A top-down survival mechanism during early marine residency explains coho salmon year-class strength in southeast Alaska

Coho salmon (Oncorhynchus kisutch) are a vital component in the southeast Alaska marine ecosystem and are an important regional fishery resource; consequently, understanding mechanisms affecting their year-class strength is necessary from both scientific and management perspectives. We examined correlations among juvenile coho salmon indices, associated biophysical variables, and adult coho salmon harvest data from southeast Alaska over the years 1997–2006. We found no relationship between summer indices of juvenile coho salmon growth, condition, or abundance with subsequent harvest of adult coho salmon in the region. However, using stepwise regression, we found that variation in adult coho salmon harvest was largely explained by indices of juvenile pink salmon (Oncorhynchus gorbuscha) abundance (67%) and zooplankton abundance (24%). To determine if high juvenile pink salmon abundance indicates favorable “bottom-up” lower trophic level environmental conditions for juvenile coho salmon, we plotted abundance of juvenile pink salmon against growth and condition of juvenile coho salmon. No change in growth or condition of juvenile coho salmon was observed in relation to the abundance index for juvenile pink salmon. Therefore, we hypothesize that coho salmon year-class strength in southeast Alaska is influenced by a “top-down” predator control mechanism that results from more abundant juvenile pink salmon, which serve as a predator buffer during early marine residency.     

Variability in scale growth rates of chum salmon (Oncorhynchus keta) in relation to climate changes in the late 1980s

Fish scales were used to investigate the interannual variability in chum salmon growth rates at specific ages in relation to climatic/environmental changes during the 1980s–1990s. Scales were obtained from adult salmon returning to the east coast of Korea between 1984 and 1998. Assuming proportionality between scale size increments and fish length, distances between scale annuli were regarded as the growth conditions in different habitat areas with respect to the life stages of chum salmon. In estuarine and coastal areas, growth rates of fingerling salmon were higher in the 1990s than in the 1980s. Zooplankton abundance off the east coast of Korea increased after the late 1980s, which may have provided favorable growth conditions for young salmon in the 1990s. Growth of juvenile chum salmon during the first summer (Okhotsk Sea) was relatively stable, and neither SST nor zooplankton biomass fluctuated significantly during the study period. However, in the Bering Sea, salmon growth rates between age-2 and age-4 (i.e. ocean-phase immature salmon) were higher in the 1980s than in the 1990s. Variability in salmon growth in the Bering Sea was correlated to zooplankton biomass. These results suggest that the climate regime shift of 1988/1989 in the subarctic North Pacific affected salmon growth mediated by changes of zooplankton biomass, revealing a bottom-up process.     

Relative abundance, age, growth, and fecundity of grubby Myoxocephalus aenaeus in Niantic River and Niantic Bay, Long Island Sound

Grubby (Myoxocephalus aenaeus, Cottidae) is a common benthic fish of inshore waters and estuaries of eastern Long Island Sound; however, little information exists on their life history or population demographics. This study utilised a long-term data series (1976–2002) to assess grubby life history and population demographics and explores trends in the Niantic River and Niantic Bay populations. In addition, we examined the age, size, and fecundity of adult grubby in 2001–02 to determine the population characteristics in the region. Mean grubby catch per unit effort (CPUE) in Niantic Bay ranged from 0.4 per trawl in 1976 to 2.9 per trawl in 1984 while river CPUE ranged from 0.4 per trawl in 1977 to 7.6 per trawl in 1989. Catch of grubby in bottom trawls varied seasonally with highest CPUE occurring in winter. Highest entrainment of grubby larvae occurred in 2001 while the lowest entrainment observed was in 1991. Four age classes, 0+ through III+, were derived from otolith analysis (N = 51) although length frequency analysis suggested the possibility of older fish in the population. The total number of eggs in ovaries ranged from 286 to 16 451 for grubby (N = 64) between 52 mm and 155 mm TL. Results of this study indicated a decline in abundance of adult grubby over the 26-year period, possibly related to concurrent declines in eelgrass (Zostera marina) abundance and/or increased water temperature.     

Sunspot activity and oceanic conditions in the northern north Pacific Ocean

During periods of sunspot maxima (approximately every 11 years) the mean winter position of the center of the Aleutian Low pressure system shifts from the Gulf of Alaska to the western Aleutian Islands, and mean, cyclonic, wind-stress transport in the Gulf of Alaska is reduced by roughly 20%. Coastal sea level data in the gulf do not reflect an 11-year cycle but spectral energy densities indicate an approximate 6-year periodicity also present in transpacific annual mean sea surface temperatures that, in the last one or two decades, parallels large year classes of Pacific herring in southeastern Alaska, large escapements of sockeye salmon fry in the Bristol Bay area, and maxima in the January catch of Dungeness crab in Alaska.     

Have there been recent changes in climate? Ask the fish

It is generally accepted that a climate shift occurred about 1977 that affected the dynamics of North Pacific marine ecosystems. Agreement on the possibility of further climate shifts in 1989 and the late 1990s is yet to be achieved. However, there have been changes in the dynamics of key commercial fishes that indicate changes in their environment occurred in the early 1990s, and possibly around 1998. One method of measuring climate change is to observe the dynamics of species that could be affected.Several studies have described decadal-scale changes in North Pacific climate–ocean conditions. Generally, these studies focus on a single index. Using principal components analysis, we use a composite index based on three aspects of climate ocean conditions: the Aleutian Low Pressure Index, the Pacific Atmospheric Circulation Index and the Pacific Interdecadal Oscillation Index. We link this composite index (Atmospheric Forcing Index) to decadal-scale changes in British Columbia salmon and other fish populations. Around 1989 there was a change from intense Aleutian Lows (above average south-westerly and westerly circulation patterns and warming of coastal sea surface temperatures) to average Aleutian Lows (less frequent south-westerly and westerly circulation and slightly cooler coastal sea surface temperatures in winter). These climate–ocean changes were associated with changes in the abundance and ocean survival of salmon (Oncorhynchus spp.), distribution and spawning behaviour of hake (Merluccius productus) and sardines (Sardinops sagax) and in recruitment patterns of several groundfish species.     

Unusual bed forms on the North Aleutian Shelf,Bristol Bay,Alaska

Side-scan sonar records collected over an area of the North Aleutian Shelf, approximately 250 km west of the head of Bristol Bay, Alaska, identified widespread evidence of active sea floor erosion processes, including sediment transport. Thousands of sea floor depressions, many linear and some containing rippled floors, were identified in water depths of 30 to 90 m. The depressions cover approximately 40 percent of the area surveyed. The sea floor depressions are interpreted to be erosional features, and in conjunction with a field of sand waves, exemplify the dynamic nature of the ocenographic processes active on this area of the sea floor.     

Spatio-temporal covariability in coho salmon (Oncorhynchus kisutch) survival,from California to southeast Alaska

One of the motivations of the GLOBEC Northeast Pacific program is to understand the apparent inverse relationship between the increase in salmon catches in the Gulf of Alaska and concurrent declines in the California Current System (CCS). We therefore used coded wire tag (CWT) data to examine the spatial and temporal patterns of covariability in the survival of hatchery coho salmon along the coast from California to southeast Alaska between release years 1980 and 2004. There is substantial covariability in coho salmon survival between neighboring regions along the coast, and there is clear evidence for increased covariability within two main groups – a northern and southern group. The dividing line between the groups lies approximately at the north end of Vancouver Island. However, CWT survivals do not support inverse covariability in hatchery coho salmon survival between southeast Alaska and the CCS over this 25 year time span. Instead, the hatchery coho survival in southeast Alaska is relatively uncorrelated with coho survival in the California Current System on inter-annual time scales. The 50% correlation and e-folding scales (distances at which magnitude of correlations decreases to 50% and e⁻¹ (32.8%), respectively) of pairwise correlations between individual hatcheries were 150 and 217 km, which are smaller than that reported for sockeye, pink, and chum salmon. The 50% correlation scale of coho salmon is also substantially smaller than those reported for upwelling indices and sea surface temperature. There are also periods of 5–10 years with high covariability between adjacent regions on the scale of hundreds of km, which may be of biological and physical significance.     

A critical size and period hypothesis to explain natural regulation of salmon abundance and the linkage to climate and climate change

We hypothesise that salmon year class strength is determined in two stages during the first year in the ocean. There is an early natural mortality that is mostly related to predation, which is followed by a physiologically-based mortality. Juvenile salmon that fail to reach a critical size by the end of their first marine summer do not survive the following winter. In this study we describe our initial tests of this critical size and critical period hypothesis using data from ocean surveys of juvenile salmon and from experimental feeding studies on coho. Conservative swept volume abundance estimates for juvenile coho, and possibly chinook, indicate that there is high mortality in fall and winter during their first year in the sea. Studies of otolith weight show that the length and otolith-weight relationship for young coho changes in the early fall of their first ocean year. Studies of growth and associated hormone levels in feeding studies show that slow growing juvenile coho are stunted and deficient in an insulin-like growth factor-I (IGF-I). Juvenile coho sampled in September had low IGF-I values, indicative of poor growth. The results of these studies provide evidence for the general hypothesis that growth-related mortality occurs late in the first marine year and may be important in determining the strength of the year class (brood year). The link between total mortality and climate could be operating via the availability of nutrients regulating the food supply and hence competition for food (i.e. bottom–up regulation).     

Recruitment variation of eastern Bering Sea crabs: Climate-forcing or top-down effects?

During the last three decades, population abundances of eastern Bering Sea (EBS) crab stocks fluctuated greatly, driven by highly variable recruitment. In recent years, abundances of these stocks have been very low compared to historical levels. This study aims to understand recruitment variation of six stocks of red king (Paralithodes camtschaticus), blue king (P. platypus), Tanner (Chionoecetes bairdi), and snow (C. opilio) crabs in the EBS. Most crab recruitment time series are not significantly correlated with each other. Spatial distributions of three broadly distributed crab stocks (EBS snow and Tanner crabs and Bristol Bay red king crab) have changed considerably over time, possibly related in part to the regime shift in climate and physical oceanography in 1976–1977. Three climate-forcing hypotheses on larval survival have been proposed to explain crab recruitment variation of Bristol Bay red king crab and EBS Tanner and snow crabs. Some empirical evidence supports speculation that groundfish predation may play an important role in crab recruitment success in the EBS. However, spatial dynamics in the geographic distributions of groundfish and crabs over time make it difficult to relate crab recruitment strength to groundfish biomass. Comprehensive field and spatially explicit modeling studies are needed to test the hypotheses and better understand the relative importance and compound effects of bottom-up and top-down controls on crab recruitment.     

An overview of the Oyashio ecosystem

The Oyashio shelf region and the seasonally ice-covered areas north of Hokkaido are highly productive, supporting a wide range of species including marine mammals, seabirds and commercially important species in the western subarctic Pacific. The fishes include gadids, such as walleye pollock and Pacific cod, and subarctic migratory pelagic fishes such as chum salmon and pink salmon. It is also an important summer feeding ground for subtropical migrants such as the Japanese sardine, Japanese anchovy, Pacific saury, mackerels, Japanese common squid, whales and seabirds. In recent decades, some components of the Oyashio ecosystem (i.e., phytoplankton, mesozooplankton, gadid fish, and subtropical migrants) have shown changes in species abundance or distribution that are correlated with environmental changes such as the 1976/1977 and 1988/1989 regime shifts. The First Oyashio Intrusion moved northward from the mid-1960s until the late 1970s, when it moved southward until the 1980s, after which it returned to the north again after the mid-1990s. The sea-surface temperature in spring decreased after the late 1970s, increased after the late 1980s, and remained high during the 1990s. The extent of ice cover in the Sea of Okhostk also decreased during the latest warming in the 1980–1990s but has increased again since the late 1990s. This and other variabilities affect the Oyashio ecosystem and the surrounding region.     

Covariance among North Sea ecosystem state indicators during the past50 years — Contrasts between coastal and open waters

Regime shift and principal component analysis of a spatially disaggregated database capturing time-series of climatic, nutrient and plankton variables in the North Sea revealed considerable covariance between groups of ecosystem indicators. Plankton and climate time-series span the period 1958–2003, those of nutrients start in 1980. In both regions, the period from 1989 to 2001 identified in principal component 1 had warmer surface waters, higher Atlantic inflow and stronger winds, than the periods before or after. However, it was preceded by a regime shift in both open (PC2) and coastal (PC3) waters during 1977 towards more hours of sunlight and higher water temperature, which lasted until 1997. The relative influence of nutrient availability and climatic forcing differed between open and coastal North Sea regions. Inter-annual variability in phytoplankton dynamics of the open North Sea was primarily regulated by climatic forcing, specifically by sea surface temperature, Atlantic inflow and co-varying wind stress and NAO. Coastal phytoplankton variability, however, was regulated by insolation and sea surface temperature, as well as Si availability, but not by N or P. Regime shifts in principal components of hydrographic and climatic variables (explaining 55 and 61% of the variance in coastal and open water variables) were detected using Rodionov's sequential t-test. These shifts in hydroclimatic variables which occurred around 1977, 1989, 1997 and 2001, were synchronized in open and coastal waters, and were tracked by open water chlorophyll and copepods, but not by coastal plankton. North–central–south or open-coastal spatial breakdowns of the North Sea explained similar amounts of variability in most ecosystem indicators with the exception of diatom abundance and chlorophyll concentration, which were clearly better explained using the open-coastal configuration.     

气候变化对鲑鱼丰富度和繁殖影响的研究进展

鲑鱼是人类重要的食品来源之一,具有巨大的经济价值。同时,鲑鱼资源通过输送营养物质对维持溪流和陆地生态系统的结构、功能和过程等有重要作用。而鲑鱼是对温度极为敏感的冷水性鱼类,对气候变化的反应尤为明显。作者通过文献综述法梳理相关研究,从气候变化引起的温度变化、淡水和海洋环境的变化以及鱼病传播4个方面探讨对鲑鱼丰富度和繁殖的影响,并提出研究展望。研究显示:温度变化对鲑鱼的产卵和迁徙有较大影响;溪流和河流流量的改变会影响鲑鱼的迁徙和幼年鲑鱼的存活数量;温度和二氧化碳浓度的变化使海洋食物供应减少,鲑鱼丰富度降低;水温的升高使病原体增加,鲑鱼死亡率增加。本研究成果可为鲑鱼种群应对气候变化的研究及管理提供一定的理论支持。     

Late Holocene sea-level rise in Tampa Bay: Integrated reconstruction using biomarkers,pollen, organic-walled dinoflagellate cysts,and diatoms

A suite of organic geochemical, micropaleontological and palynological proxies was applied to sediments from Southwest Florida, to study the Holocene environmental changes associated with sea-level rise. Sediments were recovered from Hillsborough Bay, part of Tampa Bay, and studied using biomarkers, pollen, organic-walled dinoflagellate cysts and diatoms. Analyses show that the site flooded around 7.5 ka as a consequence of Holocene transgression, progressively turning a fresh/brackish marl-marsh into a shallow, restricted marine environment. Immediately after the marine transgression started, limited water circulation and high amounts of runoff caused stratification of the water column. A shift in dinocysts and diatom assemblages to more marine species, increasing concentrations of marine biomarkers and a shift in the Diol Index indicate increasing salinity between 7.5 ka and the present, which is likely a consequence of progressing sea-level rise. Reconstructed sea surface temperatures for the past 4 kyrs are between 25 and 26 ° C, and indicate stable temperatures during the Late Holocene. A sharp increase in sedimentation rate in the top ∼50 cm of the core is attributed to human impact. The results are in agreement with parallel studies from the area, but this study further refines the environmental reconstructions having the advantage of simultaneously investigating changes in the terrestrial and marine environment.     

Ecological characteristics of Walleye pollock eggs and larvae in the southeastern Bering Sea during the late 1970s

Walleye pollock (Theragra chalcogramma) is an ecologically and economically important groundfish in the eastern Bering Sea. Its population size fluctuates widely, driving and being driven by changes in other components of the ecosystem. It is becoming apparent that dramatic shifts in climate occur on a decadal scale, and these “regime shifts” strongly affect the biota. This paper examines quantitative collections of planktonic eggs and larvae of pollock from the southeastern Bering Sea during 1976–1979. Mortality, advection, and growth rates were estimated, and compared among the years encompassing the 1970s’ regime shift. These data indicate that pollock spawning starts in late February over the basin north of Bogoslof Island. Over the shelf, most spawning occurs north of Unimak Island near the 100 m isobath in early or mid April. Pollock eggs are advected to the northwest from the main spawning area at 5–10 cm/sec. Larvae are found over the basin north of Bogoslof Island in April, and over the shelf between Unimak Island and the Priblof Islands in May. Compared to 1977, the spawning period appeared to be later in 1976 (a cold year) and earlier in 1978 (a warm year) in the study area. At the lower temperatures in 1976, egg duration would be longer and thus egg mortality would operate over a longer period than in the other years. Mean larval growth appeared to be lower in 1976 than in 1977 and 1979. Estimated egg mortality rate in 1977 was 0.6 in April and 0.3 in early May.     

Diet of landlocked sockeye salmon (Oncorhynchus nerka) and trout in the Waitaki lakes,New Zealand

Landlocked sockeye salmon (Oncorhynchus nerka), ranging in fork length (FL) from 105 to 313 mm, were captured in fine‐mesh gill nets set in the limnetic zone of the Waitaki hydro lakes (44° 30′ S, 170° 10’ E) in the South Island, New Zealand. A total of 443 stomachs was examined and the frequency of occurrence, volume and weight of prey items calculated. In the Ahuriri Arm of Lake Benmore the principal food (54% by weight) was zooplankton (Boeckella dilatata) whereas in the Haldon Arm of Lake Benmore it was larval and juvenile common bullies (Gobiomorphus cotidi‐anus) (73% by volume). In Lake Waitaki in winter, salmon had eaten insects (43% by volume) with smaller amounts of snails (Potamopyrgus antipo‐darum, 23%) and bullies (24%). In Lake Ohau adult insects may be an important food. There were also variations in diet with season and fish size. The stomachs of 147 brown trout (Salmo trutta) and 181 rainbow trout (S. gairdnerii) caught in the same gill nets were also examined. In contrast to sockeye salmon stomachs they contained negligible amounts of zooplankton (< 1% by weight) and large amounts of aquatic insects (50–58% by weight in the Ahuriri Arm of Lake Benmore). Comparisons with juvenile sockeye salmon and kokanee in North American lakes are made. The impact of introductions of sockeye salmon into other New Zealand lakes is discussed.     

The Northern Oscillation Index (NOI): a new climate index for the northeast Pacific

We introduce the Northern Oscillation Index (NOI), a new index of climate variability based on the difference in sea level pressure (SLP) anomalies at the North Pacific High (NPH) in the northeast Pacific (NEP) and near Darwin, Australia, in a climatologically low SLP region. These two locations are centers of action for the north Pacific Hadley–Walker atmospheric circulation. SLPs at these sites have a strong negative correlation that reflects their roles in this circulation. Global atmospheric circulation anomaly patterns indicate that the NEP is linked to the western tropical Pacific and southeast Asia via atmospheric wave trains associated with fluctuations in this circulation. Thus the NOI represents a wide range of tropical and extratropical climate events impacting the north Pacific on intraseasonal, interannual, and decadal scales. The NOI is roughly the north Pacific equivalent of the Southern Oscillation Index (SOI), but extends between the tropics and extratropics. Because the NOI is partially based in the NEP, it provides a more direct indication of the mechanisms by which global-scale climate events affect the north Pacific and North America.The NOI is dominated by interannual variations associated with El Niño and La Niña (EN/LN) events. Large positive (negative) index values are usually associated with LN (EN) and negative (positive) upper ocean temperature anomalies in the NEP, particularly along the North American west coast. The NOI and SOI are highly correlated, but are clearly different in several respects. EN/LN variations tend to be represented by larger swings in the NOI. Forty percent of the interannual moderate and strong interannual NOI events are seen by the SOI as events that are either weak or opposite in sign. The NOI appears to be a better index of environmental variability in the NEP than the SOI, and NPH SLP alone, suggesting the NOI is more effective at incorporating the influences of regional and remotely teleconnected climate processes.The NOI contains alternating decadal-scale periods dominated by positive and negative values, suggesting substantial climate shifts on a roughly 14-year ‘cycle’. The NOI was predominantly positive prior to 1965, during 1970–1976 and 1984–1991, and since 1998. Negative values predominated in 1965–1970, 1977–1983, and 1991–1998. In the NEP, interannual and decadal-scale negative NOI periods (e.g. EN events) are generally associated with weaker trade winds, weaker coastal upwelling-favorable winds, warmer upper ocean temperatures, lower Pacific Northwest salmon catch, higher Alaska salmon catch, and generally decreased macrozooplankton biomass off southern California. The opposite physical and biological patterns generally occur when the index is positive. Simultaneous correlations of the NOI with north Pacific upper ocean temperature anomalies are greatest during the boreal winter and spring. Lagged correlations of the winter and spring NOI with subsequent upper ocean temperatures are high for several seasons. The relationships between the NOI and atmospheric and physical and biological oceanic anomalies in the NEP indicate this index is a useful diagnostic of climate change in the NEP, and suggest mechanisms linking variations in the physical environment to marine resources on interannual to decadal climate scales. The NOI time series is available online at: http://www.pfeg.noaa.gov.     

莱州湾拟哲水蚤Paracalanus sp.发育期丰度组成和个体大小季节变化

拟哲水蚤是莱州湾桡足类中周年出现的优势种;是上层鱼类幼鱼早期开口饵料的重要贡献者;掌握其种群动态分布特点;有利于了解湾内生物环境状况、次级生产力生产水平。基于2011年5月至20 12年4月采集的浮游生物资料的分析结果显示;莱州湾拟哲水蚤的发育期丰度组成和个体大小均表现了明显的季节变化。调查季节中;4-6月以成体居多;其他季节以桡足幼体CⅢ-CⅤ较多;其中10-11月和3月主要由后期桡足幼体CⅣ和CⅤ期构成。成体中以雌体为主导;雌/雄比介于1.46~9.62;该比值在3-4月最低;10-11月最高。拟哲水蚤各期桡足幼体和成体个体大小以4-5月最大、8月最小;并与水温表现出明显的负相关。拟哲水蚤桡足幼体和成体的总生物量月变化与丰度变化相同;即以8月值最高;3-5月值最低;月均值为2.69 mg/m3（以碳计）;日生产力估算值为0.74 mg/（m3·d）（以碳计）。由上述结果;推测莱州湾拟哲水蚤的种群结构周年变化主要可分3个阶段;4-6月是莱州湾拟哲水蚤开始繁殖期;7-8月为种群数量快速增长期;9月之后为种群增长相对停滞期。     

深沪湾末次冰期以来的地层结构及古环境演变

通过深沪湾高分辨率浅地层剖面声学地层和地质钻孔沉积地层的对比,并结合沉积物的粒度、微体古生物以及AMS14C测年的综合分析,揭示了研究区晚更新世末次冰期以来的地层层序,探讨了深沪湾的古环境演变。深沪湾高分辨率浅地层剖面自上而下划分的5个声学地层单元与钻孔岩芯划分的5个沉积地层单元具有较好的对应关系。8.2 ka BP左右,全新世海侵使得海水进入深沪湾海域,海平面低于现今海平面10~12 m,气候凉爽;7 ka BP左右海水到达现今海平面位置,并于6 ka BP左右到达最高,约比现今海平面高2~3 m,气候温暖湿润,这一时期,近岸大量裸子植物被海水淹没并被沉积物快速掩埋;5 850~5 642 a BP研究区温度降低,该降温活动持续到2 ka BP左右,气候凉爽干燥;2 ka BP以来温度逐渐上升,600 a BP左右有一个相对冷期,之后温度又逐渐上升至现今水平。     

Mnemiopsis leidyi (Ctenophora) in Narragansett Bay, 1975–1979: Abundance,size composition and estimation of grazing

Surveys of the distribution, abundance and size of the ctenophore Mnemiopsis leidyi were carried out in Narragansett Bay, R.I. over a 5-year period, 1975–1979. Yearly variations were observed in time of initiation of the ctenophore increase and maximum abundance. Biomass maxima ranged from 0·2 to 3 g dry weight m^?3 at Station 2 in lower Narragansett Bay while maximum abundance varied from 20 to 100 animals m^?3. Ctenophores less than 1 cm in length generally composed up to 50% of the biomass and 95% of the numerical abundance during the peak of the M. leidyi pulse. During the 1978 maxima and the declining stages of the pulse each year, 100% of the population was composed of small animals. M. leidyi populations increased earlier, reached greater maximum abundances, and were more highly dominated by small animals in the upper bay than toward the mouth of the bay. The averageclearance rate of M. leidyi larvae feeding on A. tonsa at 22°C was 0·36 l mg^?1 dry weight day^?1, with apparent selection for nauplii relative to copepodites. Predation and excretion rates applied to ctenophore biomass estimated for Narragansett Bay indicated that M. leidyi excretion is minor but predation removed a bay-wide mean of 20% of the zooplankton standing stock daily during August of 1975 and 1976. Variation in M. leidyi predation at Station 2 was inversely related to mean zooplankton biomass during August and September, which increased 4-fold during the 5-year period.