Foraging behaviors and the interaction of alewife,Alosa pseudoharengus,and bloater,Coregonus hoyi

Synopsis Alewife, Alosa pseudoharengus, and bloater, Coregonus hoyi, are common planktivores in Lake Michigan. Both alewife and bloater use a variety of feeding modes. Alewives can filter, gulp and particulate feed; bloaters can only gulp and particulate feed. We examined handling time per prey and probability of capture for alewife and bloater particulate feeding on Mysis relicta. Using these estimates and available data for filtering alewives, cost curves were derived for alewife and bloater particulate feeding and for alewife using all three modes of feeding. Alewives filter small prey relative to their own body size and particulate feed on larger prey. Feeding mode appears to be dependent on prey size and density and shifts in feeding mode are apparently based on maximizing biomass eaten per time. The ability to filter confers a competitive advantage on alewife when small prey are abundant as they were in the mid 1960s in Lake Michigan. If the zooplankton are large, bloater young-of-year do not suffer this relative disadvantage. In fact, large bloaters can consume prey on the bottom not available to alewife. This shifting competitive balance may explain, in part, the observed dynamics of alewife and bloater.     

Taxonomic differences in the essential fatty acid composition of groups of freshwater zooplankton relate to reproductive demands and generation time

1. Marked differences were observed in the total fatty acid concentrations and essential fatty acid (EFA) distributions of co-existing freshwater copepods and cladocerans in four large lake systems (lakes Michigan, Erie, Ontario and Champlain) over two growing seasons. These patterns appeared independent of lake seston EFA composition.
2. Compared to the cladocerans, calanoid and cyclopoid copepods contained significantly higher concentrations of total fatty acids and docosahexaenoic acid (DHA), an EFA abundant in fish. Calanoids and cladocerans contained similar levels of eicosapentaenoic acid (EPA), but cladocerans showed EPA : DHA ratios consistently greater than those of the available seston food source. Alpha-linolenic acid was most abundant in the herbivorous cladocerans, Daphnia and Holopedium , while the highest concentrations of arachidonic acid were found in the predatory cladocerans, Bythotrephes longimanus and Leptodora kindtii .
3. The distinct EFA accumulation patterns between cladoceran and copepod zooplankton suggest metabolic regulation of certain EFAs to meet the particular physiological demands and ecological strategies of these different zooplankton groups. Cladocerans may accumulate EPA directly from their diet, or through transformation of dietary materials to facilitate rapid somatic growth and enhance reproduction due to their short generation time. In contrast, copepods may retain DHA to increase their cell membrane fluidity in order to remain active over the winter due to their longer generation time and life cycle.
4. Consistent EFA differences between zooplankton groups may have implications regarding the somatic growth and reproductive success of different zooplankton taxa as well as the nutritional value of various zooplankton groups for larval and planktivorous fish.     

The effect of invertebrate diets on lipids, fatty acid composition and physiological condition of pike larvae (Esox lucius)

The present study tests two invertebrate diets for pike larvae reared in floating cages. One diet contained mainly copepods and cladocerans such as Eucyclops serrulatus and Bosmina longirostris , while the other diet contained Eucyclops serrulatus together with a small proportion of large Chydoridae and chironomid larvae. During the first days of the experiment, the amount of food distributed to the larvae was insufficient (9–17.1 prey per larva and per day) and led to an increase in the mortality. Triacylglycerol contents of fry were low and dietary fatty acids were either catabolised or, concerning PUFA such as 22:6(n-3), incorporated into phospholipids. From day 13 to the end of the study (day 22), the most abundant diets distributed were accompanied by an increase in triacylglycerol PUFA and in triacylglycerol contents of larvae. During the same period weight and length growth were better for larvae reared on the copepod plus Chydoridae and chironomid diet, than for larvae reared on the copepod and Bosmina diet. The use of a lipid condition index based on the triacylglycerol/sterol ratio suggested that Chydoridae and chironomids positively influenced the growth and nutritional condition of larvae. The effects of prey type in terms of PUFA composition on pike larvae are discussed.     

Life histories of some species of Chydoridae (Cladocera: Crustacea)

SUMMARY 1. Life history parameters of two species of Chydoridae ( Leydigia leydigi and Disparalona rostrata ) were studied in the field and in the laboratory.
2. For both species the number of juvenile instars was variable and tentatively related to initial size of neonate. Egg volume increased with increasing parent length and with decreasing temperature; possible advantages gained by the offspring are discussed in relation to invertebrate predation.
3. The life history strategy employed by the littoral Chydoridae is compared to that of large and small planktonic cladocerans. Small planktonic cladocerans and the Chydoridae (except the Eurycercinae and the Saycinae) produce large young relative to their size at maturity and mature early. However, unlike the small planktonic cladocerans, growth in the Chydoridae is curtailed after the onset of reproduction and in this they resemble the large planktonic cladocerans. This strategy may be related to vertebrate predation and the presence of vegetation in the habitat.     

Assessment of Top-Down and Bottom-Up Controls on the Collapse of Alewives (<Emphasis Type="Italic">Alosa pseudoharengus</Emphasis>) in Lake Huron

Food web models are powerful tools to inform management of lake ecosystems, where top-down (predation) and bottom-up (resource) controls likely propagate through multiple trophic levels because of strong predator–prey links. We used the Ecopath with Ecosim modeling approach to assess these controls on the Lake Huron main basin food web and the 2003 collapse of an invasive pelagic prey fish, alewife (Alosa pseudoharengus). We parameterized two Ecopath models to characterize food web changes occurring between two study periods of 1981–1985 and 1998–2002. We also built an Ecosim model and simulated food web time-dynamics under scenarios representing different levels of top-down control by Chinook salmon (Oncorhynchus tshawytscha) and of bottom-up control by quagga mussels (Dreissena rostriformis bugensis) and nutrients. Ecopath results showed an increase in the relative importance of bottom-up controls between the two periods, as production decreased across all trophic levels. The production of non-dreissenid benthos decreased most, which could cause decreases in production of pelagic prey fishes feeding on them. Ecosim simulation results indicated that the alewife collapse was caused by a combination of top-down and bottom-up controls. Results showed that while controls by Chinook salmon were relatively constant before alewife collapse, controls by quagga mussels and nutrients increased jointly to unsustainable levels. Under current conditions of low nutrients and high quagga mussel biomass, simulation results showed that recovery of alewives is unlikely regardless of Chinook salmon biomass in Lake Huron, which implies that the shrinking prey base cannot support the same level of salmonine predators as that prevailed during the 1980s.     

Seasonal feeding ecology of co-existing native and invasive benthic fish along a nearshore to offshore gradient in Lake Michigan

Relative abundance, diet composition and feeding strategy were determined for three benthic fish, the native deepwater sculpin Myoxocephalus thompsonii (Girard, 1851) and slimy sculpin Cottus cognatus (Richardson, 1836), and the invasive round goby Neogobius melanostomus (Pallas, 1814), along a nearshore to offshore gradient in southeastern Lake Michigan during March–December 2010, 2015, and 2016. Round goby were most abundant in the nearshore (<25 m), slimy sculpin were most abundant in the transitional zone (35–65 m), and deepwater sculpin were most abundant in the offshore zone (>75 m). Despite a large degree of spatial separation, some species did overlap, with slimy and deepwater sculpin occurring in sympatry throughout the year in the offshore and transitional zones, and round goby overlapping with both sculpin species seasonally in the transitional zone. Deepwater sculpin exhibited specialization on Mysis diluviana in all depth regions. Slimy sculpin in the offshore reduced diet overlap with deepwater sculpin by specializing on fish eggs during spring and fall, whereas in the transitional depth zone, there was considerable overlap between sculpin species due to the high importance of Mysis in diets. The invasive round goby had a mixed diet, with some diet overlap with native sculpin, especially slimy sculpin, in the transitional zone. In the nearshore zone, round goby displayed a generalized diet with many prey contributing to the diet, but the average contribution of any prey was generally low. Spatial separation and variable feeding strategies help reduce, but not eliminate shared resource use amongst these benthic fish in Lake Michigan.     

Gape limitation and piscine prey size‐selection by yellow perch in the extreme southern area of Lake Michigan, with emphasis on two exotic prey items

The trophic linkage between yellow perch Perca flavescens and two exotic prey items, alewife Alosa pseudoharengus and round goby Neogobius melanostomus , was investigated in the extreme southern area of Lake Michigan during the summer of 2002. Yellow perch ≥100 mm total length, L _T( n  = 1293) exhibited size selective feeding, with 148 fish containing round gobies and 120 fish containing alewives. The mean round goby L _T, preyed on by yellow perch, was 23% of the predator L _T, with a range of 7 to 47%, and mean alewife L _T was 32% of yellow perch L _T, with a range of 18 to 46%. Although the selection of prey size by yellow perch increased proportionally with yellow perch L _T, prey consumed appeared smaller than theoretically possible based on gape size.     

Population dynamics of Bythotrephes cederstroemii in south-east Lake Michigan 1995–1998

1. Population characteristics (density, size, reproductive patterns) of the predatory cladoceran Bythotrephes cederstroemii in south-east Lake Michigan were monitored at an offshore station (110 m) in 1995–98 and at a nearshore station (45 m) in 1997–98.
2. The mean density of B. cederstroemii at the offshore station was generally highest in July–September (145–914 m⁻²) and at the nearshore station in October–November (168–1625 m⁻²). In 1995 and 1998, density was also high at the offshore station in November (211–284 m⁻²). Fish predation may limit B. cederstroemii in nearshore regions in the summer. The maximum annual densities of B. cederstroemii for 1995–98 were generally similar to those reported from the late 1980s, when the species arrived in Lake Michigan.
3. Body size increased rapidly each year to a maximum in August. Thereafter, body size declined and converged for stage-2 and 3 individuals, suggesting food scarcity or size-selective fish predation was affecting large individuals.
4. Most reproduction occurred asexually (90%), and by stage 2 or 3 females (99%). Asexual brood size was highest when B. cederstroemii first appeared each year, and decreased in August, when larger neonates were produced. There appeared to be differences in reproductive mode for stage 2 and 3 females, with a higher percentage of stage 2 females reproducing sexually.     

Selective Feeding of age-0 Arctic Grayling in Lake-Outlet Streams of the Northwest Territories, Canada

A paucity of information exists on the diet of Arctic grayling, Thymallus arcticus, particularly for young-of-the-year (YOY). We examined the diet of YOY Arctic grayling in relation to food availability, in the Barrenlands region of the Northwest Territories, Canada, where lake-outlet streams serve as nursery habitat for these fish. Given the small size of YOY grayling and the abundance of lake-derived microcrustacea in the drift of these lake-outlet streams, we anticipated that these prey would make up a major component of the YOY's diet. Food selectivity by YOY grayling, however, was strongly sized-biased; although microcrustacea dominated the drift, YOY primarily consumed larger taxa, especially Chironomidae and Simuliidae. Even among these taxa, grayling tended to select the larger individuals. As they grew, YOY grayling took larger numbers of both large and small prey, particularly the larger invertebrates, although prey size range did not change after mid-July. Selection of pupae and avoidance of Ephemeroptera suggest that prey characteristics other than size also contribute to selectivity by YOY grayling. The relatively limited consumption of terrestrial invertebrates and other large prey may reflect the small sizes of fish in this arctic study, as well as differences in prey availability. Despite the abundance of lake-derived prey, instream production of invertebrates should largely determine the productive capacity of Barrenlands streams as fish habitat.     

Food and habitat partitioning between young-of-year alewives and rainbow smelt in southeastern Lake Ontario

Synopsis Thermal and depth distributions, diets and time of feeding of young-of-year (YOY) alewives and YOY rainbow smelt were compared for evidence of resource partitioning in southeastern Lake Ontario. YOY alewives were largely epilimnial during August and September, but moved toward the bottom during fall turnover. Alewives were most abundant in the warmest available water. YOY rainbow smelt were concentrated at depths between 10 and 30 m in August and September, but moved into deeper water at fall turnover. Depth distribution of YOY smelt was correlated with temperatures of 8–12°C Both species fed predominantly during day on zooplankton during August and September. Cyclopoid copepods were the most common prey, but bosminids, eubosminids, and occasionally calanoid copepods were frequently eaten. As YOY rainbow smelt grew (> 60 mm), they consumed more Mysis relicta and amphipods, which became the major prey of rainbow smelt by November. YOY alewives consumed mostly zooplankton in all months. Diet overlap of the two species was greatest in warm water (> 12.0° during October (94.3% similarity) and August (80.0% similarity) and lowest in November (16.9% similarity). Positive size-selection on zooplankton was found in all months for YOY rainbow smelt, but only in late September through November for YOY alewives. Thus, during thermal stratification, the species were spatially segregated by water temperature but had a high degree of overlap in time of feeding and types of prey eaten. In contrast, after fall turnover there was a greater separation in diet but a higher overlap in habitat use.     

Perch or plankton: top‐down control of Daphnia by yellow perch (Perca flavescens) or Bythotrephes cederstroemi in an inland lake?

1. Seasonal termination of the vernal clear-water phase in Long Lake, Grand Traverse Co., Michigan coincided with severe size-selective predation on juvenile Daphnia pulicaria from 0.8 to 1.8 mm in length. This could be caused by predation by age-0 yellow perch ( Perca flavescens ) or by the exotic predatory zooplankter Bythotrephes cederstroemi .
2. During the initial decline of Daphnia , Ivlev's electivity coefficient for yellow perch from 15.0 to 20.0 mm in length was 0.50 for copepods and −0.75 for D. pulicaria .
3. Bioenergetics modelling of both yellow perch and Bythotrephes demonstrates that, during the initial Daphnia decline, Bythotrephes consumed 1.5–5 times greater total mass than yellow perch. Furthermore, models in which Bythotrephes consumed juvenile Daphnia were more consistent with the timing of the Daphnia decline than those in which yellow perch consumed juvenile Daphnia .
4. The invasion of Bythotrephes into Long Lake seems to be a significant perturbation, introducing effects that propagate throughout the food chain. Bythotrephes created a possible bottleneck for age-0 yellow perch in late June by suppressing Daphnia .     

The predatory Chironomidae of an iron-rich stream: feeding ecology and food web structure

Abstract. 1. Three species of Tanypodinae (Chironomidae) were found in an acid and iron-rich stream in southern England. Maximum abundance was achieved in summer and they were sparse at other times. Individuals were aggregated on the stream bed and were overrepresented in accumulations of leaf litter.
2. The diets of all three species consisted of a mixture of prey (prominently detritivorous chironomid larvae) and detritus. More detritus and fewer prey were taken in winter than in summer.
3. When comparing large tanypod species with small and, intraspecifically, late instars with early, the proportion of guts containing prey increased with increasing body size.
4. Stonefly larvae were more prominent in the diet of Zavrelimyia barbatipes (Kieffer) in summer than in winter but for the other two species the reverse was true. A bigger proportion of Trissopelopia longimana (Staeger) guts contained prey in early summer than in August whereas more Macropelopia goetghebueri (Kieffer) guts contained prey in August. This was apparently a consequence of seasonal differences in the distribution of body size among the populations of these two species.
5. The stream contains two further common predators, Plectrocnemia conspersa (Curtis) and Sialis fuliginosa Pict. These are important predators of tanypod larvae but might also compete with them since they severely deplete populations of prey taken in common.
6. Analysis of the food-web in Broadstone Stream reveals remarkably high values of connectance (C and C_max) and of species richness times connectance (SC_max). Such characteristics are theoretically associated with fragile and dynamically unstable food webs, and may be found in 'constant' environments. There is also an apparently unusual prevalence of omnivory in the community.     

Prey selection by the dobsonfly larva, Protohermes grandis (Megaloptera: Corydalidae)

SUMMARY 1. Prey selection by the dobsonfly larva, Protohermes grandis (Thunberg), was studied in stony riffles of the Yataro River, central Japan. The density, size distribution and taxonomic composition of available prey were assessed for 2 years. In order to know the encounter rate between prey and this ambush predator, prey mobility was also estimated from patterns of colonization of experimentally detiuded stones.
2. Foregut analyses revealed that maximum size of prey eaten increased with larval size, and large larvae did not take the smallest prey in spite of high availability in all seasons.
3. Charnov's (1976) optimal diet model quantitatively predicted such size-selective feeding under seasonally fluctuating conditions of water temperature and prey availability. Larvae maximized the feeding rate by selecting prey.
4. Maximum width of prey eaten coincided approximately with larval mandible length. Mandible size seemed to play an important role in the selection of prey in the optimal size range.     

The environmental basis of diet variation in pumpkinseed sunfish, Lepomis gibbosus, and largemouth bass, Micropterus salmoides, along an Iberian river basin

The diet composition of largemouth bass, Micropterus salmoides, and pumpkinseed sunfish, Lepomis gibbosus, and its relation to environmental factors was studied in intermittent watercourses of the lower Guadiana basin (southern Iberia) during the particularly dry summer of 1994. Overall, both species took food items similar to those found in their North American and acclimatised ranges, with pumpkinseed consuming invertebrates (chiefly Chironomidae) and bass preying on invertebrates (chiefly Micronecta meridionalis) and fish (chiefly L. gibbosus and Gambusia holbrooki). Both bass and pumpkinseed were opportunistic feeders, feeding on the most frequent and abundant prey. However, Diptera larvae were apparently preferred by pumpkinseed and avoided by bass, while the contrary occurred with respect to Heteroptera. The two main fish prey of bass were eaten on the basis of random encounter. The relative abundance of macro-prey (i.e. fish and Atyephira desmarestii) were the principal environmental variables constraining bass dietary variation along the basin. On the contrary, pumpkinseed diet variation was mainly related to pumpkinseed size and the presence of piscivorous bass nearby, although habitat size and cyprinid abundance were also influential. This revised version was published online in July 2006 with corrections to the Cover Date.     

Reaction distance of Bythotrephes longimanus, encounter rate and index of prey risk for Harp Lake, Ontario

SUMMARY 1. Some predatory zooplankton, such as the large cladoceran Bythotrephes longimanus (Leydig), have a large compound eye, hence vision may play an important role in prey detection and encounter. Our objectives were to quantify, under different light regimes, the reaction distance of Bythotrephes to an assortment of zooplankton prey and to model encounter rate with prey from Harp Lake, Ontario. Reaction distance to prey increased at higher light intensity.
2. Results from the encounter model show that small, slow-moving prey faced the greatest risk from Bythotrephes and most encounters occurred in the upper 10 m of the water column throughout the 24-h period. The model was highly sensitive to ambient light. Encounter rate and prey risk were two to three orders of magnitude more sensitive to swimming velocity of the predator than to that of the prey.     

Food selectivity and diel vertical distribution of Chaoborus edulis (Diptera, Chaoboridae) in Lake Malawi

1. Diel diet and vertical distribution patterns of the larval instars of Chaoborus edulis were studied in deep water near the central part of Lake Malawi, Africa.
2. First instar larvae contained very little food in their crops and probably depended on reserves from the egg. Second, third and fourth instars fed on zooplankton and were size-selective in their feeding. The mean size of prey eaten by the three instars was significantly different from each other, with larger instars feeding on larger prey. Smallest available prey was selected against and the upper size of prey was probably constrained by larval gape. Nauplii were not found in any of several thousand larvae examined. Phytoplankton did not form a significant part of the diet.
3. There was a progressive and related increase in diel periodicity in feeding and vertical migrations of successive instar stages. Fourth instars migrated particularly large distances. Such migrations removed them from their zooplankton food supply but avoided predators. A refuge from predators is probably found in or near the permanent zero oxygen boundary, at depths greater than 200 m.     

Influence of Chironomus plumosus larvae on ammonium flux and denitrification (measured by the acetylene blockage- and the isotope pairing-technique) in eutrophic lake sediment

Bythotrephes invaded Harp Lake, Ontario, Canada, in the early 1990s. Here we describe seasonal changes in the size, abundance and life history of the invader. The weight of Bythotrephes could be accurately estimated (r² =0.90) from the length of its body and the state of development of its brood, but substantial (22%) corrections for shrinkage in sugarformalin were required. The seasonality of the size and abundance of Bythotrephes was apparently regulated by the interaction of fish predation, temperature-mediated growth, the availability of the invader's prey, and switches in its reproductive biology. The vernal increase in the growth of the Bythotrephes population lagged behind increases in water temperature and abundances of preferred prey, most probably because of predation by lake herring (Coregonus artedi). Bythotrephes abundance increased rapidly in early July; however, the population maximum was brief. After the abundance of their preferred prey crashed in mid-July, Bythotrephes body size declined, and adult females switched from parthenogenic to resting egg production. Population size declined in consequence. In the summer, female Bythotrephes committed most of their energy to growth between instar I and II, but to egg production for instars II and III. The biology of the Harp Lake population differed from that observed in other recently invaded lakes in several ways – the lag in population growth in the spring, the small change in body size between instars II and III, the decline in body size in August, and the co-incident switch by mature females to resting egg production. These inter-lake differences in biology indicate we have more to learn about Bythotrephes before we can predict its influence on the inland lakes of North America.     

Changes in the crustacean zooplankton community of Harp Lake, Canada, following invasion by Bythotrephes cederstrœmi

1. Detecting the impacts of invading Bythotrephes cederstrœmi (Crustacea, Onychopoda, Cercopagididae) on zooplankton in North American lakes has been hampered by the brevity of pre-invasion data, and by the difficulty of distinguishing the effects of the invader from other stresses. The data from Harp Lake in Ontario, Canada, circumvent these difficulties. Bythotrephes appeared in the lake in 1993. There is a 15-year pre-invasion data set, and no significant complicating concurrent stresses.
2. The species composition and the size structure of the crustacean zooplankton community of Harp Lake changed after the invasion. Several small species either declined dramatically in abundance (e.g. Bosmina longirostris , Tropocyclops extensus ) or disappeared ( Chydorus sphaericus , Diaphanosoma birgei , Bosmina ( Neobosmina ) tubicen ). In contrast the abundance of the larger cladocerans Holopedium gibberum and Daphnia galeata mendotae and the hypolimnetic copepod Leptodiaptomus sicilis increased. Several univariate and all multivariate summarizations of zooplankton abundance, biomass and size structure highlighted the uniqueness of the post-invasion community.
3. The alterations in the zooplankton community could not be attributed to changes in lake acidity, thermal regimes, penetration by ultraviolet light, nutrient status, fish stocking or the abundances of native invertebrate predators, but they were correlated with Bythotrephes abundance, both within and among years. Hence, we hypothesize that the invasion by Bythotrephes has significantly altered the crustacean zooplankton community of Harp Lake.     

Impact of Bythotrephes longimanus on zooplankton assemblages of Harp Lake, Canada: an assessment based on predator consumption and prey production

1. Following the 1993 invasion of Harp Lake by Bythotrephes longimanus a number of small-bodied zooplankton declined dramatically compared to pre-invasion densities, and some larger species increased. 2. To test whether these changes were caused by Bythotrephes, we measured its consumption of zooplankton in 1995 and compared this to production by the prey species with which it overlapped spatially and temporally. 3. On a seasonal basis Bythotrephes consumption was 199 mg dry mass (DM) m⁻², or 25% of zooplankton production which was 783 mg DW m⁻². However, for some species, such as the small copepod Tropocyclops extensus , consumption greatly exceeded production—this was one of the species noted to decline in Harp Lake following the appearance of Bythotrephes . By contrast, for the larger cladocerans Daphnia galeata mendotae and Holopedium gibberum , consumption was much less than production—these species were observed to increase following the invasion. 4. Our results thus support the hypothesis that zooplankton changes in Harp Lake were caused by Bythotrephes . We speculate that lower quality prey remaining in Harp Lake may lead to reduced densities of Bythotrephes , or impose strong selective forces that lead to new adaptations by this predator.