Invaders Ctenophores Mnemiopsis leidyi (A. Agassiz) and Beroe ovata Mayer 1912, and Their Influence on the Pelagic Ecosystem of Northeastern Black Sea

We analyzed the state of pelagic ecosystem of northeastern Black Sea affected by invasion of a new settler Beroe ovata in 1999–2001. Development of B. ovata considerably decreased the population of another invader, Mnemiopsis leidyi, that deformed the Black Sea ecosystem for over a decade. Reduction of M. leidyi population limited its influence on the ecosystem and, consequently, we observed recovering of the main components of the Black Sea pelagic ecosystem—zooplankton and fish as well as their eggs and larvae. The relationships between annual and seasonal variability of the population development and size of the both invaders M. leidyi and B. ovata are discussed.     

Temporal variation in abundance of the egg predator Carcinonemertes epialti (Nemertea) and its effect on egg mortality of its host,the shore crab,Hemigrapsus oregonensis

An outbreak of the nemertean, Carcinonemertes epialti, was observed on Hemigrapsus oregonensis during October, 1982 at Campbell Cove, Bodega Harbor, California. Mean worm intensity (296 worms/crab) was the highest recorded for this nemertean egg predator on H. oregonensis. During the outbreak, male crabs were found to harbor more worms than both non-ovigerous and ovigerous females. Crab egg mortality was substantial; 83% of the ovigerous females experienced 75–100% brood mortality. The seasonal peak in worm abundance coincided with the seasonal low in crab reproduction at this locality. A method for estimating the impact of C. epialti on H. oregonensis natality was developed using crab size and fecundity, and worm prevalence and intensity. For a non-outbreak sampling period, a mean of 5.6% egg mortality was experienced by infested crabs for the period selected. Thus, brood mortality during the outbreak was much greater than that experienced at non-outbreak periods. Heavy fishing pressure on some commercially important crab species has been suggested as a possible factor inducing worm outbreaks and facilitating their continued persistence. These observations suggest that fisheries are not necessarily responsible for the outbreaks of nemerteans on commercially important crab species. However, fishing pressure may still be a sufficient condition to promote nemertean outbreaks.     

Timing and size of blooms of the ctenophore Mnemiopsis leidyi in relation to temperature in Narragansett Bay, RI

The ctenophore Mnemiopsis leidyi is at the northern extreme of its geographic range in Narragansett Bay, an estuary on the northeast coast of the United States. Blooms have typically been observed in late summer and fall according to records from 1950 to 1979. We document an expansion of the seasonal range of this important planktonic predator to include springtime blooms during the 1980s and 1990s. This shift to an earlier seasonal maximum is associated with increasing water temperature in Narragansett Bay. Temperatures in spring have risen, on average, 2 °C from 1950 to 1999 with warm years being associated with the positive phase of the North Atlantic Oscillation. During 1999, M. leidyi appeared earlier in spring and was more abundant than during any previous year for which records are available. Changes in the seasonal pattern and abundance of this predator are likely to have important effects on planktonic ecosystem dynamics of Narragansett Bay. These include reduction of zooplankton abundance in spring followed by increases in size and frequency of summer phytoplankton blooms. Earlier blooms of M. leidyi may also reduce survival of eggs and larvae of fish because, as in 1999, they coincide with the period of peak spawning.     

Spatio-temporal overlap of the alien invasive ctenophore <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> and ichthyoplankton in the Bornholm Basin (Baltic Sea)

In 2007 the alien invasive ctenophore Mnemiopsis leidyi A. Agassiz 1865 was recorded for the first time in the Bornholm Basin, an area which serves as important spawning ground for Baltic fish stocks. Since M. leidyi is capable of preying upon early life stages of fish and further might act as food competitor for fish larvae, it is of major concern to investigate the potential threat that this non-indigenous species poses to the pelagic ecosystem of the Baltic Sea. The present study investigates the temporal and spatial overlap of M. leidyi with eggs and larvae of Baltic cod (Gadus morhua L.) and sprat (Sprattus sprattus L.) in order to assess the potential impact of this new invader on two of the most important Baltic fish stocks. Results show variable inter-seasonal distribution and overlap dynamics and thus different seasonal threat-scenarios for the early life stages of cod and sprat. The spatial overlap between M. leidyi and ichthyoplankton was low for most of the period observed, and we conclude that M. leidyi presently does not have a strong impact. However, we detected situations with high overlaps, e.g. for sprat larvae and cod eggs in spring. As the population dynamics of M. leidyi in the central Baltic are not yet fully understood, a future population explosion of the alien ctenophore with possible effects on fish recruitment cannot be ruled out. Furthermore, a possible shift in peak spawning of cod to the early season, when ctenophore abundances were relatively high, might increase the impact of M. leidyi on cod.     

Modelling potential habitat of the invasive ctenophore <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> in Aegean Sea

The invasive ctenophore Mnemiopsis leidyi was accidentally introduced into the Black Sea in the early 1980s and it was first sighted in the Aegean Sea (Eastern Mediterranean) in the early 1990s. This article presents a first attempt to develop a predictive spatial model based on M. leidyi presence data and satellite environmental data from the Aegean Sea during early summer, in order to identify those areas in the Greek Seas and the entire Mediterranean basin that could serve as potential habitat for the species. Generalized additive models (GAM) were applied. The final GAM model indicated higher probability of finding M. leidyi present in depths of 65–135 m and sea surface temperature values of 21–25°C. Furthermore, the significant interaction between photosynthetically active radiation (PAR) and sea level anomaly (SLA) indicated a higher probability of M. leidyi presence in low values of PAR and SLA. In the next step, the final GAM was applied in a prediction grid of mean monthly satellite values for June 2004–2006 in order to estimate probability of M. leidyi presence in the Hellenic Seas and the whole Mediterranean basin at a GIS resolution of 4 km. In the Aegean Sea, species potential habitat included areas influenced by the Black Sea Water (e.g. Thracian Sea, Limnos-Imvros plateau), gulfs that are affected by river runoffs, such as the Thermaikos, Strymonikos and Patraikos gulfs, or areas with strong anthropogenic influence such as the Saronikos gulf. Areas with the same environmental conditions as those in Aegean Sea have been indicated in certain spots of the Levantine Sea as well as in coastal waters of Egypt and Libya, although their spatial extent varied largely among years examined. However, the occurrence of conditions that are linked to high probability of M. leidyi presence does not necessarily mean that these areas can support successful reproduction, high population or bloom levels, since these depend on a combination of temperature, salinity, food availability and the abundance of predators. Guest editor: V. D. Valavanis Essential Fish Habitat Mapping in the Mediterranean     

Colonization of the Mediterranean basin by the vector biting midge species Culicoides imicola: an old story

Understanding the demographic history and genetic make‐up of colonizing species is critical for inferring population sources and colonization routes. This is of main interest for designing accurate control measures in areas newly colonized by vector species of economically important pathogens. The biting midge Culicoides imicola is a major vector of orbiviruses to livestock. Historically, the distribution of this species was limited to the Afrotropical region. Entomological surveys first revealed the presence of C. imicola in the south of the Mediterranean basin by the 1970s. Following recurrent reports of massive bluetongue outbreaks since the 1990s, the presence of the species was confirmed in northern areas. In this study, we addressed the chronology and processes of C. imicola colonization in the Mediterranean basin. We characterized the genetic structure of its populations across Mediterranean and African regions using both mitochondrial and nuclear markers, and combined phylogeographical analyses with population genetics and approximate Bayesian computation. We found a west/east genetic differentiation between populations, occurring both within Africa and within the Mediterranean basin. We demonstrated that three of these groups had experienced demographic expansions in the Pleistocene, probably because of climate changes during this period. Finally, we showed that C. imicola could have colonized the Mediterranean basin in the Late Pleistocene or Early Holocene through a single event of introduction; however, we cannot exclude the hypothesis involving two routes of colonization. Thus, the recent bluetongue outbreaks are not linked to C. imicola colonization event, but rather to biological changes in the vector or the virus.     

Potential pathways of invasion and dispersal of <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> A. Agassiz 1865 in the Baltic Sea

The rapid spread of Mnemiopsis leidyi across the entire Baltic Sea after its first observation in 2006 gave rise to the question of its invasion pathway and the possible vector of its transport. To investigate pathways of M. leidyi invasion, the years 2005–2008 have been simulated by a three-dimensional coupled sea ice-ocean model of the Baltic Sea. In addition, a Lagrangian particle-tracking model has been utilized to test possible transport routes of this invader for 2006/2007. Based on the model, we exclude advection from the Kattegat as the main area of origin of M. leidyi and further spreading through the entire Baltic Sea. To explain the dispersion of M. leidyi in 2007 an earlier invasion already in 2005 is most probable. Alternatively, an invasion originating from main harbors with high ship traffic could also be a potential pathway. Drift simulations with drifter release in the main harbors are in good agreement with the observed distribution pattern of M. leidyi.     

Predator-induced vertical behavior of a ctenophore

Although many studies have focused on Mnemiopsis leidyi predation, little is known about the role of this ctenophore as prey when abundant in native and invaded pelagic systems. We examined the response of the ctenophore M. leidyi to the predatory ctenophore Beroe ovata in an experiment in which the two species could potentially sense each other while being physically separated. On average, M. leidyi responded to the predator’s presence by increasing variability in swimming speeds and by lowering their vertical distribution. Such behavior may help explain field records of vertical migration, as well as stratified and near-bottom distributions of M. leidyi.     

Ecology of Naididae (Oligochaeta) from an alkaline bog stream: life history patterns and community structure

Population and community dynamics of naidid oligochaetes were studied from June 1982–December 1983 in a sluggish, alkaline bog stream situated within Cedarburg Bog, Ozaukee County, Wisconsin, USA. Temporal differences in periods of peak abundance were observed for five species studied in detail: Chaetogaster diaphanus (mid-August and September), Chaetogaster diastrophus (mid-May), Dero digitata (mid-September), Dero nivea (mid-October), and Pristina leidyi (mid-October). Several correlations of abiotic and biotic parameters to density and percentage of naidids reproducing asexually were calculated. Mean doubling times (days) for field populations were 12.1, 22.7, 62.4, 19.6, 27.7 for C. diaphanus, C. diastrophus, D. digitata, D. nivea, and P. leidyi, respectively. Asexual reproduction by paratomy was the principal method of population increase. Sexual specimens were observed for Stylaria lacustris during June and October–December, D. digitata in September, C. diastrophus and Nais simplex during October, and C. diaphanus in October–November. Generally, a high proportion of the respective population was sexually mature when this type of reproduction occurred. Naididae community diversity index values using the Shannon-Weaver index ranged from 0.19–3.86.     

Blooms of the invasive ctenophore, <Emphasis Type="Italic">Mnemiopsis</Emphasis> <Emphasis Type="Italic">leidyi</Emphasis>, span the Mediterranean Sea in 2009

Blooms of the invasive ctenophore, Mnemiopsis leidyi, occurred in 2009 along the Mediterranean Sea coasts of Spain and Israel. This voracious zooplanktivore spread throughout the Black Sea basin after its introduction in the early 1980s, throughout northern European coastal waters, and now occurs throughout the Mediterranean Sea. M. leidyi occurred throughout the summer along the entire Catalan Spanish and Israeli coasts in 2009. Those locations had high temperatures (18–26°C) and salinities (37–38) during the blooms. The patterns of abundance of large jellyfish along the Catalan coast were unusual in 2009, with low numbers during July, August, and September when ctenophores were abundant. Small populations of those potential predators and food competitors of M. leidyi could have contributed to the ctenophore bloom. The identity of the ctenophores from Spain and Israel was confirmed as M. leidyi by molecular analysis based on DNA sequencing of the nuclear internal transcribed spacer (ITS) regions. This is the first molecular confirmation of M. leidyi in the Mediterranean Sea. Most ctenophores had an ITS genotype previously found in M. leidyi from other invaded regions (the Black, Azov, and Mediterranean seas), as well as native regions in the United States, suggesting common ancestry. Based on the circulation patterns of Mediterranean surface waters and shipping activities, we conclude that the spread of M. leidyi in the Mediterranean probably resulted from re-introductions by ballast water transport and subsequent distribution by currents. We also conclude that the near-simultaneous blooms in opposite ends of both the Mediterranean basins indicate that M. leidyi is resident around the Mediterranean. We discuss environmental conditions, food, and predators of M. leidyi in both regions that would influence the future effects of this voracious consumer on the pelagic food web of the Mediterranean Sea.     

The Impact of Annual and Seasonal Rainfall Patterns on Growth and Phenology of Emergent Tree Species in Southeastern Amazonia,Brazil

Understanding tree growth in response to rainfall distribution is critical to predicting forest and species population responses to climate change. We investigated inter‐annual and seasonal variation in stem diameter by three emergent tree species in a seasonally dry tropical forest in southeast Pará, Brazil. Annual diameter growth rates by Swietenia macrophylla demonstrated strong positive correlation with annual rainfall totals during 1997–2009; Hymenaea courbaril growth rates demonstrated weak positive correlation, whereas Parkia pendula exhibited weak negative correlation. For both Swietenia and Hymenaea, annual diameter growth rates correlated positively and significantly with rainfall totals during the first 6 mo of the growing year (July to December). Vernier dendrometer bands monitored at 4‐wk intervals during 3–5 yr confirmed strong seasonal effects on stem diameter expansion. Individuals of all three species expanded in unison during wet season months and were static or even contracted during dry season months. Stems of the deciduous Swietenia contracted as crowns were shed during the early dry season, expanded slightly as new crowns were flushed, and then contracted further during 3–5 wk flowering periods in the late dry season by newly mature crowns. The three species’ physiographic distribution patterns at the study site may partially underlie observed differences in annual and seasonal growth. With most global circulation models predicting conditions becoming gradually drier in southeast Amazonia over the coming decades, species such as Swietenia that perform best on the ‘wet end’ of current conditions may experience reduced growth rates. However, population viability will not necessarily be threatened if life history and ecophysiological responses to changing conditions are compensatory.     

A long term study of small mammal populations in a Brazilian agricultural landscape

Long-term monitoring of small mammal populations is very important to understand the variations in temporal abundance on a large time scale, which are related to ecological, economic and epidemiological phenomena. The aim of this study is to monitor the populations of the marsupials Didelphis aurita and Philander frenatus and the rodents Nectomys squamipes, Akodon cursor and Oligorysomys nigripes in a locality of typical Brazilian rural landscape, Sumidouro Municipality, Rio de Janeiro State, Brazil. A mark-recapture study was conducted during five years. We analyzed the population dynamics, the reproduction and age structure of these species. Both marsupials presented higher population sizes in the end of wet period and beginning of the dry period, which can be explained by the seasonal reproduction which begins in the middle of the dry period and ends in the last months of the wet period. N. squamipes reproduced throughout the year but mostly during rainy periods, due to the close association of this rodent to resources found in the water. Higher survivorship and recruitment rates were in the end of the wet season. The rodent A. cursor had an opportunist reproduction, resulting in high turnover rates. Survivorship increased with the effects of the dry periods. O. nigripes showed a clear annual pattern of population cycle with peaks during the dry season. The rodents did not show potential to present outbreaks and become agricultural pests. The annual population cycles of O. nigripes and the unique peak of A. cursor population during five years highlight attention to their importance as wild reservoirs of the hantavirus disease. Their ecological characteristics associated to their opportunistic behavior make these species prone to be good reservoirs of zoonoses.     

The benthic invertebrates of the Salton Sea: distribution and seasonal dynamics

The Salton Sea, California's largest inland water body, is an athalassic saline lake with an invertebrate fauna dominated by marine species. The distribution and seasonal dynamics of the benthic macroinvertebrate populations of the Salton Sea were investigated during 1999 in the first survey of the benthos since 1956. Invertebrates were sampled from sediments at depths of 2–12 m, shallow water rocky substrates, and littoral barnacle shell substrates. The macroinvertebrates of the Salton Sea consist of a few invasive, euryhaline species, several of which thrive on different substrates. The principal infaunal organisms are the polychaetes Neanthes succinea Frey & Leuckart and Streblospio benedicti Webster, and the oligochaetes Thalassodrilides gurwitschi Cook, T. belli Hrabe, and an enchytraeid. All but Neanthes are new records for the Sea. Benthic crustacean species are the amphipods Gammarus mucronatus Say, Corophium louisianum Shoemaker, and the barnacle Balanus amphitrite Darwin. Neanthes succinea is the dominant infaunal species on the Sea bottom at depths of 2–12 m. Area-weighted estimates of N. succinea standing stock in September and November 1999 were two orders of magnitude lower than biomass estimated in the same months in 1956. During 1999, population density varied spatially and temporally. Abundance declined greatly in offshore sediments at depths >2 m during spring and summer due to decreasing oxygen levels at the sediment surface, eventually resulting in the absence of Neanthes from all offshore sites >2 m between July and November. In contrast, on shoreline rocky substrate, Neanthes persisted year round, and biomass density increased nearly one order of magnitude between January and November. The rocky shoreline had the highest numbers of invertebrates per unit area, exceeding those reported by other published sources for Neanthes, Gammarus mucronatus, Corophium louisianum, and Balanus amphitrite in marine coastal habitats. The rocky shoreline habitat is highly productive, and is an important refuge during periods of seasonal anoxia for Neanthes and for the other invertebrates that also serve as prey for fish and birds.     

Changes in biodiversity of phytoplankton,zooplankton, fishes and macrobenthos in the Southern Caspian Sea after the invasion of the ctenophore <Emphasis Type="Italic">Mnemiopsis Leidyi</Emphasis>

Monitoring for 6 years (2001–2006) showed that the population explosion of the alien ctenophore Mnemiopsis leidyi in the southern Caspian Sea coincided with a decline in the abundance and species number of mesozooplankton. While this decline appeared to have reduced the nourishment of sprat (also known as kilka), it seemed to have affected phytoplankton favorably mainly due to the decrease in grazing pressure. During 2001–2002, when M. leidyi abundance and biomass were at their highest levels, abundance of dinoflagellates and cyanophytes exceeded that of diatoms. Before the invasion (1996) and in some years after the invasion (2003, 2004 and 2006) diatom abundance was higher than the abundance of other groups. In September 2005, an unprecedented bloom of the toxic cyanophyte Nodularia sp. was observed in the southern Caspian Sea. Disappearance of edible zooplankton such as Eurytemora spp. was among the first changes observed after the expansion of M. leidyi in the area. Some changes in the macrobenthic fauna were also conspicuous after the increase of this ctenophore. While the biomass of some deposit feeders, such as the polychaete Nereis diversicolor and oligochaete species increased, benthic crustaceans decreased sharply in abundance during 2001–2003 and completely disappeared during 2004–2006. Iranian catches of kilka, the most abundant and widespread zooplanktivorous fish, decreased significantly in the southern Caspian Sea after 1999. Iranian landings of kilka dropped ~70% from 69,070 ± 20,270 t during 1995–2000 to 23,430 ± 12,240 t during 2001–2006, resulting in a loss of at least 125 million US dollars to the economy. There were also changes in the total catches of large predators such as the kutum and mullet, which mainly feed on kilka, between 1991 and 2006.     

Spreading and physico-biological reproduction limitations of the invasive American comb jelly Mnemiopsis leidyi in the Baltic Sea

Hydrodynamic drift modeling was used to investigate the potential dispersion of Mnemiopsis leidyi from the Bornholm Basin in the Baltic Sea where it has been observed since 2007 further to the east and north. In the brackish surface layer dispersion is mainly driven by wind, while within the halocline dispersion is mainly controlled by the baroclinic flow field and bottom topography. Model runs showed that the natural spreading via deep water currents from the Bornholm Basin towards north and east is limited by topographic features and low advection velocities. Based on the information on ranges of salinity and temperature, which limit survival and reproduction of this ctenophore within the Baltic Sea, areas have been identified where the American comb jelly, M. leidyi could potentially survive and reproduce. While, we could show that M. leidyi might survive in vast areas of the northern Baltic Sea its reproduction is prevented by low salinity (<10 psu) and temperature (<12°C). Thus, due to the combined effect of low salinity and temperature, it is not probable that M. leidyi could establish permanent populations in the central or northern Baltic Sea. However, it seems that in the southern parts of the Baltic Sea environmental conditions are suitable for a successful reproduction of M. leidyi.     

Vertical distribution of pelagial zooplankton in a middle-sized dimictic valley reservoir

Our observations indicate the vertical distribution of zooplankton and its seasonal changes in Dubník II reservoir (Slovakia) are determined mainly by the thermal regime of the reservoir, by transparency, and by fish and invertebrate predation. During periods of circulation, zooplankton vertical distribution in the whole water column was more homogeneous, whilst during summer temperature stratification zooplankton concentrated in the epilimnion — rotifers in higher layers than crustaceans. During summer stagnation a steep thermal gradient occurred at the boundary of the epi-and hypolimnion and low temperature and low dissolved oxygen in hypolimnion offered a refuge for Chaoborus flavicans larvae against fish, enabling coexistence of vertebrate and invertebrate predation. This evidence supports our previous findings concerning dominance of rotifers in zooplankton and representation of crustaceans by small-bodied species in the study reservoir. Steep thermal gradient and the presence of Chaoborus larvae caused very low zooplankton abundance in the lower part of the water column and a reduction of cladocerans refuges against fish to layers of thermocline or closely under thermocline where Daphnia cucullata and Daphnia parvula were found. Our previous assumptions about the high density of zooplanktivorous fish in Dubník II reservoir are supported by the fact that these small cladocerans are represented by smaller individuals in the upper layers and bigger individuals in deeper layers.     

Population rate of change in the leaf-eared mouse: The role of density-dependence,seasonality and rainfall

We analysed statistically the influence of density-dependent regulation, seasonality and precipitation on the realized population rate of change of the Neotropical rodent Phyllotis darwini (Waterhouse 1837) at an intra-annual time scale. We used four years of continuous live trapping at a semiarid locality of north central Chile. Results showed that density-dependence, seasonal effects and precipitation were important factors influencing population growth rates in this species. An empirical population model including a sine function for seasonal effects, a linear form for density-dependence, and precipitation was fitted to the full data set and to the data set with the first year removed (after an outbreak). The empirical model explained 33% and 48% of the variance in population growth. The natural rate of population increase, estimated from the empirical model, was r_max = 2.51 or 5.06 years^?1. These estimates indicate a great potential for population increase and may explain the capability of this species to undergo large irruptions. We propose that merging empirical and theoretical modelling with field research is the most promising avenue to understand the outbreaks experienced by some rodent species in western South America.     

Prorocentrum lima (Dinophyceae) in northeastern USA coastal waters: II: Toxin load in the epibiota and in shellfish

The seasonal variation in diarrhetic shellfish poisoning (DSP)-type toxins was followed in the epibiotic community and in shellfish between 41° and 44°N in coastal waters of the northwest Atlantic during a 2-year period. Low levels of okadaic-acid equivalents were detected at all stations in the <90 μm fraction of the collected epibiota as measured by the protein phosphatase inhibition assay, but only 3.5% of the samples had values greater than 100 ng (g dry weight of epibiota)⁻¹. No seasonal pattern could be detected due to differences in intensity, duration and timing of toxin content in the epibiota between the 2 years and between stations. Nevertheless, the concentration of DSP-type toxins in the epibiota correlated weakly but significantly with the abundance of Prorocentrum lima, when data from all stations were considered. A very limited toxin uptake by shellfish was measured at only one station in October and November 2001 and in June and July 2002 at times of maximum cell concentration of P. lima in the epibiota. Toxin levels in shellfish remained well below regulatory limits that would have required quarantine or bans on harvesting. Results from our 2-year survey suggest that, at this time, the threat of DSP events appears minimal. However, the presence of a known toxin producer and its demonstrated ingestion by shellfish would argue for further studies to better understand conditions leading to DSP outbreaks generated by an epiphytic dinoflagellate.     

Contrasting migration behaviour of Daphnia pulicaria and D. galeata x hyalina, in avoidance of predation by 0+perch (Perca fluviatilis)

In spring and early summer, a small population of the large-bodied Daphnia pulicaria coexists with a much larger population of the medium-sized hybrid Daphnia galeata × hyalina in the epilimnion of Lake Maarsseveen (The Netherlands). When large shoals of juvenile perch (Perca fluviatilis) appear in the open water, both species start to migrate vertically. Since D. pulicaria has a larger body-size than D. galeata × hyalina, and is therefore competitive dominant over the hybrid, it is unlikely that both species interact via their common food resource, but they react both to fish predation. However, since they differ in size, and therefore in vulnerability for fish predation, both species adopt different strategies. The smaller bodied, and less vulnerable D. galeata × hyalina exhibits diel vertical migration ascending to the surface at dusk, and staying there during the night. In this way, benefiting from the higher temperatures of the surface layers. In contrast, the large-bodied, and more vulnerable D. pulicaria selects the deep cold hypolimnion water layers as refuge against fish predation. In this way it benefits from a safe habitat, free from fish predators, but on the other hand suffers from low water temperatures, which decrease its fitness. It is likely that the relatively higher temperature in the upper water layers serves as a proximate factor for the downward migration of D. pulicaria.     

Food limitation and seasonal population declines of rotifers

Summary Keratella cochlearis exhibited consistent seasonal abundance patterns during a four-year study in Little Rock Lake, Wisconsin, U.S.A. In each year, spring population peaks were followed by strong summer reductions. Concomitant with population declines, there were reductions in rotifer egg ratios. Keratella taurocephala abundance patterns were similar to those of K. cochlearis during 1984 and 1985, but not in 1986 and 1987, when spring peaks and summer declines were not apparent. However, summer declines in the egg ratio of K. taurocephala were observed during each year. The reduction in rotifer populations simultaneously with decreased egg ratios suggested that population declines were caused by food limitation. Food-addition experiments conducted in situ in small enclosures indicated that food was limiting for K. cochlearis when its populations were declining, but not during other periods of the year. Keratella taurocephala did not show a consistent response to food addition.