Temperature tolerance and latitudinal range of brown algae from temperate Pacific South America

Lower and upper survival temperatures of microthalli of 25 species of South American Phaeophyceae isolated from central Peru (14°S) to the Canal Beagle (55°S) were determined using 2-wk exposure for the upper and 4-wk exposure for the lower limit. All species survive 4 wks at -2°C. With respect to the upper limit, species reported only from southernmost South America tolerate 19.9 to 24.5°C (n=8), and species occurring from Cape Horn to central Chile 24.6 to 27.4°C (n=7). Three species that occurred as far north as northern Chile and Peru before the 1982–1983 El Niño event, and whose northern limit was dramatically shifted southwards in 1983, tolerate 20.8 to 25.3°C, whereas five species that have survived in Peru tolerate 25.6 to 28.5°C. Tinocladia falklandica which tolerates 27.8 to 28.1°C but lives only in southernmost South America and Striaria attenuata, which tolerates 31.6 to 31.9°C but occurs at ca. 42°S, are exceptional. Their high temperature tolerance may have no adaptive value in South America. They are restricted to the cold-temperate region due to low temperature requirements for reproduction or for reasons yet unknown. In general, the northern distributional limits of the Phaeophyceae studied along the temperate Pacific coast of South America are reproduction boundaries, except in El Niño years when they are redefined according to the species' upper suvival limits. Temperature tolerance of isolates from northern Chile and Peru agrees well with maximum temperatures reached during the 1983 El Niño.     

Are direct developers more locally adapted than planktonic developers?

The hypothesis that populations of direct developers exhibit greater geographic differentiation in life history features than populations of planktonic developers, was tested with two species of grazing snails of the genus Littorina from 1986 to 1987. Littorina sitkana (direct developer) and L. scutulata (planktonic developer) coexist on sun- and wave-sheltered beaches from Alaska to Oregon, USA. Seasonal patterns in growth, survival and reproduction were monitored for samples from four geographically separated populations of each species grown in population cages at a common site, Friday Harbor, Washington, USA. The environmental and population effects on growth in the two species were determined in a four-way reciprocal transplant experiment with the same populations. Both the direct and planktonic developers exhibited geographic differentiation in life history features. Differentiation in the direct developer occurred over distances shorter than 30 km, while differentiation in the planktonic developer occurred over the 500 km distance examined (greater than their larvae would likely travel).     

Comparative population genetic structure of marine gastropods (Littorina spp.) with and without pelagic larval dispersal

 Population genetic theory predicts that marine animal species with planktonic larvae will have less genetic structure than those with direct development. We compared the genetic structure of four species of littorinid snails – two with planktonic egg capsules that hatch as planktonic larvae and two with benthic egg masses that hatch as crawl-away juveniles. We used DNA sequencing and single stranded conformational polymorphism (SSCP) to assess sequence variation in a 480 bp fragment of the mitochondrial cytochrome b gene and then used an analysis of molecular variance (AMOVA) to estimate Φ_st for populations from the northeastern Pacific coast. One of the two direct-developing species, Littorina subrotundata, had a moderate amount of population structure (Φ_st=0.209) as expected but the other direct-developing species, L. sitkana, was nearly fixed for a single haplotype that made it impossible to precisely estimate Φ_st. One of the two planktonic-developing species, L. scutulata, did not show any significant population structure (Φ_st=0.004). In contrast to our expectations, the other planktonic-developing species, L. plena, showed some weak but statistically significant population structure (Φ_st=0.052). We discuss how differences in population genetic structure between species with the same type of development may reflect differences in their historical demography. Received: 22 December 1999 / Accepted: 24 July 2000     

Continuous plankton records: Changes in the composition and abundance of the phytoplankton of the north-eastern Atlantic ocean and North Sea, 1958–1974

In most areas of the north-eastern Atlantic Ocean, diatoms have declined drastically in abundance in the last decade. Additionally, in areas to the north of 59°N Ceratium species and an index of total phytoplankton have also declined. South of 59°N the phytoplankton index has increased, diatoms have declined and Ceratium species have remained at a constant level of abundance. A possible explanation of the increase in the phytoplankton index at a time when the diatoms were declinig south of 59°N is the development of unidentified phytoplankton organisms such as microflagellates. As many of the variables influencing phytoplankton standing crop are governed in turn by the prevailing weather, the phytoplankton changes may well be a consequence of the general deterioration, since 1940, of North Atlantic weather. Changes in phytoplankton which may be attributed to an amelioration of climate since 1971 are evident as yet only in the southern North Sea.     

Effects of isolation by distance and geographical discontinuity on genetic subdivision of Littoraria cingulata

Littoraria cingulata (Philippi, 1846) is a Western Australian, mangrove littorine snail, represented by two morphologically distinct subspecies, whose distributions are separated by >300 km. The southern subspecies, L. cingulata pristissini, is distinguished from the northern subspecies, L. cingulata cingulata, by having a thinner, keelless shell with more primary grooves, and lower and much more numerous ribs. In contrast with these striking differences, L. cingulata cingulata is morphologically very similar to another species, L. sulculosa, with which it also shares a nearly coincident geographic range. Allozyme comparisons at 22 presumptive loci confirmed a large genetic distance between L. cingulata and L. sulculosa, and the apparent conspecificity of the morphologically divergent subspecies of L. cingulata. Based on geological evidence, the geographical separation of the morphologically divergent forms of L. cingulata has developed within the past 5000 to 10 000 yr. The extensive continuous distribution of the northern subspecies, L. cingulatacingulata, and the large geographic disjunction between the northern and Shark Bay subspecies, L. cingulata pristissini, allowed a test of the genetic importance of this relatively recent disjunction. Within the continuous distribution of the two subspecies, a pattern of isolation by distance was visible up to distances of 300 km. Beyond 300 km, genetic subdivision, measured by pairwise G _ST (the proportion of genetic diversity due to differences between populations), averaged 0.028, whereas subdivision between Shark Bay and northern populations averaged 0.055 over the same range of distances. Although the relative paucity of barriers to gene flow tends to limit genetic subdivision in marine species with planktotrophic larvae, the results for L. cingulata suggest that subdivision can occur within a continuous distribution, but that special events leading to major disjunctions can substantially increase divergence, even over a relatively short period of time. Received: 16 February 1998 / Accepted: 23 April 1998     

Rearing Pandalus borealis larvae in the laboratory

Larvae of the northern shrimp Pandalus borealis (Krøyer) are pelagic. In the Estuary and Gulf of St. Lawrence, Canada, the early stages are found in the upper 25-m of the water column in spring and early summer and are expected to experience a range of water temperatures from as low as 0°C to as high at 6°C. Little is known of the impact of water temperature on metabolic requirements of northern shrimp larvae. In this study, routine respiration (VO₂), maximum respiration (electron transport system activity, ETSA) and metabolic scope for growth (MS, ETSA–VO₂) of northern shrimp larvae were measured as a function of temperature (3, 5 and 8°C), developmental stage (I–V at 3°C, I–VII at 5°C and 8°C) and growth rate in dry mass. After logarithmic transformation, all three metabolic variables were linearly related to dry mass. The increase in VO₂ with body mass was faster at 5°C than at 3 or 8°C, whereas with ETSA this increase was slower. As a result, MS increased more slowly with dry mass at 5°C than at 3 and 8°C. However, MS did not limit growth in this study, since it explained only 39% of the variability in growth. All three metabolic variables as well as growth varied together as a function of temperature and ontogeny. Q₁₀ of all three metabolic variables ranged from 1.6 and 2.2 for stages I–V larvae, except for VO₂ at stage I (3.9) and stage III (2.9).     

Distribution of dimorphic males of three species of Nematoscelis (Euphausiacea)

Samples collected from the eastern Atlantic Ocean between 40° and 4° N with rectangular and ring nets contained male specimens of Nematoscelis atlantica, N. microps, and N. tenella (Euphausiacea) with saddle-shaped thickenings of chitin on the dorsal surface of some abdominal segments, and with enlarged photophores on the abdominal segments immediately posterior to those with thickenings. Two forms of each species are distinguished by the number and the position of chitinous thickenings and enlarged photophores. Except in a zone off W. Africa (approximately between 22° and 16° N) each form of N. atlantica and N. microps occurs as a distinct population. These populations appear to be characteristic of either the North or South Atlantic Central water, and the overlap zone corresponds approximately with the physical boundary between these water masses and with a faunal transition zone found in this region by other workers. For N. tenella it appears that only one form occurs as a separate population south of 18° N in South Atlantic Central water. Between 28° and 18° N the two forms co-exist.     

Optimal growth and maximal survival temperatures of Atlantic Laminaria species (Phaeophyta) in culture

The effects of temperature on growth rate of rapidly-growing cultured macrosporophytes of 9 isolates of Atlantic Laminaria comprising 4 species have been investigated. No significant population variation was observed within species despite wide variations in temperature between the original collecting sites. L. saccharina showed a broad temperature optimum in the 10°–15°C range, whereas L. longicruris had a sharp optimum at 10°C. L. digitata and L. hyperborea grew more slowly, with only slightly sub-optimal growth over a wide temperature range, but with peaks at 10°C (L. digitata) and 15°C (L. hyperborea). The maximum survival temperatures of individual male and female vegetatively-growing gametophytes were ascertained for these species plus the Arctic L. solidungula, and were as follows: L. saccharina and L. longicruris, 23°C; L. digitata (male), 23°C; L. digitata (female), 22°C; L. hyperborea, 21°C; L. solidungula, 18°C. The lack of within-species differences demonstrates that the success of the genus in areas with different temperature regimes is brought about by phenotypic plasticity of individuals rather than the selection of temperature races or ecotypes.     

Latitudinal variation in aspects of the biology of Cyclothone braueri and C. microdon (Pisces: Gonostomatidae) in the eastern North Atlantic Ocean

Samples from a series of RMT-8 tows along 20°W in spring and autumn of various years were used to study the variability in aspects of the breeding biology of two species of Cyclothone: C. braueri and C. microdon in the eastern North Atlantic Ocean. Variables compared within each species included length-frequency distribution, maximum size, size at maturity, sex ratio, egg size and fecundity. Populations of both species at lower latitudes were generally smaller, reached maturity at a smaller size, and probably spawned earlier in the year and over a longer period than populations at higher latitudes. In both species, fecundity was highest at 40°N in the spring. The results suggested that C. microdon may not spawn at the northern extent of its range.     

Effects of temperature on activity,food consumption rates,and gut passage times of seaweed-eating<Emphasis Type="Italic"> Tegula</Emphasis> species (Trochidae) from California

Previous feeding studies on herbivorous marine snails rarely have focused on temperature effects on food intake. If temperature affects food intake, ectothermic snails may experience difficulty obtaining sufficient nutritional resources, limiting their ability to sustain populations at suboptimal temperatures. We hypothesized that the feeding responses of Tegula species would correspond with temperatures characteristic of their geographic distributions. We determined activity, consumption rates, and gut passage times at 11°C, 15°C, 19°C, and 23°C for three Tegula species with distinct thermal distributions: T. brunnea (cold water), T. aureotincta (warm water), and individuals from warm- and cold-water populations of T. funebralis, a broadly distributed species. Activity and consumption rates of T. aureotincta increased with increasing temperature, but were highest for T. brunnea at 19°C, a temperature rarely achieved in habitats occupied by this species, and lowest at 11°C. Warm-water T. funebralis showed significantly lower activity and consumption rates at 11°C, whereas cold-water T. funebralis consumed food fastest at 15°C and were most active at 23°C. Temperature affected gut passage time only in T. aureotincta. These data suggest that temperature might influence the northern limit of T. aureotincta by affecting activity and food consumption rates. T. brunneas activity and ability to consume food were not hindered by warmer temperatures despite the present day restriction of this species to colder waters. Also, widely separated (>300 km) T. funebralis populations may be adapted to regional conditions based on the different temperature responses of northern and southern snails.Communicated by P.W. Sammarco, Chauvin     

Effect of high temperatures on the ultrastructure of Leydig cells in Mytilus galloprovincialis

Individuals of the mussel Mytilus galloprovincialis (L), collected from the Black Sea, were subjected to relatively high temperatures (25°, 28° and 30°C) for 60 h Ultrastructural studies on Leydig cells of the mantle indicated progressive modifications in their structure. Mainly, we established that a continuous decrease of glycogen quantity in the cells took place as the temperature raised, while the lipid content increased, up to 28°C. Between 28° and 30°C the lipid content decreased as a result of a pronounced degradation process. An increase of membrane permeability and the disturbance in the synthesis of mucoproteic secretory granules were observed. It is concluded that for Mytilus galloprovincialis the thermal adaptation zone ranges between 25° and 28°C. 28°C is the upper tolerable limit temperature, while 30°C acts as a lethal temperature.     

Effects of temperature on the mortality and growth of Hawaiian reef corals

Three common species of Hawaiian reef corals, Pocillopora damicornis (L.), Montipora verrucosa (Lamarck) and Fungia scutaria Lamarck, were grown in a temperature-regulated, continuous-flow sea water system. The skeletal growth optimum occurred near 26°C, coinciding with the natural summer ambient temperature in Hawaii, and was lowest at 21° to 22°C, representing Hawaiian winter ambient. Levels of approximately 32°C produced mortality within days. Prolonged exposure to temperatures of approximately 30°C eventually caused loss of photosynthetic pigment, increased mortality, and reduced calcification. Corals lived only 1 to 2 weeks at 18°C. The corals showed greater initial resistance at the lower lethal limit, but ultimately low temperature was more deleterious than high temperature. Results suggest that a decrease in the natural water temperature of Hawaiian reefs would be more harmful to corals than a temperature increase of the same magnitude.Contribution No. 504 of the Hawaii Institute of Marine Biology.     

On the near-surface plankton of the eastern South Pacific Ocean

A study has been made of the plankton of the near-surface water layer (0 to 30 cm) of the eastern South Pacific Ocean in the region lying to the east of the meridian 90°W, between latitudes 5°N and 35°S. This region is influenced by the Peru Current: the current brings water from high latitudes, which results in a decrease in the number of species of the local fauna of copepods of the family Pontellidae, typical of tropical near-surface plankton. Some of the widely tropical and one bicentral species are absent or rare. Least affected by the Peru Current are the waters of equatorial structure in the northern part of the region. Here, 7 species of pontellids were recorded: the widely tropical Labidocera detruncata, Pontella tenuiremis, Pontellopsis regalis, the distant-neritic Pontella danae and Labidocera acuta, the bicentral Labidocera acutifrons, and the neritic Pontellopsis lubbockii. The dominant species among these are L. detruncata and L. acuta. To the west of the convergence, in the southern part of the region, live the southern central species Pontella valida and P. whiteleggei, with Pontellopsis regalis occurring occasionally. In these regions the copepod fauna is frequently dominated by pontellids. To the south of the boundary of the waters of equatorial structure, between the coast of South America and the line of convergence, lies a region most subjected to the effect of waters from high latitudes and of upwellings. It is inhabited by 2 pontellids only: Pontellopsis regalis and Labidocera acutifrons, but they too disappear close to the coast. In this particular region the copepods Calanus australis and Centropages brachiatus are common; they are found in a thicker water layer (0 to 200 m), and are often more abundant than the pontellids.     

Diel vertical migration and timing of metamorphosis of larvae of the Dungeness crab Cancer magister

Joint USA/USSR ichthyoplankton surveys off the coasts of Washington, Oregon and northern California during the years 1981 to 1985 sampled more than 120 stations each year, from 5 to 360 km offshore and between Latitude 40° and 48° N, providing information on ontogeny and diel migration of larvae of the Dungeness crab Cancer magister on a scale not studied previously. We developed a maximum likelihood method for estimating abundance and fraction in the neuston at each station from a neuston tow and an oblique bongo tow. Latestage megalopae migrate vertically on a diel basis, with the fraction in the neuston being (on average) 62% at night (19.00 to 08.00 hrs Pacific Standard Time, PST) and 8% during the day (08.00 to 19.00 hrs PST). The hourly pattern of this migration includes a peak in the early evening, possibly another in the early morning, and an intermediate level in the late afternoon. We detected no dependence of vertical migration on cloud cover or sea state. Early-stage megalopae were present in much lower fractions in the neuston, but weakly displayed the same diel pattern of migration. Zoeae appeared to be below the neuston at all times, except for 2 or 3 h in the evening. From an abrupt change in larval stage in samples from a north-south cruise, we concluded that the majority of the larvae metamorphose from zoeae to megalopae over a fairly short time span (2 to 4 wk) at a given latitude. In later cruises, 95% of the larvae were megalopae, indicating that metamorphosis over the study area either occurs at the same time or proceeds from south to north over a time span of less than a month in early spring.     

Feeding energetics and metabolism in demersal fish species from Antarctic,temperate and tropical environments

The energetics of feeding has been investigated in demersal fish with similar sedentary lifestyles from the Antarctic (Notothenia neglecta Nybelin), North Sea (Myoxocephalus scorpius L.) and Indian Ocean (Cirrhitichys bleekeri Bleeker). In general, the metabolic rates of fasting individuals were positively correlated with adaptation temperature: values for a standard 100 g fish (mg O₂/h) were 3.3 for N. neglecta at around 0 °C, 2.7 for winter-acclimatized M. scorpius at 5 °C, 4.3 for summer-acclimatized M. scorpius at 15 °C, and 7.0 for C. bleekeri at 25 °C. In all species, following a single satiating meal, oxygen consumption increased to a peak of 2 to 3.5 times the fasting values. Maximum rates of oxygen consumption after feeding were several-fold higher in the warm-than in the cold-water species. After controlling for the effects of body mass and energy intake by analysis of covariance, the duration of the increase in metabolic rate, referred to as specific dynamic action (SDA), was found to be 3 to 4 times shorter in the warm- than in the cold-water fish, ranging from 57 h in C. bleekeri to 208 h in N. neglecta. In contrast, the SDA was not significantly different in the various species, corresponding to 15 to 23% of the energy ingested. Seasonal influences on metabolism and feeding were also studied in N. neglecta acclimated to simulated winter (-1.0 to-0.5 °C; 3 h light:21 h dark) or summer (0 to 0.9 °C; 21 h light:3 h dark) conditions. The metabolic rates of fasting and fed individuals, and the characteristics of the SDA were found to be independent of acclimation conditions. This suggests that N. neglecta is capable of processing food at similar rates throughout the year. Energy stores and enzyme activities were measured in the swimming muscles and liver of fish fed ad libitum. Summer-acclimated fish had higher concentrations of liver triglyceride stores and elevated activities of some enzymes of intermediary metabolism relative to winter-acclimated fish. The observed changes in intermdiary metabolism are probably related to annual cycles of growth and reproduction. It is suggested that the low aerobic scope for physiological performance in Antarctic fish may necessitate the seasonal switching of energy allocation between growth and reproduction.     

Extensive gene flow among mytilid (Bathymodiolus thermophilus) populations from hydrothermal vents of the eastern Pacific

Prior studies of the hydrothermal vent mussel Bathymodiolus thermophilus (Bivalvia: Mytilidae), provided conflicting predictions about the dispersal ability and population structure of this highly specialized species. Analyses of morphological features associated with its larval shells revealed a feeding larval stage that might facilitate dispersal between ephemeral vent habitats. In contrast, an allozyme study revealed substantial genetic differentiation between samples taken from populations 2370 km apart on Galápagos Rift (Latitude 0°N) and the East Pacific Rise (13°N). To resolve the discrepancy between these studies, we examined allozyme and mitochondrial (mt) DNA variation in new samples from the same localities plus more recently discovered sites (9° and 11°N) along the East Pacific Rise. Although analysis of 26 enzyme-determining loci revealed relatively low levels of genetic variation within the five populations, no evidence existed for significant barriers to dispersal among populations. We estimated an average effective rate of gege flow (Nm) of 8 migrants per population per generation. Two common mtDNA variants predominated at relatively even frequencies in each population, and similarly provided no evidence for barriers to gene flow or isolation-by-distance across this species' known range. Larvae of this species appear to be capable of dispersing hundreds of kilometers along a continuous ridge system and across gaps separating non-contiguous spreading centers.     

Temperature acclimation of respiration and photosynthesis in the brown algaLaminaria saccharina

Sporophytes of the brown algaLaminaria saccharina (L.) Lamour grown at 15°C contained significantly more chlorophylla (chla) than did similar plants grown at 5°C. The increase in chla in 15°C plants was due to increased numbers of photosystem II reaction centes, and possibly to increased photosynthetic unit size, compared with 5°C plants. These changes were associated with increased values (photosynthetic efficiencies) in 15°C-grownL. saccharina relative to 5°C-grown plants. The changes in together with reduced respiration rates allowed 15°C-grownL. saccharina to achieve net photosynthesis and light-saturated photosynthesis at a lower photon fluence rate (PFR) than 5°C plants when both groups were assayed at the same temperature (15°C). The photon fluence rates necessary to reach the compensation point and achieve light-saturated photosynthesis (I _c andI _k , respectively) increased with increasing incubation temperature inL. saccharina grown at both 5 and 15°C. However, acclimation responses to growth temperature compensated for the short-term effect of temperature onI _c andI _k . Consequently, plants grown at 5 and 15°C were able to achieve similar rates of light-limited photosynthesis, and similarI _c andI _k values at their respective growth temperatures. These responses are undoubtedly important for perennial seaweeds such asL. saccharina, which frequently grow in light-limited habitats and experience pronounced seasonal changes in water temperature.Please address all correspondence and requests for reprints to I.R. Davison     

55Fe in Pacific Ocean plankton

The specific activity of ⁵⁵Fe in plankton increased from below detectable limits at 20°N to a maximum of 130 nCi/g Fe at 20°S and then decreased to 5 nCi/g Fe at 50°S. Along an east-west track near 15°S a peak of 90 nCi/g Fe occurred at 135°W. It is suggested that this maximum is the result of fallout from French nuclear tests at Mururoa Atoll (22°S; 139°W). The increase in ⁵⁵Fe at high latitudes observed in previous studies in the North Atlantic Ocean and North Pacific Ocean did not occur in the South Pacific Ocean which supports the suggestion that belts of high tropospheric fallout accounted for the increase in the Northern oceans.     

Influence of light and temperature on the growth rate of estuarine benthic diatoms in culture

Four species of estuarine benthic diatoms: Amphiprora c. f. paludosa W. Smith, Nitzschia c. f. dissipata (Kützing) Grunow, Navicula arenaria Donkin, and Nitzschia sigma (Kützing) W. Smith were grown in unialgal cultures. The growth rates of the diatoms were determined as the rate of increase of the chlorophyll a content of the cultures. The diatoms were cultured at different combinations of temperture, daylength, and quantum irradiance. The highest growth rates of Navicula arenaria occurred at 16° to 20°C; the other 3 species had their optimum at 25°C or higher. The small-celled species had higher growth rates at their optimum temperature, but at lower temperatures the growth rates of all 4 species became very similar. The minimum daily quantum irradiance that could effect light-saturated growth at 12° and 20°C ranged from 2.5 to 5.0 E.m^-2.day^-1. At 12°C, two species had their highest growth rates under an 8 h daily photoperiod. At 20°C, the three species tested all had highest growth rates under 16 h daily photoperiod. The growth response of the benthic diatoms is comparable to that of several cultures of planktonic diatoms, as described in the literature. The influence of temperature and quantum irradiance on the diatoms in the present investigation was comparable to the influence of temperature and light intensity on the ¹⁴C-fixation of marine benthic diatoms (Colijn and van Buurt, 1975).     

Larval tolerance, gene flow, and the northern geographic range limit of fiddler crabs

Despite growing interest in species' range shifts, little is known about the ecological and evolutionary factors that control geographic range boundaries. We investigated the processes that maintain the northern range limit of the mud fiddler crab (Uca pugnax) at North Scituate, Massachusetts, USA (42 degrees 14' N), located approximately 60 km north of Cape Cod. Larvae from five populations in Massachusetts were reared under controlled temperatures to test whether cooler water near the edge of this species' range inhibits planktonic development. Few larvae completed development at temperatures < 18 degrees C, a threshold that larvae would regularly encounter north of Cape Cod. Extensive salt marshes are present north of the current range boundary, and a transplant experiment using field enclosures confirmed that benthic fiddler crabs can survive severe winter conditions in this northern habitat. Taken with oceanographic data, these results suggest that the range boundary of fiddler crabs is likely maintained by the influence of cooler water temperatures on the larval phase. Analyses of mitochondrial DNA sequences from a neutral marker (COI) indicate high gene flow among U. pugnax populations in Massachusetts with little differentiation across Cape Cod. Consistent with predictions regarding the homogenizing influence of gene flow, larvae from source populations north and south of Cape Cod shared a common lower threshold for development. However, larvae produced near the range edge had faster growth rates than those from the south side of Cape Cod (typically reaching the final megalopal stage 1.0-5.5 d faster at 18 degrees C). Additional studies are needed to determine the mechanism underlying this counter-gradient variation in development time. We hypothesize that dispersal into cooler water on the north side of Cape Cod may act as a selection filter that sieves out slow developers from the larval pool by increasing planktonic duration and exposure to associated sources of mortality. Thus while high gene flow may prevent the evolution of greater cold tolerance in northern populations, recurrent selection on existing variation may lead to an unexpected concentration of favorable adaptations at the edge of the range. Such a pattern could permit edge populations to play a dominant and unrecognized role in future range extensions.