Experimentally induced effects of water temperature and immersion time on reproductive output of bivalves in the Wadden Sea

Results of experiments are reported on the effects of water temperature and immersion time in winter on egg size and egg numbers in three intertidally living bivalves in the Dutch Wadden Sea, the Baltic tellin Macoma balthica, the common cockle Cerastoderma edule and the common mussel Mytilus edulis. Macoma (14–17 mm shell length) produced large eggs (diameter of 107 μm) in relatively small numbers (20 000–70 000) in early spring. Later in spring, Cerastoderma (28–33 mm shell length) produced smaller eggs (77 μm, excluding the surrounding jelly layer) in tenfold larger numbers (200 000–700 000). Mytilus (45–55 mm shell length) spawned even smaller eggs (72 μm) in high (but not easily assessed) numbers over a more extended period. In Macoma egg size was not affected by winter temperatures or immersion time. Effects of winter–spring temperatures and immersion time on egg size could be demonstrated in Cerastoderma. Smaller eggs were produced at the higher temperatures. Effects of immersion time were non-consistent: at lower water temperatures larger, but at higher temperatures smaller eggs were produced by animals kept at longer immersion times. In Mytilus, no temperature effects were observed. However, a longer immersion time resulted in larger eggs. In Macoma as well as in Cerastoderma significantly more eggs were produced at the lower temperature. Immersion time effects were most pronounced at the lower temperature, where more eggs were produced at the subtidal level than at the tidal level. At the higher water temperature differences between egg numbers produced at the two tidal levels were small. Just prior to spawning, egg numbers were strongly positively related to body mass at a certain shell length.     

The annual reproductive cycle of the freshwater mussel Dreissena polymorpha Pallas in lakes

Summary The annual development of the gonads of Dreissena polymorpha was studied at three sampling sites in two lakes over 3 and 1 1/2 years, respectively. A resting stage occurred after the last spawning in summer/autumn. Oogenesis (accompanied by multiplying segmentation of the oogonia and early growth processes of its oocytes) restarted in specimens at least 1 year old at low temperatures (below 10° C) during winter and early spring. At one location (Fühlinger See) the onset of the spawning season was correlated with an increase of water temperatures above 12° C. At 2 m depth, two main spawning periods in May and August were normally recognized, the first at temperatures of 12–16° C, the second at 16–21° C. It was clearly demonstrated for the first time in Dreissena polymorpha that the oocytes became mature in successive cohorts within one gonad. A female mussel may spawn several times during the reproductive season. At 9 m depth, the onset of spawning also started at about 12° C; this occurred in late summer, with two spawning periods within 1 month at a temperature range of 12–16° C. At another location (Heider Bergsee) the size of the gonads and the oocytes was reduced during April of both years studied, when food supply was low simultaneously with rapidly rising water temperatures in this shallow lake. There was no spawning period during spring. The major spawning period was delayed until July (temperatures 19–22°C). This shows (1) the synchronizing influence of low winter temperatures on the annual reproductive cycle and (2) a temperature threshold of at least 12° C for the start of the spawning processes. The results are discussed with regard to the geographical limits of further spread of Dreissena polymorpha.     

Temperature impact on the midsummer decline of Daphnia galeata: an analysis of long-term data from the biomanipulated Bautzen Reservoir (Germany)

1. The influence of water temperature on occurrence and duration of a midsummer decline (MSD) of Daphnia galeata was studied in the biomanipulated Bautzen Reservoir in Germany. The proportion of piscivores in the fish community of the reservoir has been enhanced experimentally since 1981. As a consequence, Daphnia galeata has dominated the zooplankton. Over 18 years of study (1981–1998), a long‐lasting MSD (longer than 30 days) occurred in 7 years, whereas a short MSD (shorter than 30 days) was observed in 6 years. During the remaining 5 years, an MSD was not observed.
2. Two hypotheses were examined to explain the observed patterns. First, we postulated that high water temperature during winter and early spring (January–April) leads to an MSD after an early and high spring peak of daphnids. On the other hand, low temperature during winter and early spring should not cause an MSD owing to a slower increase of the population, resulting in a later peak of daphnids. Second, we hypothesized that the mean water temperature during early summer (May and June) influences the occurrence of an MSD (by controlling young‐of‐the‐year (YOY) fish predation on daphnids).
3. The water temperature during winter and early spring explains 83%, and the early summer water temperature 55%, of interannual variation in the occurrence of an MSD.
4. The interannual variation in duration of an MSD was neither explained by temperature during winter and early spring nor by early summer temperature alone, but in 14 of the 18 years (78%) by a combination of both.
5. We conclude that water temperature during winter and early spring had a strong impact on Daphnia mortality by influencing height and timing of the spring peak which, in turn, influenced the extent of overexploitation of their food resources. By contrast, the water temperature during early summer probably influenced the mortality of daphnids caused by predation of YOY fish. The relative timing of both sources of mortality, which depends on the temperature regime during the first 6 months of the year, is the key process in controlling the occurrence and duration of an MSD. A long‐lasting MSD, therefore, is likely in Bautzen Reservoir only if temperatures are high during winter and early spring, as well as during early summer.
6. As a consequence of climate warming, recent climate records reveal warming during winter, spring and early summer in middle Europe, rather than an increase in mean annual temperatures. If our findings and conclusions are related to this regional and temporal pattern of climate warming, an increasing frequency of years with a long‐lasting MSD and, consequently, a decreasing efficiency of biomanipulation can be predicted.     

The macrobenthos fraction accessible to waders may represent marginal prey

Summary The relationship between relative body condition (deviation from expected mean body weight) and burying depth was investigated in five macro-zoo benthic species living in a marine intertidal habitat. Body weight increased with depth when animals of the same size were compared. The increase amounted to 50% in the clamScrobicularia plana, ca. 40% in the wormNereis diversicolor, 25% in the clamMacoma balthica and 20% in the cockleCerastoderma edule and the clamMya arenaria. Only a part of the prey was within reach of some feeding wader species. Therefore prey value may be overestimated if one does not take into account the fact that shallow and accessible prey often have a relatively poor body condition.     

Deepwater broadcast spawning by Montastraea cavernosa, Montastraea franksi, and Diploria strigosa at the Flower Garden Banks, Gulf of Mexico

Broadcast spawning by corals is a tightly synchronized process characterized by co-ordinated gamete release within 30–60 min time windows once per year. In shallow water corals, annual water temperature cycles set the month, lunar periodicity the day, and sunset time the hour of spawning. This tight temporal regulation is critical for achieving high fertilization rates in a pelagic environment. Given the differences in light and temperature that occur with depth and the importance of these parameters in regulating spawn timing, it has been unclear whether deeper coral can respond to the same environmental cues that regulate spawning behaviour in shallower coral. In this report, a remotely operated vehicle was used to monitor coral spawning activity at the Flower Garden Banks at depths from 33 to 45 m. Three species Montastraea cavernosa, Montastraea franksi, and Diploria strigosa were documented spawning within this depth range. All recorded spawning events were within the same temporal windows as shallower conspecifics. These data indicate that deep corals at this location either sense the same environmental parameters, despite local attenuation, or communicate with shallower colonies that can sense such spawning cues.     

Thermal stratification influences maturation and timing of spawning in a local Clupea harengus population

Maturation and timing of spawning in relation to temperature were studied in a local Atlantic herring Clupea harengus population inhabiting a small semi‐enclosed ecosystem (7 km²) separated from the larger outer fjord system by narrow sills on the west coast of Norway. Ambient temperatures varied annually up to 4° C during both the pre‐spawning and spawning periods from February to April, but without affecting the spawning time. Instead, the timing of spawning was found to be related to thermal stratification in response to spring warming, which occurred about the same time every year regardless of initial temperatures.     

The influence of environment,sex, and innate timing mechanisms on body temperature patterns of free-ranging black-tailed prairie dogs (Cynomys ludovicianus)

Mechanisms that influence body temperature patterns in black-tailed prairie dogs are not well understood. Previous research on both free-ranging and laboratory populations of black-tailed prairie dogs (Cynomys ludovicianus) has suggested that reductions in ambient temperature and food and water deprivation are the primary factors that stimulate torpor in this species. In other species, however, torpor has been shown to be influenced by a multitude of factors, including innate circadian and circannual timing mechanisms, energy status, and reproductive behaviors. Our objective was to clarify the influence of weather, sex, and intrinsic timing mechanisms on the body temperature patterns of free-ranging black-tailed prairie dogs. We monitored body temperatures of eight adult (>1 yr) prairie dogs from November 1999 to June 2000. Prairie dogs showed distinct daily and seasonal body temperature patterns, which reflected changes in ambient temperatures that occurred during these periods. These patterns of daily and seasonal heterothermy suggest that body temperature patterns of black-tailed prairie dogs may be driven by an innate timing mechanism. All prairie dogs entered torpor intermittently throughout winter and spring. Torpor bouts appeared to be influenced by precipitation and reductions in ambient temperature. Our results also suggest that reproductive behaviors and circadian timing may influence torpor in this species.     

Effect of season on peripheral resistance to localised cold stress

This study was carried out to determine the effect that seasonal changes have on the effect of localised cold stress on peripheral temperatures using the foot immersion method with a cold water bath. The subjects were six males and four females. The data were obtained in April, July, October and January. Skin temperature of the right index finger, the forehead, the arm, the cheek, the second toe and the instep were measured before, during and after the immersion of the feet in water at 15°C for 10 mins, as well as oxygen consumption before immersion of the feet.The average finger temperature was highest during foot immersion in the summer, next highest in the winter, then spring, and the lowest during foot immersion in the autumn. The finger temperatures during the pre-immersion period in the autumn tended to be lower than in other seasons. The finger temperatures during the pre-immersion period affected the temperature change of the finger during the immersion period. The rate of increase of the toe temperature and the foot temperature during post-immersion in the summer and the spring were greater than those in the autumn and winter. Oxygen consumption during the pre-immersion period in the autumn was significantly lower than in the other seasons (p<0.001 or 0.010). Cooling the feet caused no significant changes in the temperatures the cheek, forehead or forearm. The cheek temperature in the summer and autumn was cooler than corresponding temperatures taken in the winter and spring.     

黄海中南部细纹狮子鱼的生物学特征及资源分布的季节变化

根据2009年7—8月、10月和2010年1月、5月黄海中南部渔业生物底拖网调查数据,对该海域细纹狮子鱼的生物学特征及其分布的季节变化作了分析。结果表明,细纹狮子鱼平均体长和平均体重从春季(4.7 cm、3.3 g)到冬季(34.2 cm、764.9 g)呈显著增加,并且雄性个体平均体长显著大于雌性个体(P<0.05,春季除外)。性比(♀∶♂)随体长组和季节变化,体长越大趋向于雄性,反之趋向于雌性;夏季雄鱼居多(0.70∶1,P<0.05),秋季则为雌鱼居多(1.35∶1,P<0.05),而冬季(产卵群体)和春季性比接近于1∶1。细纹狮子鱼各季节摄食等级均在2.5以上,冬季雄性个体摄食等级显著大于雌性个体(P<0.05),但雄性个体肥满度为全年最低(1.52)。细纹狮子鱼相对资源量和贡献率从春季(0.17 kg/h,1.54%)到秋季(15.36 kg/h,33.05%)呈上升状态,而冬季(2.37 kg/h,5.60%)有所下降。相比2000年,夏秋季相对渔获量和贡献率提高明显。全年集中分布于7.8—13.6℃,3.20%—3.38%的水域,平均体重和水深有显著的相关性(秋季除外)。另外,根据性成熟个体分布区和稚幼鱼分布的相关历史资料分析发现,除海州湾外,黄海中部深水区可能是细纹狮子鱼的产卵场。     

The influence of freshwater inflows on spawning success and early growth of an estuarine resident fish species, Acanthopagrus butcheri

The influence of freshwater inflows and salinity on spawning success of black bream Acanthopagrus butcheri (Sparidae) was investigated over 2 years in a small estuary on the east coast of Tasmania, Australia. The individual spawning seasons experienced quite different freshwater inflows; 2004-2005 was characterized by low flows throughout the season whereas during 2005-2006 there were three relatively large discharge events in the first part of the season. Macroscopic gonad staging of adults was used to define the spawning season and daily increment analysis of otoliths from recently settled recruits was used to backcalculate spawning dates. Gonad staging indicated that adults were in spawning condition over a 3 to 4 month period during spring and summer. The timing and duration of successful spawning, however, differed markedly between years and was linked to the timing of freshwater inflows and salinity conditions, with successful spawning occurring during periods of low freshwater discharge and when salinities in the upper estuary were above c. 15. Growth rates of the recently settled recruits did not differ between years, nor did the timing of spawning within the season influence growth rates. While the latter finding was unexpected, especially given within season temperature variability, these results imply that by the onset of winter earlier spawned fish would be larger than later spawned individuals, potentially conferring advantages for survival and competition for food. Climate change predictions for eastern Tasmania indicate a decrease in river flows in spring and an increase during summer, potentially increasing environmental variability between and within years, with implications for spawning success and subsequent recruitment.     

Influence of acclimation temperature and season on acute temperature preference of adult mountain whitefish,Prosopium williamsoni

Synopsis Temperatures preferred by four groups of adult mountain whitefish, Prosopium williamsoni, collected in October before spawning, in December after spawning, in late winter, and in spring were determined in the laboratory in a horizontal gradient. Acute temperature preference based on fish tested soon after capture and final preferendum estimates were 12.8 and 17.7°C, respectively (pre-spawning), 9.6 and 11.9°C (post-spawning), 10.8 and 9.9°C (winter), and 16.4 and 16.3°C (spring). Seasonal influence on temperature preference was evident on the basis of differences in final preferenda, covariance analysis of responses of laboratory-acclimated fish, and temperature preference of fish held at ambient river temperatures. Fish of the post-spawning and winter groups preferred lower temperatures than did those of pre-spawning and spring groups. Temperatures preferred by pre-spawning fish were too high for embryo survival. Caution is necessary in predicting thermal preference on the basis of a sample collected at one time of year or stage of sexual development.     

Seasonal variation in condition, growth and food habits of walleye in a Great Plains reservoir and simulated effects of an altered thermal regime

Catch rates in gillnets and relative weight ( W _r) of walleye Stizostedion vitreum , in Glen Elder Reservoir, Kansas, were lowest during the summer (June–August) and highest during the autumn (September–November). Approximately 80% of their annual growth in length and mass was attained during late summer and autumn. Growth was minimal during winter (January–February) and spring (March–May). The number of walleye with empty stomachs was highest during the summer. Invertebrates (Cladocera, Chironomidae) were common in walleye stomachs during the summer and spring, but contributed little to the ingested biomass. Gizzard shad Dorosoma cepedianum dominated walleye diets (per cent by mass) throughout the year. A bioenergetics model predicted that the proportion of maximum consumption ( P _c) was highest during the autumn and was probably due to spatial overlap of walleye and gizzard shad once water temperatures were <22° C. The bioenergetics model predicted that walleye would lose up to 65% of their body mass during the summer if water temperature increased by 10% (as predicted by some global warming models). Growth during the autumn, winter and spring was enhanced up to 150% by increased temperatures. The results of this study indicate that lower condition, reduced consumption and slow growth are a generalized response of walleye to extreme temperatures. Elevated temperatures may have a net positive effect on walleye growth if they can survive the high thermal stress during summer.     

The adult body size variation of Dryas iulia (Lepidoptera,Nymphalidae, Heliconiinae) in different populations is more influenced by temperature variation than by host‐plant availability during the seasons

The body size of insects is affected by environmental conditions during development and can present considerable intraspecific variations, which can be seen as an ultimate consequence/adaptation to environmental conditions. This paper evaluated whether the body size of the butterfly Dryas iulia from subtropical populations was influenced by changing climate conditions and food source availability during the seasons. The likely reasons behind body size variation were also investigated. First, field data on body size variation, host‐plant availability and climate fluctuation throughout the seasons were recorded. Then, the effects of host‐plant species and temperature on body size were analyzed by controlled experiments. Field data revealed that body size and host‐plant availability varied significantly through the seasons. Populations had the smallest body size during the spring and the biggest size during summer, whereas host‐plant availability was lower during winter and higher during spring. The controlled experiments revealed that both temperature and host‐plant had significant effect on the plasticity of body size. Larvae subjected to winter temperature treatment led to smaller butterflies when compared to immatures reared under summer temperature treatment, and larvae fed with Passiflora misera produced bigger adults when compared to larvae reared on Passiflora suberosa. The combination of data gathered in the field and in the laboratory suggests that seasonal body size variation in D. iulia is related mainly to differences in the temperatures to which larvae are subjected during development, while host‐plant shifts caused by differential availability of food through the seasons had slightly effects on the variation observed.     

The North Atlantic Oscillation and plankton dynamics in two European lakes –- two variations on a general theme

Long‐term data on water temperature, phytoplankton biovolume, Bosmina and Daphnia abundance and the timing of the clear‐water phase were compared and analysed with respect to the influence of the North Atlantic Oscillation (NAO) in two strongly contrasting lakes in central Europe. In small, shallow, hypertrophic Müggelsee, spring water temperatures and Daphnia abundance both increased more rapidly than in large, deep, meso/oligotrophic Lake Constance. Because of this, the clear‐water phase commenced approximately three weeks earlier in Müggelsee than in Lake Constance. In Müggelsee, the phytoplankton biovolume during late winter/early spring was related to the NAO index. In Lake Constance, where phytoplankton growth was inhibited by intense downward mixing during all years studied, this was not the case. However, in both lakes, interannual variability in water temperature, in Daphnia spring population dynamics and in the timing of the clear‐water phase, were all related to the interannual variability of the NAO index. The Daphnia spring population dynamics and the timing of the clear‐water phase appear to be synchronized by the NAO despite large differences between the lakes in morphometry, trophic status and flushing and mixis regimes, and despite the great distance between the lakes (～700 km). This suggests that a great variety of lakes in central Europe may possibly have exhibited similar interannual variability during the last 20 years.     

Upper-lethal-temperature relations of the nymphs of the stonefly,paragnetina media

Summary Continuous water flow and static test apparatus were used in evaluating the influence of acclimation temperature, size, sex, season, life stage, current velocity, and testing method upon the upper-lethal temperature of the nymphal stage of the stonefly, Paragnetina media Walker.Lethal temperature was raised by acclimation to higher temperatures throughout the thermal range examined. The rate of gain of heat resistance was approximately 5°C per day. Acclimation to colder temperatures was much slower.Body size had no significant effect on upper-lethal temperature in summer or autumn, whereas in winter and spring larger nymphs were significantly less tolerant than smaller nymphs. This difference was attributed to sex, not body size per se.Tolerance was significantly less in static waters than in flowing water, faster current significantly increasing heat resistance at all lethal temperatures examined.Upper-lethal temperatures showed seasonal differences independent of acclimation, with nymphs least tolerant in spring, when temperature resistance was influenced by sex, molting condition, and proximity of emergence. The most temperature-sensitive stage was the period immediately following egg hatching.The direct effect of upper-lethal temperature is apparently not a limiting factor for P. media in the environment in which it is found. Temperature increases do, however, effect final stages of nymphal development, emergence patterns, and survival after hatching.Contribution No. 213, W. K. Kellogg Biological Station of Michigan State University. This investigation was supported in part by Research Grant WP 01178 from the Federal Water Pollution Control Administration.     

Assessment of the Feasibility of Preventing Reproduction of Lake Charr, Salvelinus Namaycush, in Shallow Areas of Reservoirs Affected by Drawdowns

Lake charr, Salvelinus namaycush, spawn almost exclusively in lakes, in the fall. Spawning sites in inland lakes are generally located close to the shore in areas with coarse-textured substrate, and in depths of less than 2m. Because of the type of water management system applied in reservoirs where water levels are lowered during the winter, eggs deposited during the fall around the shallow shoreline of reservoirs can be exposed. The depth of spawning sites relative to drawdowns may be a limiting factor for lake charr survival in reservoirs. In 1995 and 1996, we evaluated the ability to induce lake charr reproduction in deeper water, below the range of fluctuating water levels by the sequential construction of artificial reefs and limiting access to natural spawning habitat. The creation of artificial deeper water spawning areas, adjacent to natural sites, produced a partial transfer of reproduction. However when natural spawning sites were covered with tarpaulins almost all the eggs were laid in the artificial deeper water spawning areas. Complete or partial transfer of reproduction activities to sites below the depth not affected by water level fluctuations is therefore a potential way of alleviating the effects of hydroelectric reservoir management.     

The Role of Temperature and Embryo Development Time in the Diel Timing of Spawning in a Coral-reef Damselfish with High-frequency Spawning Synchrony

We studied the diel timing of spawning in the demersally spawning Hawaiian damselfish, Dascyllus albisella, from mid-June to late-September 1997 at two small patch reefs in Hawaii. Our objectives were to elucidate daily timing of spawning in relation to water temperature, diel timing of hatching, and short-period spawning synchrony. Spawning occurred every 5–7 days at both reefs, with all spawning on a reef concluded either within a single day (1-day spawning) or within two successive days (2-day spawning). Spawning began in early morning and continued for most of the day. There was a significant, positive linear relationship between mean daily average water temperature (= daily average temperature averaged over the period starting from the day following the last spawning day of the preceding nest cycle till the day before the first spawning day of the current cycle) and peak spawning hour of day, for 1-day spawning, and the first and second days of 2-day spawning at both reefs. The relationship between mean daily average water temperature and peak spawning hour of day was comparable among all spawning-day classes and reefs. Hatching occurred on the fourth day of development throughout the study despite the 26.5–29.1°C change in water temperature during the study period, and hatching was restricted to within two hours after sunset. We propose that D. albisella's peak spawning time is positively correlated with increased water temperature because it maintains the benefits of synchronous spawning within two constraints: the narrow daily period of hatching, and the inverse relationship between water temperature and embryo developmental time.     

Timing matters: species‐specific interactions between spawning time,substrate quality,and recruitment success in three salmonid species

Substratum quality and oxygen supply to the interstitial zone are crucial for the reproductive success of salmonid fishes. At present, degradation of spawning grounds due to fine sediment deposition and colmation are recognized as main factors for reproductive failure. In addition, changes in water temperatures due to climate change, damming, and cooling water inlets are predicted to reduce hatching success. We tested the hypothesis that the biological effects of habitat degradation depend strongly on the species‐specific spawning seasons and life‐history strategies (e.g., fall‐ vs. spring‐spawners, migratory vs. resident species) and assessed temperature as an important species‐specific factor for hatching success within river substratum. We studied the species‐specific differences in their responses to such disturbances using egg‐to‐fry survival of Danube Salmon (Hucho hucho), resident brown trout (Salmo trutta fario), and migratory brown trout (Salmo trutta lacustris) as biological endpoint. The egg incubation and hatching success of the salmonids and their dependence on temperature and stream substratum quality were compared. Hatching rates of Danube salmon were lower than of brown trout, probably due to higher oxygen demands and increased interstitial respiration in spring. Increases in maximum water temperature reduced hatching rates of resident and migratory brown trout (both fall‐spawners) but were positively correlated with hatching rates of Danube salmon (a spring‐spawner). Significantly longer incubation periods of resident and migratory brown trout coincided with relatively low stream substratum quality at the end of the egg incubation. Danube salmon seem to avoid low oxygen concentrations in the hyporheic zone by faster egg development favored by higher water temperatures. Consequently, the prediction of effects of temperature changes and altered stream substratum properties on gravel‐spawning fishes and biological communities should consider the observed species‐specific variances in life‐history strategies to increase conservation success.     

Age and size-dependent variation in the seasonal timing and probability of reproduction among mature female pumpkinseed,Lepomis gibbosus

Synopsis In this study, we used pumpkinseed,Lepomis gibbosus, populations in two east-central Ontario lakes to test for age and size-dependent effects on the probability and timing of reproduction of mature females. Pumpkinseed body size characteristics differed in the two lakes; Little Round Lake harbours a stunted population and Beloporine Lake does not. Age-class distribution of mature females was determined by a maturity assessment on fish collected just prior to spawning and at the mid-spawning period, combined with an early-season capture-mark-recapture survey. Spawning females were collected throughout the breeding season to assess age and size-related temporal trends, and to compare their age and size distribution with that of mature females at large. The proportion of age 2 females in Little Round Lake that actually spawned was considerably lower than the proportion of mature age 2 females at large. Furthermore, age 2 females that spawned in this lake did not do so until late in the breeding season. In contrast, the proportion of young/small females spawning in Beloporine Lake was comparable to the proportion of young/small mature females at large, and both small and large females spawned throughout the breeding season. Small mature females in Little Round Lake may have had no other option but to spawn late in the season because of their poor body condition. In Beloporine Lake, condition factor early in the breeding season in age 2–4 females was higher than that of Little Round Lake females, suggesting that limited energy reserves in the spring may have prevented young Little Round Lake females from early spawning. Our results show that the likelihood and timing of reproduction are both age and size-dependent in some populations. Small individuals that delay seasonal maturation and spawn late in the summer probably contribute little to the population due to the restricted growth and reduced overwinter survival of their progeny.To whom correspondence should be addressed     

Effects of climate and demography on reproductive phenology of a harvested marine fish population

Shifts in phenology are a well‐documented ecological response to changes in climate, which may or may not be adaptive for a species depending on the climate sensitivity of other ecosystem processes. Furthermore, phenology may be affected by factors in addition to climate, which may accentuate or dampen climate‐driven phenological responses. In this study, we investigate how climate and population demographic structure jointly affect spawning phenology of a fish species of major commercial importance: walleye pollock (Gadus chalcogrammus). We use 32 years of data from ichthyoplankton surveys to reconstruct timing of pollock reproduction in the Gulf of Alaska and find that the mean date of spawning has varied by over 3 weeks throughout the last >3 decades. Climate clearly drives variation in spawn timing, with warmer temperatures leading to an earlier and more protracted spawning period, consistent with expectations of advanced spring phenology under warming. However, the effects of temperature were nonlinear, such that additional warming above a threshold value had no additional effect on phenology. Population demographics were equally as important as temperature: An older and more age‐diverse spawning stock tended to spawn earlier and over a longer duration than a younger stock. Our models suggest that demographic shifts associated with sustainable harvest rates could shift the mean spawning date 7 days later and shorten the spawning season by 9 days relative to an unfished population, independent of thermal conditions. Projections under climate change suggest that spawn timing will become more stable for walleye pollock in the future, but it is unknown what the consequences of this stabilization will be for the synchrony of first‐feeding larvae with production of zooplankton prey in spring. With ongoing warming in the world’s oceans, knowledge of the mechanisms underlying reproductive phenology can improve our ability to monitor and manage species under changing climate conditions.