The influence of temperature on the development of Baltic Sea sprat (Sprattus sprattus) eggs and yolk sac larvae

In spring 2004 and 2005 we performed two sets of experiments with Baltic sprat (Sprattus sprattus balticus Schneider) eggs and larvae from the Bornholm Basin simulating ten different temperature scenarios. The goal of the present study was to analyse and parameterise temperature effects on the duration of developmental stages, on the timing of important ontogenetic transitions, growth during the yolk sac phase as well as on the survival success of eggs and early larval stages. Egg development and hatching showed exponential temperature dependence. No hatching was observed above 14.7°C and hatching success was significantly reduced below 3.4°C. Time to eye pigmentation, as a proxy for mouth gape opening, decreased with increasing temperatures from 17 days post hatch at 3.4°C to 7 days at 13°C whereas the larval yolk sac phase was shortened from 20 to 10 days at 3.8 and 10°C respectively. Maximum survival duration of non-fed larvae was 25 days at 6.8°C. Comparing the experimental results of Baltic sprat with existing information on sprat from the English Channel and North Sea differences were detected in egg development rate, thermal adaptation and in yolk sac depletion rate (YSDR). Sprat eggs from the English Channel showed significantly faster development and the potential to develop at temperatures higher than 14.7°C. North Sea sprat larvae were found to have a lower YSDR compared to larvae from the Baltic Sea. In light of the predictions for global warming, Baltic sprat stocks could experience improved conditions for egg development and survival.     

Impacts of temperature and acidification on larval calcium incorporation of the spider crab <Emphasis Type="Italic">Hyas araneus</Emphasis> from different latitudes (54° vs. 79°N)

The combined effects of ocean warming and acidification were compared in larvae from two populations of the cold-eurythermal spider crab Hyas araneus, from one of its southernmost populations (around Helgoland, southern North Sea, 54°N, habitat temperature 3–18°C; collection: January 2008, hatch: January–February 2008) and from one of its northernmost populations (Svalbard, North Atlantic, 79°N, habitat temperature 0–6°C; collection: July 2008, hatch: February–April 2009). Larvae were exposed to temperatures of 3, 9 and 15°C combined with present-day normocapnic (380 ppm CO₂) and projected future CO₂ concentrations (710 and 3,000 ppm CO₂). Calcium content of whole larvae was measured in freshly hatched Zoea I and after 3, 7 and 14 days during the Megalopa stage. Significant differences between Helgoland and Svalbard Megalopae were observed at all investigated temperatures and CO₂ conditions. Under 380 ppm CO₂, the calcium content increased with rising temperature and age of the larvae. At 3 and 9°C, Helgoland Megalopae accumulated more calcium than Svalbard Megalopae. Elevated CO₂ levels, especially 3,000 ppm, caused a reduction in larval calcium contents at 3 and 9°C in both populations. This effect set in early, at 710 ppm CO₂ only in Svalbard Megalopae at 9°C. Furthermore, at 3 and 9°C Megalopae from Helgoland replenished their calcium content to normocapnic levels and more rapidly than Svalbard Megalopae. However, Svalbard Megalopae displayed higher calcium contents under 3,000 ppm CO₂ at 15°C. The findings of a lower capacity for calcium incorporation in crab larvae living at the cold end of their distribution range suggests that they might be more sensitive to ocean acidification than those in temperate regions.     

Elevated temperature elicits greater effects than decreased pH on the development, feeding and metabolism of northern shrimp (Pandalus borealis) larvae

Climate models predict that the average temperature in the North Sea could increase 3–5 °C and surface-waters pH could decrease 0.3–0.5 pH units by the end of this century. Consequently, we investigated the combined effect of decreased pH (control pH 8.1; decreased pH 7.6) and temperature (control 6.7 °C; elevated 9.5 °C) on the hatching timing and success, and the zoeal development, survival, feeding, respiration and growth (up to stage IV zoea) of the northern shrimp, Pandalus borealis. At elevated temperature, embryos hatched 3 days earlier, but experienced 2–4 % reduced survival. Larvae developed 9 days faster until stage IV zoea under elevated temperature and exhibited an increase in metabolic rates (ca 20 %) and an increase in feeding rates (ca 15–20 %). Decreased pH increased the development time, but only at the low temperature. We conclude that warming will likely exert a greater effect on shrimp larval development than ocean acidification manifesting itself as accelerated developmental rates with greater maintenance costs and decreased recruitment in terms of number and size.     

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.     

Larval development rate predicts range expansion of an introduced crab

Introduced populations can cause ecological and economic damage and are difficult to eradicate once they have established. It is therefore important to be able to predict both where species may become established and their capacity to spread within recipient regions. Here, we use a new method to assess potential for intraregional spread of a marine crab introduced to North America, Carcinus maenas. We determined survivorship and development rates throughout a range of temperatures in the laboratory for C. maenas larvae from non-native populations on the Atlantic and Pacific coasts of North America. The larvae exhibited narrower physiological tolerances than adults, and no lab-cultured larvae completed larval development below 10.0°C or above 22.5°C. Survivorship peaked at intermediate water temperatures of 12.5–20.0°C, and development time decreased with increasing temperatures within this range. Based upon these laboratory development rates, we used nearshore sea-surface temperature data from both coasts of North America to predict development times required for larvae at different months and sites. Taken together, survivorship and development data indicate that C. maenas has the capacity to continue its northward spread and establish populations at numerous additional sites in North America. Moreover, decadal temperature data at two Alaskan sites predicted little variability in development duration across years, suggesting that development duration predictions are robust to interannual water temperature differences.     

Thermal tolerance of early development in tropical and temperate sea urchins: inferences for the tropicalization of eastern Australia

The thermal envelope of development to the larval stage of two echinoids from eastern Australia was characterized to determine whether they fill their potential latitudinal ranges as indicated by tolerance limits. The tropical sand dollar, Arachnoides placenta, a species that is not known to have shifted its range, was investigated in Townsville, northern Australia (19°20′S, 146°77′E), during its autumn spawning season (May 2012). The subtropical/temperate sea urchin, Centrostephanus rodgersii, a species that has undergone poleward range expansion, was investigated in Sydney, southern Australia (33°58′S, 151°14′E), during its winter spawning season (August 2012). The thermal tolerance of development was determined in embryos and larvae reared at twelve temperatures. For A. placenta, the ambient water temperature near Townsville and experimental control were 24 °C and treatments ranged from 14 to 37 °C. For C. rodgersii, ambient Sydney water temperature and experimental control were 17 °C, and the treatment range was 9–31 °C. A. placenta had a broader developmental thermal envelope (14 °C range 17–31 °C) than C. rodgersii (9 °C range 13–22 °C). Both species developed successfully at temperatures well below ambient, suggesting that cooler water is not a barrier to poleward migration for either species. Both species presently live near the upper thermal limits for larval development, and future ocean warming could lead to contractions of their northern range limits. This study provides insights into the factors influencing the realized and potential distribution of planktonic life stages and changes to adult distribution in response to global change.     

The invasive gastropod Crepidula fornicata: reproduction and recruitment in the intertidal at its northernmost range in Wales, UK, and implications for its secondary spread

The establishment and spread of a non-native species in an introduced range depends to a large extent on the performance of the species under the prevailing environmental conditions. The spawning, larval and spatfall periods of the invasive gastropod Crepidula fornicata were monitored in the intertidal zone at its northernmost range in Wales, UK, between February 2010 and January 2011. The duration of the reproductive season was similar to that recorded from more southerly European populations. Spawning and larval release occurred throughout most of the year even at low seawater temperatures of <7 °C, but benthic recruitment was observed over a much shorter period at seawater temperatures >16 °C. Recruitment was low and likely controlled by post-settlement mortality. These observations suggest that C. fornicata’s northwards spread in Welsh waters will not be limited by seawater temperature negatively affecting reproduction, but by processes acting after larval release. These data show the importance of incorporating settlement and post-settlement processes into studies on recruitment success when aiming to predict the potential spread of a potentially harmful invader such as C. fornicata.     

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.     

Energy budgets reveal equal benefits of varied migration strategies in northern gannets

We investigated migration and wintering of adult northern gannets (Morus bassanus) breeding in east Scotland, North Sea, by deploying geolocation loggers over three winters. The wintering ranges of these birds varied from the North Sea to the Atlantic off West Africa. Flight time was taken as a proxy for migration and foraging effort. Gannets wintering off Africa had higher total flight times during migration than birds wintering further north. Total flight times in different wintering regions were generally low. Birds off West Africa consistently spent < 25 % of daylight hours in flight, but birds further north showed more variable values that may reflect more variable weather or food availability. Winter sea surface temperatures ranged from 9 °C (North Sea) to 16 °C (West Africa). Thermostatic costs in winter as estimated by measuring thermal conductance in carcasses in still air and water were 28 % higher in North Sea than off West Africa. This effect is aggravated by higher thermostatic costs caused by stronger wind chills in the North Sea compared to the conditions off West Africa. Birds wintering close to the UK arrived at the colony on average 12 days earlier than birds wintering off West Africa. We conclude that the net cost-benefit analysis may be similar for all wintering areas investigated.     

Strengthening recruitment of exploited scallops Pecten maximus with ocean warming

There is evidence that ocean warming has effects on the ecology, including recruitment dynamics, of marine organisms. In association with rising mean spring temperatures in the Irish Sea, a time-series of juvenile scallop Pecten maximus density around the Isle of Man showed a significant increasing trend since 1991. Favorable conditions (warmer water and correspondingly greater food availability) during gonad development can increase scallop gamete production. We examined the possibility that ocean warming has directly increased recruitment of exploited P. maximus around the Isle of Man by enhancing gonad development. From 1991–2007, there was a significant positive correlation between scallop recruitment and mean spring (the main period of gonad development) temperature in the year of larval settlement. Detrended (i.e., accounting for a time effect) recruitment data showed a marginally non-significant correlation to temperature. Gonadal somatic index of adult scallops and temperature were positively correlated. These relationships support the hypothesis that greater gamete production associated with ocean warming may be primarily responsible for observed increases in recruitment success and CPUE in a commercially important shellfish stock.     

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.     

Effect of temperature on viability and axial muscle development in embryos and yolk sac larvae of the Northeast Arctic cod (Gadus morhua)

Cod (Gadus morhua L.) eggs may develop and hatch within temperatures of −1.5 to 12 °C, but little is known about the effects of very low temperatures on larval characteristics. Eggs of the Northeast Arctic cod (Gadus morhua) were incubated at 1, 5 or 8 °C from Day 1 after fertilisation until hatching, and transferred to 5 °C after hatching. Histological samples of the axial musculature were taken at hatching and 5 d after hatching, and the data on muscle cellularity from these samples were related to survival and hatching, size, developmental data and viability of the yolk sac larvae. All larvae hatched at the same developmental stage. Incubation of eggs at 1 °C produced shorter larvae with a larger yolk sac and more, small deep fibres at hatching than larvae from eggs incubated at 5 or 8 °C. The larval size difference was still present 5 d after hatching, a time at which the larvae from 1 °C-incubated eggs were less developed and less resistant to an acute viability stress test (65 ppt salinity). Although there were no differences between temperature groups in number and size of muscle fibres 5 d after hatching, the deep fibres of the 1 °C-group contained less myofibrils than the two other groups. The phenotype of the larvae at hatching was thus affected within these incubation temperatures. Although all groups were transferred to the same temperature after hatching, the lowest egg incubation temperature (1 °C) still had a negative effect 5 d after hatching, as these larvae were both smaller, less resistant to stress and had less functional muscles at the time of first feeding. Our conclusion is therefore that 1 °C is close to, or below, the lower thermal tolerance limit for normal functional development of Northeast Arctic cod. The results are discussed in relation to larval viability and recruitment of this species in the wild. Received: 4 February 1998 / Accepted: 10 July 1998     

Effect of development rate on the swimming, escape responses, and morphology of yolk-sac stage larval American plaice, Hippoglossoides platessoides

To examine the impact of development rate on swimming performance, escape response, and morphology, yolk-sac larvae of American plaice (Hippoglossoides platessoides, Fabricius) were reared at two temperatures (5 and 10 °C). Videomicroscopy and silhouette collimation videography were used to examine swimming, escape behaviour, and morphology (standard length, finfold area, and yolk-sac area) of individual larvae. Larvae were examined from 0 d post hatch (dph) to 14 dph for the 5 °C treatment group and from 0 to 6 dph for the 10 °C treatment group (3 August to 17 August 1996). Since larvae were not fed, yolk-sac reserves were essentially exhausted by 14 and 6 dph for the 5 and 10 °C treatment groups, respectively. To control for the effect of testing temperature on behaviour, larvae from each temperature treatment were tested at both 5 and 10 °C. Testing temperature had an effect on some swimming parameters but not on escape response. Swimming performance, escape response, and morphology varied with age, while only morphology and escape response varied with development rate. Morphology and swimming performance, and morphology and escape response were found to be correlated as determined by canonical correlation. This study suggests that both types of swimming behaviours should be examined when developing models of the impacts of predation on the early life history of larval fish. Received: 13 September 1999 / Accepted: 21 June 2000     

Metabolic rate and lipofuscin accumulation in juvenile European lobster (Homarus gammarus ) in relation to simulated seasonal changes in temperature

 The accumulation of lipofuscin, which is an indictor of physiological age, in the brain of juvenile European lobster (Homarus gammarus L.) was monitored for 22 mo in three experimental temperature regimes that simulated seasonal variation in temperature in the geographic range of this species. Metabolic rate responses to changes in temperature were estimated by measuring the activity of the electron transport system (ETS) in muscle tissue and in vivo rates of oxygen consumption. Lipofuscin accumulation oscillated with simulated seasonal changes in temperature and was described by seasonalised von Bertalanffy growth functions. The incremental accumulation in lipofuscin between sampling dates was linearly related to the number of degree days that accumulated between dates, irrespective of the amplitude of temperature fluctuation that had occurred. ETS activity increased with acclimation temperature and was modelled using a polynomial function. This indicated a lower temperature sensitivity in the temperature mid-range (12 to 16 °C), although the Q₁₀ for this mid-range was 2.1. ETS activity in lobsters acclimated to 8 and 18 °C and assayed at 13 °C was similar, indicating no compensation for changes in environmental temperature. Oxygen consumption rate was significantly higher at 14 °C than at 10.5 °C and had a Q₁₀ of 3.6, again suggesting no compensation to temperature change. The absence of metabolic compensation in response to temperature change in H. gammarus is consistent with the predictability of changes in temperature and food availability in the sub-littoral environment of this species. As lipofuscin accumulates according to metabolic rate, and metabolic rate in H. gammarus is directly correlated with temperature, geographic differences and long-term temporal trends in temperature will need to be considered when converting physiological age indices, obtained from lipofuscin estimates, to a chronological scale. Received: 27 April 2000 / Accepted: 21 July 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.     

Cephalopod hatchling growth: the effects of initial size and seasonal temperatures

Temperature is known to have a strong influence on cephalopod growth during the early exponential growth phase. Most captive growth studies have used constant temperature regimes and assumed that populations are composed of identically sized individuals at hatching, overlooking the effects of seasonal temperature variation and individual hatchling size heterogeneity. This study investigated the relative roles of initial hatchling size and simulated natural seasonal temperature regimes on the growth of 64 captive Octopus pallidus over a 4-month period. Initial weights were recorded, and daily food consumption and fortnightly growth monitored. Two temperature treatments were applied replicating local seasonal water temperatures: spring/summer (14–18°C) and summer/autumn (18–14°C). Overall octopuses in the spring/summer treatment grew at a rate of 1.42% bwd⁻¹ (% body weight per day) compared to 1.72% bwd⁻¹ in the summer/autumn treatment. Initial size influenced growth rate in the summer/autumn treatment with smaller octopuses (<0.25 g) growing faster at 1.82% bwd⁻¹ compared to larger octopuses at 1.68% bwd⁻¹. This was opposite to individuals in the spring/summer treatment where smaller octopuses grew slower at 1.29% bwd⁻¹ compared to larger octopuses at 1.60% bwd⁻¹. Initial size influenced subsequent growth, however, this was dependent on feeding rate and appears to be secondary to the effects of temperature.     

Der einfluß der temperatur auf oocytenentwicklung und eiablage bei dem interstitiellen archiannelidenTrilobodrilus axi

The interstitial archiannelidTrilobodrilus axi Westheide 1967 inhabits sandy beaches of the Island of Sylt (North Sea). Spawning occurs from May to July, approximately. Development of oocytes and oviposition occur as soon as the water temperature has risen above 5 °C. In rearing experiments, the influence of temperature on maturation and release of gametes was studied in the female. Animals were collected in winter and reared at temperatures between 3° and 15°C. Within this range, development of oocytes and shedding of eggs were accelerated progressively with increasing test temperatures. Successful reproduction occurred only at temperatures above a critical level (between 3° and 6 °C). It was possible to shift the time of spawning by several months under experimental conditions; this suggests that, in the field, spawning is mainly controlled by temperature. The results indicate thatTrilobodrilus axi is probably a boreo-mediterranean species.     

Behaviour moderates climate warming vulnerability in high-rocky-shore snails: interactions of habitat use, energy consumption and environmental temperature

High-rocky-shore intertidal animals are predicted to be generally more vulnerable to climate warming than lower-shore species, because their thermal tolerances lie closer to maximum environmental temperatures (T _e). However, this prediction is based on taxonomically and ecologically limited information. The present study investigated the effect of habitat use on climate warming vulnerability of the tropical high-shore snail, Echinolittorina malaccana (from Brunei Darussalam, 5°N), which aestivates in sun-exposed or shaded habitats. The thermal regimes of these habitats differed vastly, but snails showed similar daily energy consumption in either habitat, due to temperature-insensitive metabolism (TIM) between 35 and 46 °C in the sun-resting snails. However, maximum T _e values in the shade and the sun were 35 and 46 °C, respectively, suggesting that sun-resting snails, which presently experience temperatures near the incipient lethal temperature range (46–56 °C), should be more threatened by further warming than shade-resting snails, which have an 11 °C ‘safety margin’. Thus, vulnerability of high-shore species to climate warming could be moderated by availability of shaded habitat, making predictions for these species more complex than previously realized.     

Temperature acclimation of upper and lower thermal limits and freeze resistance in the nonindigenous brown mussel, Perna perna (L.), from the Gulf of Mexico

Acute and chronic upper and lower thermal limits and freeze resistance were investigated in the nonindigenous brown mussel, Perna perna, from the Texas Gulf of Mexico coast in order to assess its potential distribution in North American coastal waters. This species' long-term, incipient lower and upper thermal limits were 7.5°C and 30°C, congruent with the seasonal ambient water temperature range of 10-30°C reported for other populations worldwide. Effects of temperature acclimation and individual size on survival time were most pronounced on chronic exposures to lethal temperatures approaching incipient lower or upper thermal limits. When exposed to temperature increasing at 0.1°C min^-1, the acute upper lethal limit was 44°C regardless of acclimation temperature or individual size. P. perna had a limited freeze resistance, being intolerant of emersion at -2.5°C. This species' narrow incipient thermal limits, limited capacity for temperature acclimation and poor freeze resistance may account for its restriction to subtidal and lower eulittoral zones of cooler subtropical rocky shores. Near extinction of P. perna from Texas Gulf of Mexico waters occurred in the summer of 1997 when mean surface-water temperatures approached its incipient upper limit of 30°C.