Diet dynamics of the juvenile piscivorous fish community in Spirit Lake, Iowa, USA, 1997–1998

Abstract – We assessed temporal dynamics and variation among species and age-classes in the diets of age 0 and age 1 piscivorous fish species in Spirit Lake, Iowa, USA during 1997 and 1998. Species included walleye Stizostedion vitreum , yellow perch Perca flavescens , smallmouth bass Micropterus dolomieui , largemouth bass Micropterus salmoides , black crappie Pomoxis nigromaculatus and white bass Morone chrysops. Thirty taxa were identified in diets, including 12 species of fish. We found dramatic differences in diets among species, among age-classes within species and over time. Walleye, largemouth bass, smallmouth bass and white bass were piscivorous at age 0. Black crappie began piscivory at age 1. Yellow perch also began piscivory at age 1, but fish were a very small fraction of age-1 diets. The primary temporal pattern, seen in several species and age-classes, was an increase in piscivory from spring to fall. This pattern was due to the lack of small, age-0 prey fish in spring. Although some patterns were evident, the taxonomic composition of the diets of all species was highly variable over time, making generalizations difficult. A surprising result was the absence of yellow perch in the diet of age-0 walleye, despite their abundance in Spirit Lake and prominence in diets of age-1 walleye and other age 1-piscivores. Age-0 yellow perch were consistently too large to be eaten by age-0 piscivores, which preyed primarily on invertebrates and smaller fish such as johnny darters Etheostoma nigrum and age 0 bluegill Lepomis macrochirus. This finding suggests that predator-prey interactions and resulting population dynamics may be quite different in Spirit Lake than in other systems dominated by walleye and yellow perch.     

Piscivory in larval perch (Perca fluviatilis): mechanisms structuring larval roach (Rutilus rutilus) cohorts

Abstract – Perch ( Perca fluviatilis ) can act as a piscivore from larval stage VI (body size 10.3 mm) on newly hatched larval roach ( Rutilus rutilus ), bream ( Abramis brama ) and smaller siblings of its own cohort. Consumption rates at this stage were approx. 0.5 prey/perch*h at 21°C. Larval perch predation was strongly gap-limited, and the maximum size of roach consumed by perch (perch length interval 10.3–62.0 mm) under experimental conditions followed the linear regression, P_prey-max.=0.478*L_Pred.+1.829 ( r ²=0.99, P <0.001, n =12). Under experimental conditions, predatory 0⁺ perch substantially affected the size distributions of 0⁺ roach prey cohorts, since smaller prey individuals were predated more frequently than larger ones. In both unimodal and bimodal size distributions of prey roach, the distributions changed according to the maximum prey size consumed by the added predatory perch. Unimodal prey distributions were positively skewed when piscivorous perch were added compared with controls without predators. According to the size distributions of lake-living 0⁺ roach and 0⁺ perch and the relative size difference between prey and predator, the vulnerability of 0⁺ roach cohort to 0⁺ perch predation changed from June to September. Prey vulnerability was extremely sensitive to the relative size difference between predator and prey. Therefore differences in hatching time and growth rates between the two species will strongly influence the potential for predator-prey interactions. ^Note     

Predation as a determinant of size structure in populations of crucian carp (Carassius carassius) and tench (Tinea tinea)

Predation is an important structuring force in many fish assemblages and may have large, direct lethal effects on prey fish populations but also may affect growth rate and survival indirectly through shifts in behavioral patterns. Some species appear more sensitive to predation and earlier studies in a limited set of Finnish ponds suggests that crucian carp (Carassius carassius) is very sensitive to predation, which results in changes in population size-structure and density. In this study we tested the effect of piscivore presence or absence on crucian carp populations by using data from a large number (>600) of Swedish lakes across a wide geographical area and ranging in size from small ponds to large lakes (0.02–4400 ha). We also included a second species, tench (Tinea tinea), in our analyses. The population size structures of crucian carp and tench were clearly related to the presence or absence of piscivores. In lakes and ponds without piscivores. populations were dominated by small-bodied individuals. When piscivores were present, populations consisted almost exclusively of large individuals. Densities of crucian carp and tench were much higher in ponds and lakes without piscivores. There were no differences in size structure or relative abundance when catches from electrofishing were compared with gill net catches, suggesting that our results are not an effect of a behavioral shift in the presence of piscivores.     

Diets of double-crested cormorants in the Lake Winnebago System,Wisconsin

Double-crested cormorant Phalacrocorox auritus Lesson (cormorant) populations have increased throughout the Great Lakes region of North America causing concern related to the impact of cormorant predation on fish communities. A recent decline in yellow perch Perca flavescens (Mitchill) abundance within the Lake Winnebago System, Wisconsin, USA, prompted an assessment of cormorant diets to evaluate potential effects of cormorant predation on the sportfish community. Diets were collected from 883 cormorants (417 from Lake Winnebago and 466 from Lake Butte des Morts) between 2015 and 2017. Cormorant diets on both waterbodies consisted mostly of freshwater drum Aplodinotus grunniens Rafinesque and gizzard shad Dorosoma cepedianum (Lesueur). Yellow perch and walleye Sander vitreus (Mitchill) observations were infrequent and represented < 5% of cormorant diets by weight each year. Under current conditions, cormorant predation likely has minimal impact on the Lake Winnebago sportfish community, but more research is needed to assess potential impacts on Lake Butte des Morts.     

Spatio‐temporal,ontogenetic and interindividual variation of age‐0 diets in a population of yellow perch

Dietary niches of fishes have traditionally been evaluated at the population level, with diet pattern central tendencies compared spatio‐temporally among habitats and populations. More recently, however, studies have emphasised the importance of within‐population diet variation and niche partitioning. Several studies have examined diets of young yellow perch (Perca flavescens) at the population level and have described an ontogenetic transition from zooplankton to benthic prey during the first year of life. However, independent of ontogenetic diet shifts, intrapopulation variation of young yellow perch diets remains largely unexplored. We quantified patterns of diet composition in age‐0 yellow perch collected from Saginaw Bay, Lake Huron, USA during July–October, 2009 and 2010. We observed substantial variation in diet composition among individuals across and within sites, but found relatively weak evidence indicating an ontogenetic diet shift. Zooplankton were the dominant prey for age‐0 yellow perch on most occasions, and individual diets were composed primarily of either zooplankton (e.g. Daphnia spp., Calanoida) or benthic (i.e. Chironomidae larvae, Chydoridae) prey. These patterns were not simply attributable to differences in prey availability and ontogenetic diet shifts, because a) not only diet composition, but also prey selectivity (Chesson's α) varied among sites and b) individual and spatial diet differences were evident independent of ontogeny. Within‐cohort differences in diet composition may be an important, but often overlooked, phenomenon with implications for cumulative trophic interactions and intracohort growth and survival among young fish.     

Diet dynamics of the adult piscivorous fish community in Spirit Lake, Iowa, USA 1995–1997

Abstract – Diets of adults of six important piscivorous fish species, black crappie Pomoxis nigromaculatus , largemouth bass Micropterus salmoides , northern pike Esox lucius , smallmouth bass Micropterus dolomieui , walleye Stizostedion vitreum , and yellow perch Perca flavescens were quantified in Spirit Lake, Iowa, USA from May to October in 1995–1997. Forty-one prey taxa were found in the diets of these species, including 19 species of fish. The most important prey taxa overall were yellow perch, amphipods and dipterans. Diets of northern pike and walleye were dominated by yellow perch. Largemouth bass diets included large percentages of both yellow perch and black bullhead Ameiurus melas . Smallmouth bass diets included large percentages of both yellow perch and crayfish. Black crappie and yellow perch diets were dominated by invertebrates, primarily amphipods and dipterans. There were pronounced differences in diets among species, among size classes within species and over time. Most of the dominant prey taxa we documented in the diets of piscivorous species were in accordance with previous studies, but a few deviated significantly from expectations. Many of the temporal diet changes were asynchronous among piscivorous species and size classes, suggesting different responses to common prey resources over time.     

The shift to smaller zooplankton in Lake Victoria cannot be attributed to the ‘sardine’ Rastrineobola argentea (Cyprinidae)

After the population increase of introduced Nile perch (Lates niloticus) in Lake Victoria, a decrease in the relative abundance of herbivorous cladocerans and calanoids was reported for the pelagic zone. The shift from large herbivores to small-bodied predatory cyclopoids has been attributed to the increase of the zooplanktivorous cyprinid dagaa (Rastrineobola argentea), which rose concomitantly with the Nile perch. However, we observed that in the Mwanza Gulf of the lake cyclopoids dominated the zooplankton community already before the rise of dagaa. Furthermore, there are indications that dagaa takes fewer calanoids than expected from their representation in the environment, although a slight preference for cladocerans may exist. Calanoids dominated the diet of small Nile perch of about 4 cm length, but juvenile Nile perch did not occur in the study area until after the size shift in zooplankton. The lumped biomass of dagaa and small Nile perch is lower than the biomass of the original haplochromine-dominated community of zooplanktivorous fish. Thus, the decline of large zooplankters in the lake cannot be explained by intensified predation after the replacement of zooplanktivorous haplochromines by dagaa and Nile perch. Increased eutrophication, resulting in a shift in predominance from diatoms to cyanophytes, is suggested as an alternative explanation for the shift in zooplankton composition.     

Effects of intensive fishing on the perch population in a large oligotrophic lake in eastern Finland

The perch population in Lake Höytiäinen was intensively fished to reduce the density of the population and hence also the predation pressure by perch on vendace larvae. A hypothesis suggests that this predation can prevent recovery of commercially important vendace stock from a state of low-density. In the 1990s the density of the perch population in Lake Höytiäinen has increased, while the vendace stock has been sparse. Intensive fishing was conducted by professional fishermen with a paired bottom trawl, seine net, hoop net and small fish traps and by recreational fishermen with small fish traps. The size of perch population during the intensive fishing period was studied by test fishing with multi-mesh gillnets and the Leslie method in which trawl YPUE was regressed on the cumulative yield. The results suggest that the population size clearly diminished in the area where the fishing pressure was strongest. YPUE of test fishing decreased ca. 30% whereas the Leslie method gave almost a decrease twice as large as that of the former. Mean weight of perch increased in the trawl and test fishing catches during the intensive fishing period. The most effective fishing gear was trawl (62% of the total yield), but importance of trap net fishing by recreational fishermen was also high as they caught 22% of the total yield.     

Diet overlap of non‐native alewife with native yellow perch and spottail shiner in nearshore waters of southwestern Lake Michigan, 2000–2007

Abstract – Lake Michigan has experienced many ecological changes as a result of introductions of non‐native species. Arguably the most significant was that of alewife (Alosa pseudoharengus), yet studies on diet overlap with native species are lacking. We analysed diet trends of alewife, spottail shiner (Notropis hudsonius) and yellow perch (Perca flavescens) collected in summer and fall 2000–2007 near Waukegan, IL, in 3–10 m depths. Mean percentage composition by dry weight for 23 prey taxa was used in multivariate analysis to test whether diet differed across species and size classes. We also tested whether zooplankton and benthic invertebrate community composition changed over time. Fish diets were similar over all years but differed seasonally. In summer, diets of large alewife were similar to both small alewife and small yellow perch, with Bosminidae, chironomid larvae and copepods as primary common prey. During fall, alewife and yellow perch size classes exhibited strong intraspecific diet overlap, while there was low diet overlap between species. Primary distinctions between species’ diets in fall were higher consumption of amphipods by yellow perch and dreissenids by spottail shiners compared to alewife, which consumed higher proportions of zooplankton. Overall, high yellow perch diet overlap with alewife during summer and with their larger conspecifics during fall could lead to negative implications for yellow perch growth before the critical overwintering period. Detailed insights into diet overlap and prey availability are critical first steps in understanding competitive interactions between native and non‐native fish that dominate the nearshore community in southwestern Lake Michigan.     

Predation by native sunfishes (Centrarchidae) on the invasive crayfish Orconectes rusticus in four northern Wisconsin lakes

Tetzlaff JC, Roth BM, Weidel BC, Kitchell JF. Predation by native sunfishes (Centrarchidae) on the invasive crayfish Orconectes rusticus in four northern Wisconsin lakes.
Ecology of Freshwater Fish 2011: 20: 133–143. © 2010 John Wiley & Sons A/S Abstract – Introduced rusty crayfish (Orconectes rusticus) often establish abundant populations in lakes where they negatively impact native food webs. However, rusty crayfish do not become abundant in all lakes. Fish predation has been posed as one potential mechanism limiting rusty crayfish density, but there is limited empirical evidence to support this hypothesis. Using a case study approach, we quantified the extent and timing of predation on nonnative rusty crayfish by three native centrarchids in four northern Wisconsin lakes with contrasting crayfish densities. We estimated population‐level predation effects on rusty crayfish using bioenergetics models in combination with fish and crayfish abundance estimates. Bluegill (Lepomis machrochirus) and pumpkinseed (Lepomis gibbosus) consumed young‐of‐year crayfish exclusively whereas rock bass (Ambloplites ruspestris) consumed a wide size range of crayfish. Rusty crayfish were found in Lepomis diets primarily in late June and early July while rock bass consumed crayfish throughout the summer. Per individual, rock bass consumed more crayfish biomass than Lepomis, but at the population‐level, higher Lepomis densities resulted in greater numbers of crayfish and more crayfish biomass consumed by Lepomis than rock bass. Lepomis populations consumed 8 × , 14 × , 22 × and 120 × more individual crayfish than rock bass populations in the four study lakes. Bioenergetics simulations revealed that dense populations of Lepomis were capable of consuming a large percentage of the juvenile crayfish population in two of our study lakes. In systems with high Lepomis densities, Lepomis consumption of juvenile rusty crayfish may play a significant role in the invasion and establishment dynamics of rusty crayfish.     

Predation risk allocation or direct vigilance response in the predator interaction between perch (Perca fluviatilis L.) and pike (Esox lucius L.)?

Abstract –  Predation risk allocation hypothesis predicts that a prey's response to predator depends on prey's previous experience on predator. Here we tested whether the group of three perch respond differentially to pike, predator of perch, depending on the timing of high constant (HC) and high unpredictable (HU) risk periods within low constant risk periods in short-term (10 h) experiments, and whether the response is stronger during a HU risk period than during a HC risk period. Perch clearly erected the dorsal fin in response to predation risk treatments (pike odour only, odour and visible pike). Decrease in activity and increase in shoaling behaviour were observed mainly during high risk periods. However, the perch's responses to pike did not differ statistically between periods of various levels of predation risk or depending on the timing of high risk situations within constant low risk periods, and thus, suggesting that perch respond mainly to changes in the current predation risk.     

Population‐level effects of egg predation on a native planktivore in a large freshwater lake

Using a 37‐year recruitment time series, we uncovered a field pattern revealing a strong, inverse relationship between bloater Coregonus hoyi recruitment success and slimy sculpin Cottus cognatus biomass in Lake Michigan (United States), one of the largest freshwater lakes of the world. Given that slimy sculpins (and deepwater sculpin Myoxocephalus thompsonii) are known egg predators that spatiotemporally overlap with incubating bloater eggs, we used recently published data on sculpin diets and daily ration to model annual bloater egg consumption by sculpins for the 1973–2010 year‐classes. Although several strong year‐classes were produced in the late 1980s when the proportion of eggs consumed by slimy sculpins was extremely low (i.e., <0.001) and several weak year‐classes were produced when the proportion of bloater eggs consumed was at its highest (i.e., >0.10–1.0), egg predation failed to explain why recruitment was weak for the 1995–2005 year‐classes when the proportion consumed was also low (i.e., <0.02). We concluded that egg predation by slimy and deepwater sculpins could have limited bloater recruitment in some years, but that some undetermined factor was more important in many other years. Given that slimy sculpin densities are influenced by piscivorous lake trout Salvelinus namaycush, the restoration of which in Lake Michigan has lagged behind those in lakes Superior and Huron, our study highlights the importance of an ecosystem perspective when considering population dynamics of fishes.     

The effects of mechanically reducing northern pike density on the sport fish community of West Long Lake, Nebraska, USA

Abstract  Removal of piscivores may affect population size structure, abundance and growth rates of the remaining fish community. A total of 572 northern pike, Esox lucius L., was removed from 25-ha West Long Lake, Nebraska from 2003 to 2006. The northern pike population was reduced from 36 fish ha⁻¹ and 22 kg ha⁻¹ in 2002 to 20 fish ha⁻¹ and 9 kg ha⁻¹ in 2006. The northern pike population shifted to dominance by smaller individuals. Relative abundance increased for bluegill, Lepomis macrochirus Rafinesque, but remained similar between years for largemouth bass, Micropterus salmoides (Lacepède), and yellow perch, Perca flavescens (Mitchill). Bluegill and yellow perch size structure increased probably because of reduced predation by northern pike. The effects of reduced northern pike density on largemouth bass were less evident. The largemouth bass population may be returning to higher density with smaller size structure observed before pike were introduced. The sport fish community responded positively to the removal of northern pike in this small lake.     

Feeding ecology of the walleye (Percidae,Sander vitreus), a resurgent piscivore in Lake Huron (Laurentian Great Lakes) after shifts in the prey community

Recovering populations of piscivores can challenge understanding of ecosystem function due to impacts on prey and to potentially altered food webs supporting their production. Stocks of walleye (Percidae, Sander vitreus), an apex predator in the Laurentian Great Lakes, crashed in the mid‐1900s. Management efforts led to recovery by 2009, but recovery coincided with environmental and fish community changes that also had implications for the feeding ecology of walleye. To evaluate potential changes in feeding ecology for this apex predator, we assessed diets in the main basin of Lake Huron and in Saginaw Bay, a large embayment of Lake Huron, during 2009–2011. Walleye switched their diets differently in the main basin and Saginaw Bay, with non‐native round goby (Gobiidae, Neogobius melanostomus) and rainbow smelt (Osmeridae, Osmerus mordax) more prevalent in diets in the main basin, and invertebrates, yellow perch (Percidae, Perca flavescens) and gizzard shad (Clupeidae, Dorosoma cepedianum) more prevalent in diets in the bay. Feeding strategy plots indicated that there was a high degree of individual specialisation by walleye in the bay and the main basin. Bioenergetic simulations indicated that walleye in Saginaw Bay need to consume 10%–18% more food than a walleye that spends part or all of the year in the main basin, respectively, in order to achieve the same growth rate. The differences in diets between the bay and main basin highlight the flexibility of this apex predator in the face of environmental changes, but changes in diet can alter energy pathways supporting piscivore production.     

Does morphological variation between young‐of‐the‐year perch from two Swedish lakes depend on genetic differences?

Heynen M, Hellström G, Magnhagen C, Borcherding J. Does morphological variation between young‐of‐the‐year perch from two Swedish lakes depend on genetic differences?
Ecology of Freshwater Fish 2010: 19: 163–169. © 2009 John Wiley & Sons A/S Abstract – Different local environmental conditions have often been found to generate phenotypic diversity. In the present study we examined morphological differences between young‐of‐the‐year perch from two lake populations with differences in size‐specific predation risk. A common garden setup was used to examine the genetic and environmental components of the morphological variation. We found differences in head and jaw length and slight differences in body depth between the wild young‐of‐the‐year perch from Lake Ängersjön and Lake Fisksjön. The differences found between the wild fish from the two lakes were not maintained under common garden rearing. The observed morphological divergence between the wild young‐of‐the‐year perch from Lake Ängersjön and Lake Fisksjön seems to stem mainly from a plastic response to different environmental conditions in the two lakes. It is clear that the morphological traits are not influenced by direct reaction to the size‐specific risk of cannibalism, but probably stem from a combination of different environment characteristics, including resource and habitat use, and the density of other piscivores, such as pike.     

The effect of depth on the interaction between perch (Perca fluviatilis) and minnow (Phoxinus phoxinus)

We tested the hypothesis that small fishes avoid piscivores by increasing their use of shallow water areas, thereby reducing the capture success of the piscivore. Experiments were conducted in 600-1 aquaria, using perch (Perca fluviatilis) as the piscivore and minnow (Phoxinus phoxinus) as the prey. Two size classes of minnows, presented alone and together, were placed together with perch in aquaria with and without shallow water areas. The capture success of perch and use of depth by minnows changed as predicted by our hypothesis. In addition, we observed a decrease in the shoal size of minnows when perch were present. Perch selected large minnows over small minnows when both size classes were present, probably because large minnows were a more energetically profitable prey size.     

Abiotic and biotic correlates of yellow perch recruitment to age‐2 in southern Lake Michigan, 1984–2007

Abstract Seven abiotic and biotic variables were tested to determine whether they influence the recruitment to age two of yellow perch, Perca flavescens (Mitchill), in waters of southern Lake Michigan, USA. Recruitment was analysed using a Ricker stock‐recruitment relationship that accounts for density dependency. Significant model variables tested individually and ranked by total variance explained included the abundance of sexually mature yellow perch, alewife, Alosa pseudoharengus (Wilson) and spottail shiner, Notropis hudsonius (Clinton). Variables found unrelated to recruitment or generally less explanatory included the abundance of round goby, Neogobius melanostomus (Pallus), mean water temperature, variation in water temperature and water clarity. The best‐fitting multivariable model explained 69% of the recruitment variability and included abundances of sexually mature yellow perch, alewife and spottail shiner. These results suggest that yellow perch recruitment in southern Lake Michigan is regulated in part by biotic interactions with other species of the near‐shore community, including alewife and spottail shiners, in addition to the abundance of reproductively mature yellow perch.     

Patterns of foraging and distribution of bluegill sunfish in a Mississippi River backwater: influence of macrophytes and predation

Abstract— We studied the trophic interactions and spatial distributions of bluegills Lepomis macrochirus and largemouth bass Micropterus salmoides in a macrophyte bed in Lake Onalaska, a backwater lake in the upper Mississippi River. The diets of adult and age-0 bluegills were similar and changed seasonally probably in response to changes in life stages of macroinvertebrates (i. e. emergence of winged adults). Diets and diel patterns of abundance of bluegill suggest that age-O and adults were feeding in the vegetated, littoral zone. Predation by age-O largemouth bass appears to influence use of vegetated habitat by age-0 bluegills. In summer, when most age-O bluegills were vulnerable to predation by age-O largemouth bass, bluegill abundance was strongly correlated with vegetation biomass. In October and November, piscivory by age-O largemouth bass was limited by gape. Consequently, the relationship between the abundance of age-O bluegills and vegetation biomass was weakened because predation risk by age-O largemouth bass was reduced.     

A modelling approach to evaluate potential management actions designed to increase growth of white perch in a high‐density population

Abstract A deterministic, age‐structured population model was developed to simulate potential management scenarios designed to increase growth of individuals (quantified by maximum length) in a hypothetical population of white perch, Morone americana (Gmelin). Four scenarios were developed that included non‐selective mortality of adult white perch, increased mortality of age groups most influential on population growth, increased age‐0 mortality and inhibiting recruitment after spawning. The greatest increase in maximum length occurred with non‐selective adult mortality when population biomass was reduced by 97%; lesser increases in maximum length were achieved with the other management scenarios. Populations returned to their original state after control efforts ceased for each scenario.     

Impact of environmental and innate factors on the food habit of Chinese perch Siniperca chuatsi (Basilewsky) (Percichthyidae)

Laboratory and field investigations were conducted to study the food habit of Chinese perch Siniperca chuatsi (Basilewsky) from first feeding through adult stage. Only fish larvae were consumed by Chinese perch larvae (2–21 days from hatching), and the presence of zooplankton did not have any significant effect on their survival rate. The ability of Chinese perch to feed on zooplankton is clearly limited by some innate factor. Instead of gill rakers, Chinese perch larvae have well‐developed sharp teeth at the first feeding stage, and are well adapted to the piscivorous feeding habit unique to the larvae of Chinese perch, e.g. they bite and ingest the tails of other fish larvae. At the first feeding stage (2 days from hatching), daily rations were both very low, either in light or complete darkness. Although early‐staged Chinese perch larvae (7–17 days from hatching) could feed in complete darkness, their daily rations were always significantly higher in light than in complete darkness. Late‐staged Chinese perch larvae (21 days from hatching) were able to feed in complete darkness as well as in light, similar to the case of Chinese perch yearlings. Chinese perch yearlings (total length, 14–16 cm) consumed prey fish only and refused shrimp when visual cues were available (in light), but they consumed both prey when visual cues were not available (in complete darkness), suggesting that prey consumption by Chinese perch yearlings is affected by their sensory modality in predation. Both prey were found in the stomachs of similar‐sized Chinese perch (total length, 14–32 cm) from their natural habitat, suggesting that shrimp are consumed by Chinese perch at night. Prey selection of Chinese perch with a length >38 cm, which consumed only fish in the field, appears to be based upon prey size instead of prey type. These results suggest that although environmental factors (e.g. light intensity) affect prey detection by Chinese perch, this fish is anatomically and behaviourally predisposed to prey on live fish from first feeding. This makes it a difficult fish to cultivate using conventional feeds.