Sex-specific temperature dependence of foraging and growth of intertidal snails

Predicting the biological impacts of climate change requires an understanding of how temperature alters organismal physiology and behavior. Given differences in reproductive physiology between sexes, increases in global temperature may be experienced differently by the males and females of a species. This study tested for sex-specific effects of increased air temperature on foraging, growth, and survival of an intertidal snail, Nucella ostrina (San Juan Island, Washington, 48–30′44″N, 123–08′43″W). Snails exhibited periodic peaks in foraging. Subjecting snails to elevated low tide air temperatures did not alter the timing or magnitude of this pattern. Despite similar temporal patterns in foraging, females foraged more than males, even when the risk of thermal stress was high. While males and females appear to have a similar body temperature threshold for optimal growth, females were more likely to cross that threshold resulting in a loss of body mass when exposed to daily increases in air temperature. These results suggest that the consequences of a warming climate in the short term may be different for males and females of N. ostrina, but also imply longer-term costs of reduced reproductive output, abundance, and distribution of this ubiquitous intertidal predator. Generally, this study points to the possible significance of sex-specific responses in an increasingly warm world.     

Effects of ocean acidification on the early developmental stages of the horned turban,Turbo cornutus

To estimate the impact of CO₂-driven ocean acidification on the early life stages of gastropods, the effects of increased partial pressure of seawater carbon dioxide (pCO₂) (800–2,000 μatm) on the early developmental stages and larval shell length of the commercially important gastropod, the horned turban snail, Turbo cornutus were investigated. Increase in experimental seawater pCO₂ had an increasingly negative impact on the early developmental rate; the proportion of embryos or larvae displaying retarded development increased at higher pCO₂. The proportion of embryos that developed to the 4-cell stage at 2 h after fertilization decreased linearly with increasing pCO₂. At ~1,000 μatm pCO₂, retarded development was observed in ~50 % of larvae. No embryos developed to the 4-cell stage at 2,000 μatm pCO₂ within 2 h of fertilization. A similar trend continued until 24–26 h after fertilization; the proportion of larvae attaining veliger stage by 24–26 h also decreased with increasing pCO₂. The shell length of T. cornutus veligers decreased gradually as seawater pCO₂ increased, but markedly decreased in seawater under nearly unsaturated and unsaturated conditions (≤1.04) of the aragonite saturation state (Ω _aragonite). The results indicate that increased pCO₂ seawater has a progressive and acute effect on embryonic and larval T. cornutus, and imply that the extended early developmental period and/or the downsized larval shell produced by ocean acidification will have a negative impact on survival, settlement and recruitment well into the future.     

Comparative larval development of the shipworms Bankia gouldi and Teredo navalis

Larvae of Bankia gouldi (Bartsch) and Teredo navalis L. were reared in the laboratory at various temperatures and salinities. T. navalis spawned at lower temperatures than B. gouldi. T. navalis larvae were released at temperatures from 13° to 30°C; in B. gouldi, spawning occurred from 17.5° to 30°C. Both species released offspring at salinities of 20 and 30‰. Larvae of the two species can be distinguished morphometrically at the earliest pelagic veliger stage and in the pediveliger stage. Average dimensions of newly released T. navalis larvae are 88 x 75 μ (length x height), while the youngest B. gouldi veligers measure 61 x 50 μ. Pediveligers of T. navalis (205 x 239 μ) are smaller than those of B. gouldi (221 x 260 μ). At other stages of larval development the two species appear so similar that they cannot yet readily be distinguished. Under laboratory conditions of 25°C and 30 ‰, the free-swimming life of B. gouldi to the pediveliger stage was about 10 days longer than that of T. navalis (25 and 15 days, respectively). Incubated larval development of T. navalis was estimated to be 5 days at 25°C. Potential competition between larval stages of the two species, and modification of settling behavior by dissolved humic material (Gelbstoff), is discussed.     

Thermotolerance and heat-shock protein expression in Northeastern Pacific<Emphasis Type="Italic"> Nucella</Emphasis> species with different biogeographical ranges

We investigated physiological traits responsible for determining the tide-height and latitudinal distributions of Northeastern Pacific Nucella congeners. First, we determined the thermotolerances of two species of intertidal dogwhelks, N. ostrina and N. canaliculata, which co-occur on the Oregon coast. We found that N. ostrina, which are distributed higher on the shore, and thus experience higher habitat temperatures, than N. canaliculata, had correspondingly higher heat-coma temperatures. Second, we acclimated individuals of all five Northeastern Pacific Nucella congeners to a common temperature and determined their thermotolerances, measured as recovery from thermal exposure, after a 5-day, 3-week, and 7-week acclimation period. The south-latitude (N. emarginata) and mid-latitude (N. ostrina) high-intertidal species were more thermotolerant than the mid-latitude low-intertidal (N. canaliculata and N. lamellosa) and north-latitude high-intertidal (N. lima) species. The results of these two experiments suggest that temperature plays a role in determining the tide-height and latitudinal distributions of these Nucella species. Finally, we measured total and inducible levels of an evolutionarily conserved and ecologically relevant protein, the 70-kDa heat-shock protein (Hsp70), which has been found to confer thermotolerance in model laboratory organisms. The results showed that the level of total, not stress inducible, Hsp70 was a better predictor of thermotolerance and that there were species-specific differences in the relationship between Hsp70 expression and thermotolerance. We suggest that Hsp70 expression may be important in conferring thermotolerance in Nucella species in nature and that higher levels of molecular chaperones may underlie increased thermotolerance between conspecifics.Communicated by P.W. Sammarco, Chauvin     

Latitudinal patterns of shell thickness and metabolism in the eastern oyster Crassostrea virginica along the east coast of North America

The goal of this study was to quantify growth and metabolic responses of oysters to increased temperatures like those that will occur due to global warming. Impact of temperature on eastern oyster (Crassostrea virginica) shell growth and metabolism was investigated by sampling 24 sites along the eastern North American seaboard ranging from New Brunswick, Canada, to Florida, USA, in March and August 2013. There was a positive correlation between oyster shell thickness and site temperature. At southern sites, shells were up to 65 % thicker than at the northernmost site, likely due to higher precipitation of CaCO₃ in warmer water. This was supported by laboratory experiments showing that thicker shells were produced in response to temperatures 2, 4, and 6 °C above ambient seawater temperatures (8–14 °C) in Connecticut, USA. Field experiments with oyster respiration were conducted during winter and summer at 13 sites to compare responses to thermal stress with latitude. Respiration rates were much higher during summer than winter, but the combination of summer and winter data fell along the same exponential curve with respect to temperature. At all sites, temperature-specific metabolic rates at elevated temperatures were lower than predicted, indicating significant seasonal acclimatization by C. virginica.     

Thermal tolerance of Strongylocentrotus purpuratus early life history stages: mortality, stress-induced gene expression and biogeographic patterns

In this study, we examined the differential thermal tolerance of Strongylocentrotus purpuratus early life history stages by comparing high temperature–induced mortality and the relative levels of the stress-induced gene, hsp70, between S. purpuratus embryos and larvae from adults collected throughout the species range. There was no significant difference between gastrulae and 4-arm plutei mortality from all sites examined. Furthermore, there was little variability in temperature tolerance across the biogeographic range as southern gastrulae and 4-arm plutei exhibited similar tolerances to northern individuals. Relative levels of hsp70 mRNA expression did not differ overall between the two developmental stages at each site. Across sites, all gastrulae and 4-arm plutei exhibited maximum hsp70 expression at approximately 25°C; however, the range of hsp70 expression was narrower in southern individuals, suggesting they are living closer to their upper thermal limit than northern individuals.     

The Arctic sea butterfly<Emphasis Type="Italic"> Limacina helicina</Emphasis>: lipids and life strategy

The sea butterfly Limacina helicina was collected from May to September 2001 in Kongsfjorden, Spitsbergen, to investigate population structure and body and lipid composition with regard to life cycle and reproductive strategy. Veligers and juveniles were only found in late autumn and spring, whereas females occurred from July to September. The size of the females increased until mid-August and decreased in September. Dry and lipid mass were closely related to size; dry mass increased exponentially and lipids linearly with size. The lipid content was highest in veligers (31.5% of dry mass) and juveniles (23.6%) but low in females (<10%). Phospholipids were the dominating lipid class followed by triacylglycerols. Females, veligers, and egg ribbons, all from September, were richest in phospholipids. Juveniles contained the highest amounts of triacylglycerols, whereas females had low levels in July and the beginning of August. In mid-August, levels of triacylglycerols were variable and higher. This suggests that females were in their main spawning period and the high variability in triacylglycerols points to different stages within the spawning cycle. Enhanced amounts of free fatty acids in females from July may be related to gonad development. The 16:1(n-7) fatty acid was more dominant in spring whereas 18:4(n-3) increased in summer and autumn, which reflects a change in diet from diatom-dominated food items in spring to dinoflagellates in summer/autumn. Small amounts of long-chain monounsaturated fatty acids suggest ingestion of copepods, and the fatty acid composition of veligers feeding on particulate matter. L. helicina has a one-year life cycle with peak spawning in August and over-winters as veligers that may grow to juveniles during the winter period. They metamorphose into juveniles during spring, develop to males in early summer, and mature into females in July and August.Communicated by M. Kühl, Helsingør     

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.     

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.     

Juvenile life stages of the brown alga Fucus serratus L. are more sensitive to combined stress from high copper concentration and temperature than adults

The combined effects of exposure to copper and temperature were investigated in adult specimens and germlings of the canopy-forming brown alga Fucus serratus. A matrix of four temperatures, 6, 12, 17 and 22 °C, and three concentrations of copper, 0, 100 and 1,000 nM total copper were used. Measured endpoints were growth rate, chlorophyll fluorescence parameters and for germlings also survival. The growth rate of adult specimens of F. serratus changed with increasing temperature. Growth tended to be negatively affected by high concentrations of copper when exposed to heat (22 °C) though not significantly so. The photosynthetic performance (i.e., chlorophyll fluorescence parameters: F _v/F _m, maximum electron transport rate (ETR_max) and maximum non-photosynthetic quenching (NPQ_max) of adults was largely unaffected by both copper and temperature. Germling survival, growth rate and chlorophyll fluorescence parameters were affected by the combination of copper concentration and temperature. Increasing temperature led to reduced survival, increased rhizoid growth and higher F _v/F _m and ETR_max, whereas high copper concentration had a negative effect on the latter three endpoints. The negative effect of high copper concentration was amplified by high temperature. We conclude that juveniles of F. serratus are more susceptible to environmental stressors than adult specimens and recommend therefore including early life stages when assessing the risk of exposure to toxic compounds. Considering the response of adult specimens only may lead to false conclusions regarding the ecological impact of environmental stress.     

Effects of mesh size on estimation of population densities of tropical euthecosomatous pteropods

Euthecosomatous pteropods were collected in successive oblique plankton tows with 1 m diameter nets equipped with No. 0, No. 6, and No. 20 nylon mesh. Comparisons of samples revealed that the same species were collected by all 3 nets, but the No. 0 and No. 6 nets failed to collect veligers and juveniles of Spiratella inflata and Creseis virgula conica. Collections with nets of larger mesh size may erroneously suggest seasonal variations in population densities due, not to actual changes in total density, but to changes in the size of individuals and changes in the relative numbers of the different life stages. A large diameter net equipped with No. 20 mesh is recommended for work that includes the smallest life stages of tropical pteropods.     

Growth rate of the bathypelagic crustacean Gnathophausia ingens (Mysidacea: Lophogastridae)

The content of water, ash, carbohydrate, lipid, protein, chitin, carbon, hydrogen, nitrogen and energy was measured for all life stages of the bathypelagic mysid Gnathophausia ingens collected in San Clemente Basin off Southern California, USA between January 1969 and February 1982. These data are used to examine the life history, growth rates, growth pattern and partitioning of material and energy over the life of this species. Females reproduce only once and brood their young for about 1.5 yr, during which time the females do not feed. This species has a very high reproductive effort: of the energy accumulated over its life, 61.3% is used in egg production, 13.4% in parental care of young, 5.6% in cast exoskeletons and only 19.6% remains in the females after brooding. The relative rate of growth (percentage of energy d^?1) is about 1% immediately after leaving the mother and declines to about 0.2% as maturity is approached. Such growth rates appear to be typical for an animal of this size living at low temperatures. The evolutionary context of this mysid's life history is discussed.     

Comparing the effect of elevated pCO2 and temperature on the fertilization and early development of two species of oysters

This study compared the synergistic effects of elevated pCO₂ and temperature on the early life history stages of two ecologically and economically important oysters: the Sydney rock oyster, Saccostrea glomerata and the Pacific oyster, Crassostrea gigas. Gametes, embryos, larvae and spat were exposed to four pCO₂ (375, 600, 750, 1,000 μatm) and four temperature (18, 22, 26, 30°C) levels. At elevated pCO₂ and suboptimal temperatures, there was a reduction in the fertilization success of gametes, a reduction in the development of embryos and size of larvae and spat and an increase in abnormal morphology of larvae. These effects varied between species and fertilization treatments with S. glomerata having greater sensitivity than C. gigas. In the absence of adaptation, C. gigas may become the more dominant species along the south-eastern coast of Australia, recruiting into estuaries currently dominated by the native S. glomerata.     

Egg and early larval stages of Baltic cod, Gadus morhua, are robust to high levels of ocean acidification

The accumulation of carbon dioxide in the atmosphere will lower the pH in ocean waters, a process termed ocean acidification (OA). Despite its potentially detrimental effects on calcifying organisms, experimental studies on the possible impacts on fish remain scarce. While adults will most likely remain relatively unaffected by changes in seawater pH, early life-history stages are potentially more sensitive, due to the lack of gills with specialized ion-regulatory mechanisms. We tested the effects of OA on growth and development of embryos and larvae of eastern Baltic cod, the commercially most important fish stock in the Baltic Sea. Cod were reared from newly fertilized eggs to early non-feeding larvae in 5 different experiments looking at a range of response variables to OA, as well as the combined effect of CO₂ and temperature. No effect on hatching, survival, development, and otolith size was found at any stage in the development of Baltic cod. Field data show that in the Bornholm Basin, the main spawning site of eastern Baltic cod, in situ levels of pCO₂ are already at levels of 1,100 μatm with a pH of 7.2, mainly due to high eutrophication supporting microbial activity and permanent stratification with little water exchange. Our data show that the eggs and early larval stages of Baltic cod seem to be robust to even high levels of OA (3,200 μatm), indicating an adaptational response to CO₂.     

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     

Ecological commonalities among pelagic fishes: comparison of freshwater ciscoes and marine herring and sprat

Systematic comparisons of the ecology between functionally similar fish species from freshwater and marine aquatic systems are surprisingly rare. Here, we discuss commonalities and differences in evolutionary history, population genetics, reproduction and life history, ecological interactions, behavioural ecology and physiological ecology of temperate and Arctic freshwater coregonids (vendace and ciscoes, Coregonus spp.) and marine clupeids (herring, Clupea harengus, and sprat, Sprattus sprattus). We further elucidate potential effects of climate warming on these groups of fish based on the ecological features of coregonids and clupeids documented in the previous parts of the review. These freshwater and marine fishes share a surprisingly high number of similarities. Both groups are relatively short-lived, pelagic planktivorous fishes. The genetic differentiation of local populations is weak and seems to be in part correlated to an astonishing variability of spawning times. The discrete thermal window of each species influences habitat use, diel vertical migrations and supposedly also life history variations. Complex life cycles and preference for cool or cold water make all species vulnerable to the effects of global warming. It is suggested that future research on the functional interdependence between spawning time, life history characteristics, thermal windows and genetic differentiation may profit from a systematic comparison of the patterns found in either coregonids or clupeids.     

Impact of ocean acidification in the metabolism and swimming behavior of the dolphinfish (Coryphaena hippurus) early larvae

Since the industrial revolution, [CO₂]_atm has increased from 280 μatm to levels now exceeding 380 μatm and is expected to rise to 730–1,020 μatm by the end of this century. The consequent changes in the ocean’s chemistry (e.g., lower pH and availability of the carbonate ions) are expected to pose particular problems for marine organisms, especially in the more vulnerable early life stages. The aim of this study was to investigate how the future predictions of ocean acidification may compromise the metabolism and swimming capabilities of the recently hatched larvae of the tropical dolphinfish (Coryphaena hippurus). Here, we show that the future environmental hypercapnia (ΔpH 0.5; 0.16 % CO₂, ~1,600 μatm) significantly (p < 0.05) reduced oxygen consumption rate up to 17 %. Moreover, the swimming duration and orientation frequency also decreased with increasing pCO₂ (50 and 62.5 %, respectively). We argue that these hypercapnia-driven metabolic and locomotory challenges may potentially influence recruitment, dispersal success, and the population dynamics of this circumtropical oceanic top predator.     

Temperature effects on vital rates of different life stages and implications for population growth of Baltic sprat

Baltic sprat (Sprattus sprattus balticus S.) is a key species in the pelagic ecosystem of the Baltic Sea. Most stocks of small pelagic species are characterized by natural, fishery-independent fluctuations, which make it difficult to predict stock development. Baltic sprat recruitment is highly variable, which can partly be related to climate-driven variability in hydrographic conditions. Results from experimental studies and field observations demonstrate that a number of important life history traits of sprat are affected by temperature, especially the survival and growth of early life stages. Projected climate-driven warming may impact important processes affecting various life stages of sprat, from survival and development during the egg and larval phases to the reproductive output of adults. This study presents a stage-based matrix model approach to simulate sprat population dynamics in relation to different climate change scenarios. Data obtained from experimental studies and field observations were used to estimate and incorporate stage-specific growth and survival rates into the model. Model-based estimates of population growth rate were affected most by changes in the transition probability of the feeding larval stage at all temperatures (+0, +2, +4, +6?°C). The maximum increase in population growth rate was expected when ambient temperature was elevated by 4?°C. Coupling our stage-based model and more complex, biophysical individual-based models may reveal the processes driving these expected climate-driven changes in Baltic Sea sprat population dynamics.     

Metal contamination increases the sensitivity of larvae but not gametes to ocean acidification in the polychaete Pomatoceros lamarckii (Quatrefages)

Ocean acidification is not happening in isolation but against a background of chronic low-level pollution for most coastal marine environments. The reproductive and larval stages of marine invertebrates can be highly sensitive to the impacts of both environmental pollutants and ocean acidification, but very little is currently known regarding the potential impacts of combined contaminant and high CO₂ exposures on the health of marine organisms. Ocean acidification research to date has focused heavily on the responses of calcifying marine invertebrate larvae and algae, and as such the polychaetes as a group, despite their ecological importance, remain understudied. Here, we investigate the effects of elevated seawater CO₂ (pH range 8.1–7.4, plus an extreme pH of 7.2 in the sperm motility experiments), in combination with the environmental pollutant copper (0.002 μM), on the early life history stages of the intertidal polychaete Pomatoceros lamarckii from two populations. P. lamarckii sperm appear to be robust to elevated seawater CO₂. Whilst all three of the sperm motility end points measured showed a response to elevated CO₂, these responses were small and not linear. The percentage of motile sperm and sperm curvilinear velocity were significantly reduced in the lower pH treatments of 7.4 and 7.2, whereas sperm straight-line velocity (VSL) was mostly unaffected except for an increased VSL at pH 8.0. Fertilisation success was investigated using two populations from the South West (UK), one from Torquay and one from Plymouth Sound. Fertilisation success was slightly but significantly reduced at the 7.6 and 7.4 pH treatments for both populations (a 9.0 % reduction in fertilisation success from pH 8.1 to 7.4 for Torquay), but with a greater effect observed in the population from Plymouth Sound (a 13.33 % reduction in fertilisation success). No additional impact of 0.002 μM copper exposure on fertilisation success was found. Larval survival was found to be much more sensitive to elevated CO₂ than sperm function or fertilisation, and a significant interaction with copper exposure was observed. These results demonstrate the potential for polychaete larvae to be affected by predicted ocean acidification conditions and that chronic coastal pollutants, such as copper, have the potential to alter larval susceptibility to ocean acidification conditions.     

Thermal tolerance of larvae of Pollicipes elegans, a marine species with an antitropical distribution

For the antitropical gooseneck barnacle Pollicipes elegans, population-specific physiological temperature tolerance of larvae may serve as a barrier to larval dispersal across the warmest regions of the tropical Pacific Ocean. Thermal tolerance ranges of larvae of three different populations of P. elegans sampled in 2011 and 2012 (Mexico [MX], El Salvador [ES], and Peru [PE]) were investigated by measuring three indicators of physiological performance: swimming activity, oxygen consumption, and lethality or LT₅₀. The thermal tolerance profiles, which include measurable optimum (maximum aerobic performance), pejus (“getting worse”) and pessimum (worst aerobic performance) ranges, of larvae from the three populations were consistent with their characteristic environmental temperatures. In MX, larvae live close to the upper border of their optimum during warm months and so have a limited capacity to tolerate higher-than-normal temperatures. Larvae from the ES population likewise appear to live within their optimum temperature range, but these larvae lack a detectable pessimum range, suggesting they would be unable to cope with temperatures above their pejus range. Larvae from PE have a broad optimum but no pejus range. Different thermal tolerance ranges provide strong evidence for population-dependent physiological adaptations in P. elegans. For the southern (PE) and northern (MX) P. elegans populations, high tropical temperatures are likely to be a strong direct physiological barrier to larval survival and dispersal, which is in contrast to the more thermally tolerant ES population.