Seasonal drying influences odonate adults and nymphs in wetlands in an urban mediterranean-climate landscape

Little is known about odonate ecology and phenology in warm mediterranean climates where most wetlands are intermittent. We identified variables associated with the presence, relative abundance, assemblage composition and body size of odonates in spring and summer, to understand the influence of intermittency and season. We hypothesized that size at eclosion would be smaller in seasonal wetlands in summer due to time stress caused by drying. Nymphs, exuviae and adults were sampled in spring and summer from 22 intermittent and perennial suburban wetlands in south-western Australia. Spatial and environmental variables within and between wetlands were measured and associated with adult and nymph distributions. Exuviae were collected to quantify size at eclosion. As in temperate perennial wetlands, wetland-scale variables (submerged, emergent and terrestrial vegetation, water temperature) were most strongly associated with assemblage composition of adults and nymphs, but landscape-scale variables (distance to nearest patch of native vegetation, distance to nearest large lake) were also associated with adult assemblages. Two abundant dragonfly (Orthetrum caledonicum Libellulidae, Hemicordulia tau Hemicorduliidae) and one damselfly (Xanthagrion erythroneurum Coenagrionidae) species emerged at smaller body sizes in summer than spring, but from both intermittent and perennial wetlands. Consequently, declining photoperiod and warmer summer temperatures, rather than wetland drying, probably caused reduced size at eclosion. Although the influence of vegetation, temperature and photoperiod on odonate assemblages appears similar in temperate and mediterranean climates, odonate phenology differs markedly. Fitness cost of emerging at a smaller adult size may be outweighed by the increased likelihood of successfully reaching emergence in drying waterbodies in summer. More field data on size at eclosion in warm climate regions, and laboratory experiments manipulating temperature and photoperiod, are needed to confirm the generality of patterns shown here.     

Bottom fauna and littoral vegetation fauna in Lake Maarsseveen

Summary The littoral vegetation starts its development only in June and reaches its highest expansion in autumn. By that time the organic debris from these plants starts to play an important role for detritivorous organisms. The quantitative development of the fauna in the littoral is highest in autumn and early winter. The bottom fauna in the hypolimnetic parts of the lake is less dependent of the season. The experiments with artificial plants have shown that most species are present when the littoral vegetation is absent. Owing to the absence of substrate in the form of vegetation with epiphyton as a major food resource the numbers in spring and early summer are low. The most productive invertebrate isDreissena polymorpha.     

Community size structure of zooplanktonic assemblages in three lakes on the upper River Paraná floodplain, PR-MS, Brazil

Abundance and body size of zooplanktonic organisms, testate amoebae, rotifers, cladocerans and copepods from the littoral and pelagic regions of three lakes were analyzed in February and August 2000. The lakes belong to three different river systems (Baía, Ivinhema and Paraná rivers) of the upper River Paraná floodplain. It was expected that average body size was different in space (regions of the lake and limnological characteristics) and time (summer and winter) because the variation of depth, pH, oxygen dissolved, chlorophyll-a and water temperature of each lake. Zooplankton community was represented by 119 species. Sorensen's coefficient showed that the three communities were similar. Larger organisms were found in the lakes' pelagic region, and seasonally larger individuals were registered in the winter and smaller individuals in the summer. The relationship between body size and density was slight, positively significant. The body size frequency distribution was bi-modal. ANOVA results showed a significant influence of the interaction of the littoral and pelagic regions, lakes, and seasons in the body size of zooplankton organisms. Spatial and temporal changes of the community size structure of zooplanktonic assemblages were related to the food resource (microbial-loop or herbivore chain), species habitat preference and life strategies (growth and reproduction).     

The influence of productivity and width of littoral zone on the trophic position of a large-bodied omnivore

Omnivory is common in many food webs. Omnivores in different habitats can potentially change their feeding behaviour and alter their trophic position and role according to habitat conditions. Here we examine the trophic level and diet of the omnivorous signal crayfish (Pacifastacus leniusculus) in gradients of trophic status and lake size, both of which have been previously suggested to affect trophic position of predators separately or combined as productive space. We found the trophic position of omnivorous crayfish to be positively correlated with lake trophic status, but found no evidence for any influence of lake size or productive space on crayfish trophic position. The higher trophic position of crayfish in eutrophic lakes was largely caused by a shift in crayfish diet and not by an increase in trophic links in basal parts of the food web. Hence, our results support the “productivity hypothesis,” suggesting that food chains can be longer in more productive systems. Furthermore, stable isotope data indicated that larger crayfish are more predatory than smaller crayfish in lakes with wider littoral zones. Wider littoral zones promoted the development of intrapopulation differences in trophic position whereas narrow littoral zones did not. Hence, differences in habitat quality between and within lakes seem to influence the trophic positions of omnivorous crayfish. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The diet and growth of roach (Rutilus rutilus (L.)) in Lake Vesijärvi and possible changes in the course of biomanipulation

Previous studies have suggested that the roach Rutilus rutilus (L.) stock of Lake Vesijärvi is one of the main factors delaying the recovery of the lake after sewage diversion. This study is concerned with the documentation of the diet of roach in the lake. In total, 531 roach were examined. Both in the pelagial and in the littoral the roach had mixed diets in May and in September—October. The importance of zooplankton decreased and the importance of benthos and plants increased with increasing size of roach. In July, in the pelagic zone all sizes of roach fed exclusively on zooplankton (Bosmina spp.), while in the littoral zooplankton had the highest volume proportions only in the smallest (<130 mm) roach. The frequent use of plant food and slow growth rate of large roach indicate a low availability of animal prey. As the fish densities decrease due to the mass removal taking place in the lake, the percentage of plant food in the diets of roach will probably decrease and the growth of roach will increase. Additionally, the tendency of the roach to migrate into the pelagic zone in early summer may be reduced, which would decrease their predation on the zooplankton.     

Movement of plankton through lake-stream systems

1. River plankton are often assumed to come from upstream lakes, but the factors controlling the movement of plankton between lakes and rivers into outflow streams are unclear. We tested the possibility that the physical structure of the littoral zone near the lake outlet (depth, presence of macrophytes) and diurnal differences in plankton composition at the lake surface influence the movement of plankton from the lake into the stream and determine their persistence downstream. 2. Zooplankton and phytoplankton biomass, community composition and mean body size were compared between two deep lakes without macrophytes at the lake edge and two shallow lakes with macrophytes at the lake edge. Samples were collected day and night on three dates, in the lake centre, in the littoral zone adjacent to the lake outlet, at the outlet and at two sites downstream in Algonquin Park, Ontario, Canada. 3. The morphology of lake edges clearly affects the movement of lake zooplankton into outlet streams. Outlets draining deeper littoral zones had higher zooplankton biomass than shallow littoral outlets (P < 0.0001), but these differences disappeared within 50 m downstream of the lake. There was no difference in mean zooplankton body size among lake outlets or between littoral and outlet samples. However, shallow littoral zones were dominated by cyclopoid copepods and deeper littoral zones were dominated by Bosmina longirostris. In contrast, phytoplankton biomass entering the outlet was similar to that found within the lake and did not vary with lake outlet morphology. These effects were consistent across several sampling weeks and were not affected by surface zooplankton biomass changes associated with diurnal vertical migration in the lake centre. 4. A comparison with published river zooplankton data suggests that zooplankton are rapidly eliminated from shallow outlet streams (≤1 m deep) but persist in most deeper outlet rivers (≥2 m deep). Because the depth of an outlet river determines downstream zooplankton community development, the contribution of lakes to river plankton communities may be influenced by the location of each lake within the drainage basin. These findings suggest that lake and outflow physical structure influences connection strength between spatially successive habitats.     

Ecology under lake ice

Winter conditions are rapidly changing in temperate ecosystems, particularly for those that experience periods of snow and ice cover. Relatively little is known of winter ecology in these systems, due to a historical research focus on summer ‘growing seasons’. We executed the first global quantitative synthesis on under‐ice lake ecology, including 36 abiotic and biotic variables from 42 research groups and 101 lakes, examining seasonal differences and connections as well as how seasonal differences vary with geophysical factors. Plankton were more abundant under ice than expected; mean winter values were 43.2% of summer values for chlorophyll a, 15.8% of summer phytoplankton biovolume and 25.3% of summer zooplankton density. Dissolved nitrogen concentrations were typically higher during winter, and these differences were exaggerated in smaller lakes. Lake size also influenced winter‐summer patterns for dissolved organic carbon (DOC), with higher winter DOC in smaller lakes. At coarse levels of taxonomic aggregation, phytoplankton and zooplankton community composition showed few systematic differences between seasons, although literature suggests that seasonal differences are frequently lake‐specific, species‐specific, or occur at the level of functional group. Within the subset of lakes that had longer time series, winter influenced the subsequent summer for some nutrient variables and zooplankton biomass.     

Feeding behaviour of major 0 + fish species in a shallow, eutrophic lake (Tjeukemeer, The Netherlands)

The forage base and the food selectivity of 0+ representatives of six abundant freshwater fish species were studied in a shallow, eutrophic Dutch lake. Most species relied on the zooplankton; the size-selective predation in early summer was directed to the smaller copepods and in late summer to larger cladocerans and copepods than concurrently present in the lake. Daphnia spp. and cyclopoid copepods were the main zooplankton taxa for smelt, perch and pikeperch. Energetically, the large cladoceran, Leptodora kindtii, was especially important for pikeperch. Bream and roach preyed upon smaller zooplankton than the other fish species. The influence of the zooplankton predation by abundant 0+ fish was clear from a small mean Daphnia size in September; this size is to be used as an indicator in fishery management. Neomysis integer, the most important macrofauna species, was consumed by perch, pikeperch and ruffe; pikeperch was most size-selective in this respect. The 0+ ruffe was à typically benthivorous fish. Only the 0+ pikeperch became piscivorous, especially in years when smelt was abundant.     

Intraspecific variability of the phenology and morphology of three protected dragonflies between natural and artificial habitats

Changes in phenology and in body size are two of the three main consequences of global warming on organisms. We investigated whether living in a warm artificial habitat would induce changes in the phenology and body size of dragonflies. We monitored in natura the emergence pattern of three protected and red-listed dragonfly species in three geographically close systems which differ in thermal profiles: a medium-sized river, one of its tributaries and an artificial lake fed by the water of the tributary. We also investigated the morphological variability of one of the species between the three systems. We showed an asynchrony of emergence for the three species, as well as morphological variability between the lake and the two rivers. Individuals from the lake emerged earlier and were smaller than those from the two rivers. These results are in agreement with a temperature-induced response hypothesis as the lake is warmer than the two rivers. Asynchrony of emergence between neighbouring populations triggers questions related to metapopulation functioning and about the fitness and the fate of the early-emerging individuals. Understanding the response of these species to local thermal conditions will help to improve population monitoring and conservation.     

The More the Merrier: Swarming as an Antipredator Strategy in the Mysid <Emphasis Type="Italic">Neomysis Integer</Emphasis>

The swarming behaviour of a Baltic littoral mysid shrimp, Neomysis integer, was studied both in the presence and absence of a predator (European perch, Perca fluviatilis L.). I performed two kinds of laboratory experiments. First, the swarming tendency of mysids and the effect of swarm size on swarm choice were studied. Second, the ingestion rate of mysids was measured when feeding alone versus in a swarm. The results indicate that N. integer actively join swarms. The avoidance of the perch by N. integer individuals was stronger when there was a swarm present. Larger swarms were preferred over smaller ones regardless of presence or absence of the predator. The overall feeding rate was similar when feeding alone and in swarm, but predator cues reduced feeding rate only when the mysids were feeding alone. This study demonstrates the capability of N. integer to assess predation risk and social context and alter their behaviour accordingly.     

Distribution and summer diapause of Canthocamptus staphylinus (Jurine) (Copepoda: Harpacticoida) in Lake Ladoga

The spatial distribution of Canthocamptus staphylinus (Jurine) in Lake Ladoga was examined. C. staphylinus is one of the most numerous species of harpacticoids in the lake. It can dwell in almost all littoral biotopes. In profundal biotopes strongly exposed to anthropogenic influence (pollution and eutrophication) only this species of harpacticoid was found. A peculiarity of C. staphylinus biology is that in summer individuals are in the resting stage as cysts. In summer active individuals were found in littoral habitats at a water temperature of 20 °C, but their number was small in comparison with cold periods of the year. In the littoral zone the relative number of individuals in cysts was clearly related to water temperature. Twelve degrees temperature appears to be critical for development of littoral populations of C. staphylinus.     

Effect of body size on swarming behavior and mating success of maleAnopheles freeborni (Diptera: Culicidae)

We examined whether body size affects the swarming behavior and mating success of male Anopheles freeborninear California rice fields. Swarms formed after dusk and persisted for approximately 30 min. The proportion of males in 33 swarms sampled n=6028 ranged from 100 to 92% but decreased over time (r=0.73, t=6.03, P<0.001).On average, swarming males (n=1058) were larger than males sampled from the resting population (n=735, H=35.6, P<0.0001),indicating that some males never swarm at all. Males swarming early were significantly smaller than those swarming during the peak (H=6.71, P=0.009)or final minutes of the swarm (H=4.86, P=0.002). Mated males returned to the swarm after mating, and larger males enjoyed greater mating success than did smaller ones (n=398, H=16.1, P=0.0005).     

Seasonal Vertical Migration and Aestivation of <Emphasis Type="Italic">Rhyacodrilus hiemalis</Emphasis> (Tubificidae,Clitellata) in the Sediment of Lake Biwa,Japan

The vertical distribution of the tubificid worm Rhyacodrilus hiemalis Ohtaka, the numerically dominant species of oligochaete in the littoral of Lake Biwa, was studied with special reference to seasonal vertical migration in the lake sediment. Monthly collections of lake sediment cores were made using PVC tubes. Core sections of sample sediments ranged from 76 to 117 cm. The vertical distribution of the worms showed no diurnal variation; therefore diel vertical migration was not evident. Seasonal downward migration started in April, and upward migration started in October. From December to March, almost all worms remained in the near-surface sediment layer (surface to 30 cm deep), while from July to September almost all worms remained deeper than 30 cm. However, few individuals migrated deeper than 90 cm. No discontinuous layers were found in grain size composition, water content, loss on ignition, particulate carbon, nitrogen or phosphorus. In deep sediment there was no free oxygen, as evidenced by negative ORP values. For 4 months in summer, R. hiemalis aestivated, probably utilizing anaerobic respiration. It appeared that R. hiemalis moved deeper in the sediment in response to sediment temperature, because sediment temperatures in the deep layers seemed to converge at around 20–21 °C in the summer months. The life history traits of seasonal vertical migration and summer aestivation perhaps arose as an adaptation to the climatic conditions accompanying the geographical origin of R. hiemalis, and they also serve to minimize predation risk during summer when most invertebrate predatory fishes are active.     

Ontogenetic habitat shifts by Galaxias gracilis (Galaxiidae) between the littoral and limnetic zones of Lake Kanono, New Zealand

Synopsis Diel and spatial differences in distribution were determined for the larvae, juveniles, and adults of Galaxias gracilis (Galaxiidae) in a New Zealand dune lake during summer months. Larvae (mostly 10–25 mm TL) and juveniles (25–40 mm TL) inhabited shallow (0–3 m) waters of the limnetic zone and fed predominantly on two limnetic zooplankton species; Bosmina meridionalis and a calanoid copepod. At about 40 mm TL, fish moved from the limnetic to the littoral zone and expanded dietary breadth from two to over seven main prey species, including five species of littoral invertebrates. After reaching a size of about 60 mm TL, most fish moved back offshore to the deeper waters (5–15 m) of the limnetic zone during the day, moving back to the littoral zone at night to feed on invertebrates. The selection of different intra-lacustrine habitats by the various size groups of G. gracilis, and the movements between them, are interpreted as adaptive responses to the interaction between ontogenetic changes in feeding requirements and predation risk.     

Carbon,phosphorus and nitrogen budgets of the littoral Equisetum belt in an oligotrophic lake

Stores and flows of carbon, phosphorus and nitrogen in a littoral Equisetum stand were studied in 1978–1980 in the oligotrophic, mesohumic lake Pääjärvi, southern Finland. The major carbon and nutrient stores were sediment and Equisetum. The seasonal cycle of the macrophyte vegetation had a profound influence on the whole littoral ecosystem. In spring, when only dead remains of Equisetum were present above ground, there were few differences in nutrient, chlorophyll a and zooplankton concentrations between the littoral and the open lake; phytoplankton and epiphytes were the major producers.In early June, when new shoots of Equisetum reached the water surface, water exchange between the littoral and the open lake started to diminish, and the characteristic features of a closed macrophyte zone gradually developed: by August the P, Chl a and zooplankton concentrations in the littoral were 5–10 times those in the open lake. From late June until autumn Equisetum was overwhelmingly dominant both in biomass and in production.The measured total primary production and respiration values indicated a high rate of internal cycling of carbon and nutrients. The daily P requirements of plant growth exceeded the total P stored in the water by a factor of 2–4, and also exceeded the release of nutrients in excretion. High N:P ratios in the water (total 10–64, inorganic 18–171) suggested that P was probably always the limiting nutrient.The P content of the annual production of Equisetum in Pääjärvi was 2.3% of the mean annual P load, and 5.3% of the mean total P storage in the water volume of the lake.     

Trading safety for food: evidence from gut contents in roach and bleak captured at different distances offshore from their daytime littoral refuge

1. Regular diel habitat shifts in roach were detected by hydro‐acoustics in five moderately eutrophic, stratifying (maximum depth 24–27 m) and approximately circular lakes (of surface area 15, 75, 125, 300 and 900 ha and diameters 250, 600, 1000, 1700 and 2600 m) in north‐eastern Poland in the years 1998–2000, when the lakes were free of smelt and other typical offshore planktivores, and their offshore areas were completely free of fish during the day. 2. The diel change in roach distribution was shown to assume a similar pattern in each lake: fish migrated from a daytime littoral refuge towards the centre of the lake at dusk, and returned to the littoral refuge at dawn. After sunset, fish gradually dispersed offshore until they covered the entire lake area in each of the three smaller lakes. In each of the two larger lakes, only small numbers of fish were seen in the central area at night, implying that the centre of the lake retained high food availability throughout the summer. 3. Inshore–offshore gradients in zooplankton prey density, body size, and numbers of eggs per clutch were weak or undetectable in the two smallest lakes, but strong and persistent in the three larger lakes, with Daphnia densities 5–30 times as high and body length 1.2–1.5 times as great in the central area as inshore. 4. The likely increase in the potential predation risk with distance from the littoral daytime refuge was found to be compensated by increased food gains in those fish which moved offshore at dusk to feed within a short time window, when light intensity was lower to make the risk reduced, but still high enough to see zooplankton prey. The benefit because of increased prey acquisition was greatest in the centre of the largest lake (at 1300 m from the shore), as revealed from gut inspections of roach and bleak trawl‐sampled at different distances from the edge of the reed belt, and seen as a gradual, order‐of‐magnitude increase in the volume of food in the foregut, The food volume against distance‐from‐shore regression was highly significant on each of the four sampling dates in the largest lake, in spite of the wide variability of food volume in individual fish.     

Horizontal distribution of crustaceans in Lake Glubokoe

A study has been made on the distribution of crustaceans along a profile of 6–9 stations from depths of less than 2 m to 20 m, in summer, autumn and early spring. The species composition, abundance, size-frequency distribution, biomass and diurnal dynamics were estimated. Many species abundant in the pelagic zone were found to be absent or scarce out to depths of 5 m. In the daytime, at depths out to 3 m, crustaceans greater than 1.0 mm were seldom found, while those 0.15–0.40 mm long were most abundant. At depths out to 5 m, not only was the total average size of crustaceans found to be smaller, but also that of individual species, including small ones: Ceriodaphnia and Chydorus. The biomass of crustaceans in shallow water and in the open lake differs in summer by more than 100 times, and in late autumn and early spring by more than 10 times. At the depth of 1 m, in none of the seasons did it exceed 0.026 mg l^-1. All the crustaceans except for nauplii and Chydorus considerably increased in numbers in the shallow water at night. The impoverishment of the littoral plankton in Lake Glubokoe is regarded as a specific case of the well-known phenomenon of ‘avoidance of the littoral’ by euplankton (Hutchinson, 1967; Kiselyov, 1980). It is pointed out that there is a possible relationship between diurnal horizontal migrations of plantivorous fish and the impoverishment of plankton near the shore. An attempt has been made to substantiate the assumption that the ‘avoidance of the littoral’ by pelagic crustaceans is due to high numbers of fish in shallow water.     

Accounting for littoral primary production by periphyton shifts a highly humic boreal lake towards net autotrophy

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Evidence for the retention of genetic variation in Erodium seed dormancy by variable rainfall

Summary The periodic occurrence of summer/early autumn precipitation in the California annual grassland can result in the formation of early and late emerging cohorts of Erodium botrys and E. brachycarpum. The occurrence of early rainfall and the timing of such rainfall are highly variable from year to year. A series of field watering experiments in 1980–81 were used to simulate early emergence conditions that would result from significant rainfall (1 cm) occurring in mid-July, late August, and mid-September. Net reproduction was used to estimate fitness differentials between Erodium cohorts emerging in response to a watering treatment (early emerging cohorts) and Erodium cohorts emerging with the onset of winter rains in mid-October (late emerging cohorts). Survival was lower and gross reproduction was higher among early emerging cohorts than late emerging cohorts. For both species, net reproduction of the early cohort was lower than that of the late cohort under the July watering treatment and higher than that of the late cohort under the August watering treatment.Early cohorts, formed in response to rainfall in mid-September, 1982, were also compared demographically to later cohorts emerging in October. Compared to late cohorts, net reproduction, gross reproduction and survival were higher for the early cohorts.Common garden experiments indicate that differences in the duration of seed dormancy between the progenies of early and late emerging plants reflect a significant genetic component. Progency produced by early cohorts of E. brachycarpum from all three watering treatments possessed more extended seed dormancy than progeny of late cohorts. In E. botrys, progeny from early cohorts emerging in response to the July watering treatment were also more dormant than late progeny. In contrast, early cohorts of E. botrys emerging in response to the September watering treatment produced seed less dormant than seed produced by late cohorts. When combined with demographic data, indicating that fitness differentials between early and late cohorts varied with changes in the date of early emergence, genetic results suggest that year to year variation in early rainfall may act to retain genetic variation in the duration of seed dormancy.     

Effects of habitat and food resources on morphology and ontogenetic growth trajectories in perch

Studies on resource polymorphism have mainly been considered at the end stage of ontogeny, whereas many species undergo diet changes as they grow. We conducted a field survey to analyze the role of adaptive variation during ontogeny in Eurasian perch (Perca fluviatilis). We caught perch from the littoral and pelagic zones of a lake to investigate whether perch differ in morphology and diet between these habitats. We also investigated whether there were any differences in morphological trajectories during the ontogeny of perch collected from the two habitats. We found that perch caught in the littoral habitat, independently of size, had a deeper body, larger head and mouth and longer fins than perch caught in the pelagic zone. Macroinvertebrates and fish dominated the diet of littoral perch, whereas the diet of the pelagic perch consisted mainly of zooplankton and to some extent fish. Independently of size, the more streamlined individuals had a larger proportion of zooplankton and a smaller proportion of macroinvertebrates in their diet than the deeper-bodied individuals, indicating a relation between diet and morphology. Some morphological characters followed different ontogenetic trajectories in the two habitats; e.g. the changes to a deeper body and a larger head were faster in the littoral than in the pelagic perch. The relationship between the length of perch and the size of the mouth and fins also differed between perch from the two habitats, where the increase in the length of the pelvic fin and the area of the mouth increased faster with size in the littoral perch. Our findings show that variation in morphology between habitats differs during ontogeny in a way that corresponds to functional expectations for fish species that occupy these habitats.