Salamander predation and vertical distributions of zooplankton

SUMMARY. 1. The impact of different predation regimes of zooplanktivorous larval tiger salamanders, Ambystoma tigrinum nebulosum , on the vertical temporal distributions of Daphnia pulex, Diaptomus nudus and larval Chaoborus flavicans was studied in limnetic enclosures in a pond in east-central Arizona. Zooplankton densities were monitored 1-2 h before sunset, at sunset, and 1-2 h after sunset at three depths (0-0.4, 1.0 and 2.0 m) in four salamander treatments (0, 20, 40 salamanders per enclosure, and unenclosed limnetic areas).
2. Densities and body size of Daphnia pulex were consistently greater in salamander-free enclosures than in other treatments. More Daphnia were generally nearer the surface (0-1.0 m) than along the bottom before and at sunset than after sunset in enclosures without salamanders. After sunset, Daphnia was by far most dense along the bottom. Densities were uniform among depths and sample times in treatments with salamanders. Salamanders moved throughout the water column. Frequencies of D. pulex spined morphs (with prominent dorsal crest) did not differ among salamander treatments.
3. Chaoborus densities were positively correlated with D. pulex distributions among depths and sample times in treatments. Spatio-temporal distributions of Chaoborus were influenced more by their daphnid prey than by risk of predation from larval salamanders.
4. Densities of Diaptomus nudus were highest in open water controls and lowest in enclosures without salamanders. Diaptomus was generally more dense between 0 and 1 m than at 2.0 m depths in all treatments, and vertical distributions were not dependent on salamander density.     

A comparison of the effects of prey and non-prey neighbours on foraging rates of Epischura nevadensis (Copepoda: Calanoida)

SUMMARY. 1. Adults of the calanoid copepod, Epischura nevadensis , aggregate in situ near the thermocline in Lake Tahoe, California-Nevada. together with adults of another species of calanoid copepod, Diaptomus tyrelli and juveniles of both species. With a series of laboratory predation and algal clearance trials, we show that foraging rates of adult E. nevadensis are determined not only by the density of co-occurring potential prey (small copepods), but also by the presence of co-occurring non-prey neighbours (large, adult copepods). These effects occur at densities and in zooplankton assemblages found naturally, emphasizing the ecological importance of neighbours other than prey on zooplankton feeding.
2. Neighbours are distinguished primarily by size. Although predation rates increase linearly with the densities of small copepods. both algal clearance and predation rates decrease in the presence of large copepods. We also show, with a field predation experiment using small enclosures, that adults are size selective within species and that Diaptomus are selected over conspecifics of the same size.
3. We hypothesize that by reducing foraging rates in the presence of large zooplankton. E. nevadensis avoids predators and reduces predation risk at the cost of reduced energy consumption.     

Seasonal dynamics of Skistodiaptomus pallidus Herrick and other zooplankton populations in Deer Lake, S.W. British Columbia

Seasonal environmental conditions and dynamics of a Skistodiaptomuspallidus population are given for a 2-year period in eutrophicDeer Lake, an urban waterbody near the northern limit of thiscalanoid's distribution. Life history characteristics of S.pallidus in the lake appear similar to those in more southerlyhabitats. It was perennial with breeding occurring between Marchand November; there were probably five generations producedper year. The annual cycles of Daphnia rosea, D. pulicaria.Diacyclops bicuspidatus thomasi, Bosmina longirostris, rotifersand Chaoborus flavicans are briefly described.     

Predation by chaoborids on typical and spined Daphnia pulex

SUMMARY. 1. In laboratory predation experiments, predation rates were determined for fourth-instar larvae of the chaoborids (Insecta: Diptera) Mochlonyx sp. and Chaoborus americanus preying on four juvenile instars and adult Daphnia pulex (clone SBL). In the case of Mochlonyx , predation was most intense on second-instar juvenile Daphnia while Chaoborus preyed equally on second and third instars.
2. Mochlonyx and Chaoborus induced an antipredator defence (neck-teeth) which was most frequently expressed in second-instar juvenile Daphnia . Expression of this antipredator phenotype was correlated with a reduced rate of Mochlonyx predation.     

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 .     

Hypolimnetic anoxia hampers top–down food-web manipulation in a eutrophic lake

SUMMARY 1. A biomanipulation experiment was carried out in a small (10 ha), but relatively deep (17 m) and highly eutrophic lake in northern Poland. The lake had been stocked in 1996, 1997 and 1998 with a variety of piscivorous fish (pike, catfish, trout and pikeperch), in order to reduce numbers of cyprinid planktivores.
2. Piscivore stocking was associated with a threefold decrease in the offshore fish density (night echosounding). Despite this reduction, the large planktonic cladoceran, Daphnia hyalina , remained scarce, whereas the density of small-sized zooplankton increased greatly.
3. The lack of demographic response in D. hyalina was probably due to the anoxia in the hypolimnetic refuge of this vertically migrating species. The anoxic hypolimnion, below 3–4 m depth, was inhabited day and night by numerous Chaoborus flavicans larvae.
4. Changes in zooplankton were associated with shifts in the taxonomic composition (from single-cell green algae to filamentous cyanobacteria), size structure (from nano- to net phytoplankton) and seasonal dynamics of phytoplankton, but not in the average biomass of planktonic algae. A clear-water phase, which was absent in the prestocking years, developed in spring, with Secchi depth reaching 2.5 m, a value which had never been recorded in the 20 years preceding the biomanipulation. In general, the lake's status was switched from hypertrophic to eutrophic.
5. Deteriorating food conditions, resulting from qualitative changes in the phytoplankton community, combined with predation pressure by the remaining fish and Chaoborus larvae were associated with the ultimate elimination of D. hyalina from the lake.     

Influence of food density on the effects of a Chaoborus-released chemical on Daphnia ambigua

SUMMARY 1. Daphnia ambigua was reared individually at four different food levels in water conditioned by a predator, Chaoborus flavicans. Daphnia ambigua developed spike-like helmets during the first four instars in each treatment with the Chaoborus-conditioned water, but only during the first instar in untreated water. The helmet development at instars 2–4 was probably induced by a chemical released from the Chaoborus .
2. The helmets decreased in size with decreasing food levels, but never ceased to form, even at a very low food concentration.
3. Daphnids cultured in the Chaoboras -conditioned water exhibited reduced growth, reproduction and survival rates at low food levels. These results might be induced by the energy loss associated with producing the helmets.
4. It is suggested that the predator releases a chemical, which reduces the tolerance of the cyclomorphic Daphnia to food deficiency, thus accentuating the summer decline in the Daphnia population caused by food shortage.     

Implications of Chaoborus pupation and ecdysis in cold water

1. Cold water acted differently to delay and lengthen the pupation period for the larvae of two species of the zooplankton predator Chaoborus (Diptera: Chaoboridae). During Chaoborus pupation, the zooplankton community is released from predation, while the dark-coloured Chaoborus pupae are more susceptible to their own predators.
2. Fourth instar larvae of Chaoborus americanus and C. trivittatus , collected from an oligotrophic lake, were reared individually at 5 °C in the dark. Chaoborus americanus was also reared at 9 and 12 °C under spring photoperiod conditions (L : D, 16 : 8 h). Individuals were observed through pupation to emergence (ecdysis) or death.
3.  Chaoborus americanus pupated at 5, 9 and 12 °C with substantial emergence only at 12 °C. In comparison, C. trivittatus emerged at 5 °C. Light was not a necessary cue for pupation and ecdysis, contrary to previous reports. Cold water delayed the onset and lengthened and increased the variability of the duration of pupation.
4. In Shirley Lake, C. americanus pupated in late June–early July while C. trivittatus pupated first in April and again in June–July.
5.  Chaoborus americanus pupae needed a temperature cue to complete ecdysis. The ecdysis temperature threshold helps to explain the difference in pupation timing, and the geographical distribution, of C. americanus and its relatively inflexible life history contrasted with C. trivittatus . Delayed predator pupation in years with low spring temperature can affect the community dynamics of the prey.     

Zooplankton Migration in Three Lakes of Western New York

We examined the diel vertical migration (DVM) of zooplankton in three lakes of western New York; Lakes Conesus, Lime, and McCargo. In all three lakes, the dipteran predator Chaoborus was a more dramatic migrator than any of the cladocerans or copepods. In contrast, another invertebrate predator, Leptodora, seemed to have the least vertical dispersal. In Conesus Lake, following the evening ascent and upper-water night-positioning of Chaoborus flavicans, Daphnia pulicaria expanded its own vertical range of dispersal thereby decreasing its degree of overlap with Chaoborus. The vertical distribution of Diaptomus sicilis was mostly below that of Daphnia pulicaria, thus reducing possible competition for food resources. Poor oxygen conditions in the lower waters of Lakes Lime and McCargo restricted all zooplankton (except Chaoborus) to an epilimnetic zone where the regions of niche overlap and predation were narrowed. Such annually-induced restrictions on vertical dispersal are probably common features of numerous stratified lakes with hypolimnetic reducing conditions. No fixed pattern of vertical dispersal or migration is likely in lakes where stratification and non-stratification follow each other seasonally. Vertical migration does provide some flexibility in niche separation, but the potential for chaotic behavior in interacting predators and prey may confound simple explanations of DVM.     

Zooplankton populations and the diets of three Chaoborus species (Diptera, Chaoboridae) in a tropical lake

SUMMARY. 1. Zooplankton populations in a small, natural, tropical lake are dominated by a few, small-sized taxa including the copepod Thermo- cyclops consimilis , the cladoceran Moina micrura and several rotifer species.
2. Moina micrura and adults of Thermocyclops consimilis undergo diel vertical migrations within the water column. Population densities of T. consimilis show marked intra-annual variations which may to some extent be related to variations in rainfall and to lunar periodicities In predator abundance.
3. The diets of Chaoborus lavae include other Chaoborus , Cladocera, Copepoda. Rotifera and the dinoflagellate Peridinium. The diets of late instars of the largest species, edulis , were dominated by Crustacea, while those of the two smaller species. C, ceratopogones and C. anomalus , were dominated by Rotifera and the dinoflagellate alga Peridinium , as were the diets of early instars of all species. Algae have not been previously reported to be a large component of the diet of Chaoborus populations in nature.
4. Some of the dietary differences among Chaoborus instars and species are related to the size of each prey species in relation to the mouth gape of each instar. However, there are also important differences in electivity among instars of different species of the same size.     

Studies on the biology of Chaoborus flavicans (Meigen) (Diptera: Chaoboridae) in a fish-free eutrophic pond,Japan

Population dynamics of Chaoborus flavicans larvae of various instars was studied from November 1986 to December 1987 in a eutrophic, fish-free pond, Japan. First and 2nd instar larvae were observed from late April to late October, indicating a reproductive period of about half a year. C. flavicans overwintered in the 4th instar larvae. In water column samples, total density of all instars was 680–23680 m^-2, and pupal density 0–2600 m^-2; larvae of the 1st, 2nd, and 3rd instars showed 5–6 density peaks in 1987, suggesting that 5–6 generations occur during a year (peaks of the 4th instar larvae were not clear, probably due to their longer development than those of younger instars). In sediment samples, no 1st and 2nd instar larvae were found, 3rd instar larvae were found occasionally but density of the 4th instar larvae was 280–18600 m^-2, and pupal density varied between 0–502 m^-2. Fouth instar larvae accumulated in sediment in the cold season and in the water column in the warm season; high temperature and low oxygen concentration were the most important factors limiting the distribution of larvae in the sediment in summer in the NIES pond. The dry weight of total C. flavicans larvae was 0.08–4.2 g m^-2 in sediment samples and 24–599 μg l^-1 (0.10–2.40 g m^-2) in water column samples. Comparisons of maximum densities in the NIES pond in different years and in waters of different trophic status show that density is generally higher in eutrophic than in oligotrophic habitats. This revised version was published online in July 2006 with corrections to the Cover Date.     

How well do laboratory experiments explain field patterns of zooplankton emergence?

1. We conducted a laboratory experiment to explore potential mechanisms driving variation in zooplankton emergence from diapausing eggs observed in Oneida Lake, NY, U.S.A. We hypothesized that variation in timing of ice-out (date of thawing of ice) between 1994 and 1995, which resulted in variation in photoperiod–temperature cues, contributed to the differences in the observed field patterns. Environmental chambers were used to establish weekly photoperiod–temperature combinations that reproduced natural conditions in Oneida Lake in 1994 and 1995. In addition, a third treatment ('dark') exposed eggs only to the changes in temperature. We recorded zooplankton emergence for 2.5 simulated ice-free seasons.
2. Nine cladoceran taxa were found to hatch, but only Daphnia pulicaria in large numbers. Hatching of D. pulicaria was recorded throughout the season in the two light treatments and sporadically in the dark treatment. The early ice-out treatment had the highest emergence, followed by the late ice-out and dark treatments. Among taxa, there was temporal segregation with five hatching in the early weeks of sampling and two taxa hatching during the middle weeks. Alona hatched late in the first year, but earlier in the second year.
3. We compared our laboratory results of D. pulicaria hatching with the field data obtained by Cáceres (1998) . Hatching was continuous in the laboratory, whereas a synchronous spring emergence was found in the field. However, in both the laboratory and the field, more D. pulicaria hatched under conditions reflecting ice-out occurring in March as opposed to April. Because differences in rates and timing of emergence can affect the population and community dynamics of pelagic systems, we suggest caution when applying laboratory results to field populations.     

Where to stay by night and day: Size-specific and seasonal differences in horizontal and vertical distribution of Chaoborus flavicans larvae

1. Data on the distributions of pelagic and benthic Chaoborus flavicans larvae were gathered in 1994 and tested for their agreement with the predator avoidance hypotheses. The development of all Chaoborus life stages, as well as the horizontal and vertical distribution in the four larval instars, was followed from May until October. We expected the largest larvae to dwell deeper by day, thus avoiding predation by visually foraging fish.
2. In agreement with this prediction body size increased with daytime depth, and this was true both between and within instars. The migration amplitude consequently increased with larval instar.
3. There was also evidence for horizontal migration, mainly in the third but also in the fourth instar.
4. Along a horizontal transect with increasing depth, locations with many benthic larvae had fewer pelagic larvae. Oxygen concentration was a good predictor of maximum benthic larval depth for most of the season but failed to predict their distribution in autumn.     

The seasonal dynamics and distribution of Chaoborus flavicans larvae in adjacent lake basins of different morphometry and degree of eutrophication

1.  Seasonal dynamics, spatial distribution and population size of the phantom midge Chaoborus flavicans in different parts of the eutrophic Lake Hiidenvesi (30.3 km²) were studied.
2.  Density of larvae was low in the shallow, most eutrophic parts of the lake, while the deep Kiihkelyksenselkä basin was inhabited by a dense population. In the deepest part of Kiihkelyksenselkä (33 m) density was 13 989 ± 3542 m^–2 in May, declined to 1102 ± 274 m^–2 in July and recovered to 7225 ± 1314 m^–2 by October. In spring and autumn the majority of larvae were benthic while, during high summer, few larvae were found in the sediment.
3.  Horizontal distribution fluctuated seasonally. On 3 June < 5% of the population inhabited areas shallower than 10 m. On 6 July the limnetic fraction was still restricted to regions deeper than 10 m, but 43% of benthic larvae were found between 6 and 10 m depths. In October both limnetic and benthic larvae were concentrated in areas deeper than 20 m.
4.  Within the lake, distribution was mainly regulated by stratification characteristics, degree of eutrophy being less important. The seasonal horizontal movements were probably induced by food shortage. Larvae could not meet their energetic demands in stratified areas and dispersed to shallower water, reducing predation risk by use of the benthic habitat.     

Regulation of rotifer densities by crustacean zooplankton in an oligotrophic montane lake in British Columbia

Summary To examine the relative demographic effects of predation and competition for food in rotifers during spring and summer in an oligotrophic lake, predator and competitor densities and food supplies were experimentally altered inside large enclosures. Abundances of rotifer species were positively correlated with experimental densities of fourth instar Chaoborus trivittatus larvae, a major crustacean predator in 1976. Experimental alteration of the densities of Daphnia rosea and Diaptomus leptopus and D. kenai in 1978 produced highly significant increases in rotifer biomass only under Daphnia removal, but not under copepod removal. Inorganic fertilizer additions to enclosures in 1978 and 1979 revealed minimal rotifer increases unless pulsed additions were large or Daphnia were also excluded. Large demographic reponses of rotifers to low fertilizer loadings in the absence of Daphnia confirmed the pre-eminence of competitive food limitation in producing rotifer scarcity in summer.     

Seasonal analysis of Daphnia pulicaria swimming behavior

Our study documents individual swimming behavior of Daphnia pulicaria over a yearly cycle in a temperate lake. We collected D. pulicaria, a common freshwater zooplankton, from Lake Mendota on 10 dates between July 1994 and June 1995 from two depths, 2 m and 10 m. The Daphnia were rushed to the laboratory and video-taped as they swam in lake water under lake-ambient temperature and light conditions. Five-second swimming tracks of individual Daphnia were filmed and digitized using a motion analysis system. We measured average turning angle, swimming speed and sinking rate for each track. D. pulicaria swimming behavior varied over the annual cycle. We found significant differences in turning angle between depths and among months. Sinking rate and swimming speed were significantly different among months but not depths. Sinking rate and swimming speed were not significantly correlated with water temperature. Our results were contrary to Stokes' Law predictions, in that D. pulicaria had the slowest sinking speed in June, not in the winter when water temperatures were lowest and viscosity was highest. Body length was significantly correlated with all three swimming variables. We also studied swimming behavior in clonal populations of D. pulicaria in different concentrations of the alga, Chlamydomonas reinhardtii. D. pulicaria did not change swimming speed, turning angle or sinking rate over a range of food concentrations. Finally, swimming behavior in a D. pulicaria clone, tested at two temperatures in the laboratory, confirmed the results from our seasonal study; Daphnia did not sink as predicted by changes in viscosity.     

Some effects of enclosure on the zooplankton in a small lake

Observations on the effect of enclosure on zooplankton by introducing two experimental tubes, each holding some 18 000 m³ of water, into a small lake, showed that a limnetic community could be maintained within tubes of this capacity throughout a period of 22 months. Two species, the copepod Diaptomus gracilis and the cladoceran Daphnia hyalina were dominant both in the open lake and in each tube, but Diaptomus gracilis was relatively more abundant in the tubes than in the lake, whereas Daphnia hyalina, especially in spring and autumn, was more abundant in the lake than in the tubes. Several scarce species became relatively more abundant in the tubes than in the lake. These changes are more probably associated with reduced predation, especially by larvae of Chaoborus, than with artificially induced changes in the phytoplankton, changes only poorly correlated with changes in the zooplankton.     

Cladoceran assemblages, seasonal succession and the importance of a hypolimnetic refuge

SUMMARY. 1. We investigate the importance of a refuge from fish predation to the abundance, species composition and seasonal succession of zooplankton. Thirty lakes representing a range of depths were sampled twice in summer for physical/chemical parameters and zooplankton community structure.
2. We define the refuge from centrarchid predators to be that space between the thermocline and the zone of anoxia. As lakes vary in rate of oxygen depletion from the hypolimnion. the refuge size and lake depth are independent: refuge size decreases during the summer period.
3. Lake depth and refuge size independently explain variation among lakes in zooplankton species composition, but seasonal community change within lakes is best predicted by loss of refuge size.
4. Refuge size also explains the substantial variation in the relative dominance of the two major daphnid species. Lakes possessing a large refuge are dominated by D. pulicaria ; those with a small refuge are dominated by the smaller, D. galeata mendotae . We suggest that lakes of intermediate refuge size, which are characterized by high species diversity, represent a more equitable balance of predation and competition.     

Accounting for Spatial Variability in the Design of Sampling Programmes for Chaoborus Larvae

We developed a model to estimate the number of net hauls (n)needed to quantify the mean abundance (m) of vertically migratingspecies of Chaoborus larvae (C. flavicans, C. punctipennis andC.trivittatus) in the entire pelagic zone of lakes at a specifiedlevel of precision (D = SE/m, where SE is the standard error).The model was n = 1.94m^–0.30D^–2. It was derivedby combining Taylor's power law with a model that estimatedsample size (i.e. number of net hauls) for a specified levelof precision. The model indicates that for a typical temperatelake Chaoborus population of 50 larvae m^–3, 15 net haulsare required to obtain a precision of 0.2. As the precisionof a single pelagic, vertical haul is only 0.77 at such typicalChaoborus densities, caution should be exercised in assessingthe role of Chaoborus in food webs using data based on the traditionalapproach to plankton sampling, i.e. one mid-lake station.     

Effects of algal food quality on fecundity and population growth rates of Daphnia

1. Food quality was at least as important as food quantity for both fecundity and population growth responses of the cladoceran Daphnia pulicaria fed the green alga Ankistrodesmus falcatus grown under N limitation, P limitation, or non-limited condition.
2. The fecundity of D. pulicaria was reduced under conditions of low food quality (low N or low P) compared with that for animals fed control non-limited algae regardless of ration size. The reduced fecundity of D. pulicaria fed P-limited food could be partially alleviated by increasing the ration (hence, compensation), but such was not the case for animals fed N-limited food.
3. Population growth rates of D. pulicaria ( r _max) were significantly reduced under conditions of low-quality food for both N-limited and P-limited algae. Population growth rates were unaffected by ration size, indicating no compensation.