have low rates of tissue growth.     

Temporal genetic variation in populations of the limpet Cellana grata from Hong Kong shores

Variations in the relative contributions of gene flow and spatial and temporal variation in recruitment are considered the major determinants of population genetic structure in marine organisms. Such variation can be assessed through repeated measures of the genetic structure of a species over time. To test the relative importance of these two phenomena, temporal variation in genetic composition was measured in the limpet Cellana grata, among four annual cohorts over 10 years at four rocky shores in Hong Kong. A total of 408 limpets, comprising individuals from 1998, 1999, 2006 and 2007 cohorts were screened for genetic variation using five microsatellite loci. Minor but significant genetic differentiation was detected among samples from the 1998/1999 collection (F _ST = 0.0023), but there was no significant differentiation among the 2006/2007 collection (F _ST = 0.0008). Partitioning of genetic variation among shores was also significant in 1998/1999 but not in the 2006/2007 collection, although there was no correlation between genetic and geographic distances. There was no significant difference between collections made in 1998/1999 and 2006/2007. This lack of clear structure implies a high level of gene flow, but differentiation with time may be the result of stochastic recruitment variation among shores. Estimates of effective population size were not high (599, 95% C.L. 352–11397), suggesting the potential susceptibility of the populations to genetic drift, although a significant bottleneck effect was not detected. These findings indicate that genetic structuring between populations of C. grata in space and time may result from spatio-temporal variation in recruitment, but the potential development of biologically significant differentiation is suppressed by a lack of consistency in recruitment variability and high connectivity among shores.     

Demography and reproductive biology ofMelinna palmata (Ampharetidae: Polychaeta) in Inner Galway Bay on the west coast of Ireland

A high-density population of the polychaeteMelinna palmata Grube was the focus of a two-year study at a shallow-water location in Inner Galway Bay on the west coast of Ireland. Mean densities were highest in the second year of the study following successful recruitment. Size-frequency analysis of population structure (April 1983 to March 1985) revealed the presence of four cohorts, with the 1983 settlement becoming the dominant modal class. Size comparison of this cohort with the 1982 settlement showed that growth rates were lower and mortality higher in the 1983 settlement, which may be related to density dependent intra-specific competition.M. palmata at this location live for 2 to 2.5 yr with a small number surviving to 3 yr of age.M. palmata in Galway Bay is dioecious and has an even sex ratio. Reproductively mature worms were approximately 2 yr old when they first spawned. Spawning took place between May and July, when mature oocytes had a modal size of ca. 290µm. Proliferation of previtellogenic oocytes (ca. 10µm) from the gonads occurred following spawning. Growth over the winter was slow and was followed by a period of rapid growth between March and May, coincident with rising water temperatures. The presence of mature unspawned oocytes, which were resorbed after commencement of new proliferation, was taken as evidence of the species propensity for polytely. However, post-spawning mortality appeared to be high, so that only a small number of individuals survived to spawn the following year. The highest abundance of mature males was encountered between May and August, but lower levels of mature stages were present throughout the year.