Population structure, life histories and abundance of copepods in Tjeukemeer, the Netherlands

SUMMARY. The copepod zooplankton of Tjeukemeer was sampled at weekly intervals for 3 years (1969–1971), and ten species were recorded. Five species contributed more than 99% of all individuals collected. Acanthocyclops robustus is the most abundant species (66%), followed by Mesocyclops leuckarti (13%), Cyclops vicinus (11%), Eurytemora affinis (9%) and Diacyclops bicuspidatus (1%).
The ecology of the A. robustus population in Tjeukemeer differs from other A. robustus populations in that it dominates the open water zone. This is an additional argument for a taxonomic revision of the A. vernalis-robustus complex.
Seasonal periodicity in abundance of each species was very similar in 3 successive years.
All cyclopoid species went into diapause, A. robustus and M. leuckarti during winter, D. bicuspidatus and C. v. vicinus during summer. Most species diapause in a particular copepodite stage.
The seasonal variation in the population structure, the fecundity and the sex ratio are described. Both population structure and sex ratio were influenced by diapause and size-dependent mortality. The fecundity was more influenced by the water temperature and the age structure of the female population than by food situation or size of the females.     

Projected impacts of 21st century climate change on diapause in Calanus finmarchicus

Diapause plays a key role in the life cycle of high latitude zooplankton. During diapause, animals avoid starving in winter by living in deep waters where metabolism is lower and met by lipid reserves. Global warming is therefore expected to shorten the maximum potential diapause duration by increasing metabolic rates and by reducing body size and lipid reserves. This will alter the phenology of zooplankton, impact higher trophic levels and disrupt biological carbon pumps. Here, we project the impacts of climate change on the key North Atlantic copepod Calanus finmarchicus under IPCC RCP 8.5. Potential diapause duration is modelled in relation to body size and overwintering temperature. The projections show pronounced geographic variations. Potential diapause duration reduces by more than 30% in the Western Atlantic, whereas in the key overwintering centre of the Norwegian Sea it changes only marginally. Surface temperature rises, which reduce body size and lipid reserves, will have a similar impact to deep‐water changes on diapause in many regions. Because deep‐water warming lags that at the surface, animals in the Labrador Sea could offset warming impacts by diapausing in deeper waters. However, the ability to control diapause depth may be limited.     

The effect of diapause emergence on the seasonal dynamics of a zooplankton assemblage

1. The role of seasonal phenology in the emergence of zooplankton from diapause in patterns of seasonal abundance in the water column was investigated in Oneida Lake, New York. Replicate emergence traps, placed in contact with the lake sediments at two locations (one at a shallow site and one at a deep site), were monitored between May and August.
2. Although six rotifer taxa showed a clear seasonal succession in the water column throughout the study period, all but one taxon emerged exclusively in spring. Three cladoceran and three calanoid copepod species, also present in the water column throughout the study period, again showed predominantly spring emergence. In contrast, three cyclopoid copepod species had distinct seasonal periods of emergence that corresponded, at least in part, to the timing of abundance peaks in the plankton.
3. These results for a single lake are largely consistent with patterns observed or inferred by other investigators for other lakes: variable dependence of abundance in the plankton on diapause emergence for species with long-lived diapausing eggs (i.e. rotifers, cladocerans and calanoid copepods), and much closer dependence for species with short-lived diapausing immature stages (i.e. cyclopoid copepods).     

Embryonic diapause is conserved across mammals

Embryonic diapause (ED) is a temporary arrest of embryo development and is characterized by delayed implantation in the uterus. ED occurs in blastocysts of less than 2% of mammalian species, including the mouse (Mus musculus). If ED were an evolutionarily conserved phenomenon, then it should be inducible in blastocysts of normally non-diapausing mammals, such as domestic species. To prove this hypothesis, we examined whether blastocysts from domestic sheep (Ovis aries) could enter into diapause following their transfer into mouse uteri in which diapause conditions were induced. Sheep blastocysts entered into diapause, as demonstrated by growth arrest, viability maintenance and their ED-specific pattern of gene expression. Seven days after transfer, diapausing ovine blastocysts were able to resume growth in vitro and, after transfer to surrogate ewe recipients, to develop into normal lambs. The finding that non-diapausing ovine embryos can enter into diapause implies that this phenomenon is phylogenetically conserved and not secondarily acquired by embryos of diapausing species. Our study questions the current model of independent evolution of ED in different mammalian orders.     

Life-history traits of lake plankton species may govern their phenological response to climate warming

A prominent response of temperate aquatic ecosystems to climate warming is changes in phenology – advancements or delays in annually reoccurring events in an organism's life cycle. The exact seasonal timing of warming, in conjunction with species-specific life-history events such as emergence from resting stages, timing of spawning, generation times, or stage-specific prey requirements, may determine the nature of a species' response. We demonstrate that recent climate-induced shifts in the phenology of lake phytoplankton and zooplankton species in a temperate eutrophic lake (Müggelsee, Germany) differed according to differences in their characteristic life cycles. Fast-growing plankton in spring (diatoms, Daphnia ) showed significant and synchronous forward movements by about 1 month, induced by concurrent earlier ice break-up dates (diatoms) and higher spring water temperature ( Daphnia ). No such synchrony was observed for slow-growing summer zooplankton species with longer and more complex life cycles (copepods, larvae of the mussel Dreissena polymorpha ). Although coexisting, the summer plankton responded species specifically to seasonal warming trends, depending on whether the timing of warming matched their individual thermal requirements at decisive developmental stages such as emergence from diapause (copepods), or spawning ( Dreissena ). Others did not change their phenology significantly, but nevertheless, increased in abundances. We show that the detailed seasonal pattern of warming influences the response of phyto- and zooplankton species to climate change, and point to the diverse nature of responses for species exhibiting complex life-history traits.     

Impact of predation by Mysis relicta on zooplankton in Flathead Lake, Montana, USA

We documented major changes in the zooplankton community of Flathead Lake following the appearance of Mysis relicta. The three common cladocerans found in the lake decreased in abundance, most notably Daphnia longiremis which disappeared from the lake. Copepods were also affected by M.relicta, especially Diacyclops thomasi which decreased in abundance by an order of magnitude. The only macrozooplankton species which seemed to benefit from the presence of M.relicta was Leptodiaptomus ashlandi. The varied responses of dominant zooplankton species appeared to be the result of a combination of factors. Laboratory feeding studies indicate that M.relicta preferred Cladocera over Copepoda, with the following order of feeding preference: Bosmina longirostris > D.longiremis >, Daphnia thorata > L.ashlandi > D.thomasi. The two zooplankton species which declined most following the appearance of M.relicta showed the greatest degree of habitat overlap with M.relicta. Daphnia longiremis and D.thomasi, together with M.relicta, are cold stenotherms and were concentrated in the hypolimnion during the summer. Finally, another trait shared by D.longiremis and D.thomasi was the absence of a diapause stage, which made them vulnerable to predation by M.relicta throughout the year.      

Evolution of the Arctic Calanus complex: an Arctic marine avocado?

Before man hunted the large baleen whales to near extinction by the end of the nineteenth century, Arctic ecosystems were strongly influenced by these large predators. Their main prey were zooplankton, among which the calanoid copepod species of the genus Calanus, long considered key elements of polar marine ecosystems, are particularly abundant. These herbivorous zooplankters display a range of adaptations to the highly seasonal environments of the polar oceans, most notably extensive energy reserves and seasonal migrations to deep waters where the non-feeding season is spent in diapause. Classical work in marine ecology has suggested that slow growth, long lifespan and large body size in zooplankton are specific adaptations to life in cold waters with short and unpredictable feeding seasons. Here, we challenge this understanding and, by using an analogy from the evolutionary and contemporary history of the avocado, argue that predation pressure by the now nearly extinct baleen whales was an important driving force in the evolution of life history diversity in the Arctic Calanus complex.     

Imidazole derivative KK-42 boosts pupal diapause incidence and delays diapause termination in several insect species

The imidazole derivative KK-42 is a synthetic insect growth regulator known previously to be capable of averting embryonic diapause in several Lepidoptera, but whether it also affects diapauses occurring in other developmental stages remains unknown. In the present study, we examined the effect of KK-42 on pupal diapause in two species of Lepidoptera, the Chinese oak silkworm Antheraea pernyi and the corn earworm Helicoverpa zea, and in one species of Diptera, the flesh fly Sarcophaga crassipalpis. In A. pernyi, KK-42 delayed pupal diapause termination under the long day conditions that normally break diapause in this species. Likewise, in H. zea, KK-42 delayed termination of pupal diapause, a diapause that, in this species, is normally broken by high temperature. KK-42-treated pupae of these two species eventually terminated diapause and successfully emerged as adults, but the timing of diapause termination was significantly delayed. KK-42 also significantly increased the incidence of pupal diapause in H. zea and S. crassipalpis when administered to larvae that were environmentally programmed for diapause, but it was not capable of inducing pupal diapause in H. zea if larvae were reared under environmental conditions that do not normally evoke the diapause response. Experiments with H. zea showed that the effect of KK-42 on pupal diapause was dose- and stage-dependent, but not temperature-dependent. Results presented here are consistent with a link between KK-42 and the ecdysteroid signaling pathway that regulates pupal diapause.     

Biological characteristics of a non‐photoperiodic‐diapause strain of the cabbage beetle Colaphellus bowringi (Coleoptera: Chrysomelidae)

Comparison of biological characteristics between diapausing and non‐diapausing strains of insects provides some insights into the mechanisms regulating diapause. In this study, biological characteristics, especially diapause characteristics and life‐history traits, of a non‐photoperiodic‐diapause (NPD) strain of the cabbage beetle Colaphellus bowringi were compared with those of a normal, high‐diapause (HD) strain that enters diapause in response to either long day length or low temperature. The NPD strain did not enter diapause at any photoperiod at 22°C or higher, but still had a capacity to enter diapause at low temperatures. Although diapause could be induced in both strains by exposure to 20°C, the proportion of individuals having shorter diapause duration was greater in the NPD strain compared to the HD strain. The NPD strain had significantly lower hatching and larval survival rates, longer developmental duration of immature stages, smaller body size and lower longevity and female fecundity compared to the HD strain. The NPD strain of this species is a promising model for investigating diapause regulation in insects in general.     

Distribution and abundance of zooplankton populations in Crater Lake,Oregon

The zooplankton assemblages in Crater Lake exhibited consistency in species richness and general taxonomic composition, but varied in density and biomass during the period between 1988 and 2000. Collectively, the assemblages included 2 cladoceran taxa and 10 rotifer taxa (excluding rare taxa). Vertical habitat partitioning of the water column to a depth of 200 m was observed for most species with similar food habits and/or feeding mechanisms. No congeneric replacement was observed. The dominant species in the assemblages were variable, switching primarily between periods of dominance of Polyarthra-Keratella cochlearis and Daphnia. The unexpected occurrence and dominance of Asplanchna in 1991 and 1992 resulted in a major change in this typical temporal shift between Polyarthra-K. cochlearis and Daphnia. Following a collapse of the zooplankton biomass in 1993 that was probably caused by predation from Asplanchna, Kellicottia dominated the zooplankton assemblage biomass between 1994 and 1997. The decline in biomass of Kellicottia by 1998 coincided with a dramatic increase in Daphnia biomass. When Daphnia biomass declined by 2000, Keratella biomass increased again. Thus, by 1998 the assemblage returned to the typical shift between Keratella-Polyarthra and Daphnia. Although these observations provided considerable insight about the interannual variability of the zooplankton assemblages in Crater Lake, little was discovered about mechanisms behind the variability. When abundant, kokanee salmon may have played an important role in the disappearance of Daphnia in 1990 and 2000 either through predation, inducing diapause, or both. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users.     

Induction of diapause in the pupa of Phalaenoides glycinae by a hormone from the suboesophageal ganglion

The pupal diapause in Phalaenoides glycinae is facultative, and diapause is induced when larvae are reared in short days at low temperatures. By varying the stimulus and especially by changing larvae from one régime to the opposite during their development it is possible to induce diapause covering a wide and continuous spectrum of intensity. Wounding tends to intensify the diapause if it already exists no matter how weakly, but superficial wounding will not put a strongly non-diapausing individual into diapause. In P. glycinae diapause is caused by a hormone secreted by the suboesophageal ganglion. It is suggested that the intensity of the diapause depends on the concentration of the hormone and the duration of the period that the target (the neurosecretory cells of the brain) remains vulnerable. It is suggested that the suboesophageal ganglion may have the function of secreting a diapause-inducing hormone quite generally in short-day and long-day insects with facultative diapause and in species with obligate diapause as well.     

ADAPTATIONS TO TEMPERATE CLIMATES AND EVOLUTION OF OVERWINTERING STRATEGIES IN THE DROSOPHILA MELANOGASTER SPECIES GROUP

The Drosophila melanogaster species group is considered to have originated in the tropics and only recently invaded temperate habitats. The temperate species of this group that were studied here may be subdivided into the warm-temperate species (D. lutescens and D. rufa) and the cool-temperate species (species of the auraria complex). The warm-temperate species were more cold-hardy than were their tropical relatives (D. takahashii or D. melanogaster) at the larval and imaginal stages, and the cool-temperate species were more cold-hardy than the warm-temperate species, although only at the imaginal stage. However, these species showed little or no intraspecific variation in cold-hardiness, in spite of great variation in winter temperature within the species' ranges. It is assumed that cold-hardiness is one of the main factors restricting their distributions at high latitudes and that it is the key for evolution of the warm- and cool-temperate species from their subtropical or warm-temperate ancestors. Both warm- and cool-temperate species had photoperiodically controlled reproductive diapause. In the cool-temperate species, the development of cold-hardiness was affected by diapause, but diapause had little or no effect on cold-hardiness in the warm-temperate species. Critical daylengths and the diapause rates varied from species to species according to variation in their overwintering plans and also varied geographically in consequence of their adaptation to local climates. Species showed different responses to temperature in preimaginal and ovarian development. These differences are considered to reflect adaptation to different environmental temperatures.     

Regional versus local processes in determining zooplankton community composition of Little Rock Lake, Wisconsin, USA

The species present within a community result from a combination of local and regional processes. We experimentally tested the importance of these two processes for lake zooplankton communities by examining the ability of additional species to persist when introduced into mesocosms in Little Rock Lake, Wisconsin, from other nearby lakes in the Northern Highland Lake District. We established a control treatment with only Little Rock Lake zooplankton and two treatments that supplemented the Little Rock communities with zooplankton from nearby lakes. Species richness declined during the 3 weeks of the experiment so that, at the end of the third week, the treatments with added zooplankton species had the same number of species as the controls; increasing the initial number of species in the community did not increase its final species richness. A plot of the mean species richness in the local habitat against the mean species richness of the regional pool fell below a 1:1 slope. This suggested that local processes were more important in structuring Little Rock Lake zooplankton communities.      

Possible impact of mild winters on zooplankton succession in eutrophic lakes of the Atlantic European area

1. We report on a long-term study (1975–94) of water temperatures and plankton in a eutrophic lake (Heiligensee, Berlin, Germany). Using a phenomenological approach, we use historical data to infer how an increase in air temperature has influenced a natural zooplankton community.
2. Air temperatures in Berlin showed a significantly rising trend between 1975 and 1994. Mean winter air temperatures in the last 8 years always exceeded the long-term mean.
3. A rising trend was also found for April water temperature, which increased significantly beginning in 1988–89. An increase of 2.58°C in the last 21 years was recorded using a linear model. A significantly decreasing trend was found in June but no trend was noted for the other summer months.
4. Phytoplankton composition shifted from a dominance of diatoms and cryptophytes during winter and spring in the 1980s towards a dominance of cyanobacteria in 1990–94.
5. The dominant zooplankton species in spring shifted in recent years from the large-bodied Daphnia galeata to the smaller D. cucullata . Cyclops kolensis , previously the only invertebrate predator during winter, decreased in abundance while C. vicinus , usually present during spring and autumn, increased in abundance and was numerous during winter, a season passed in diapause in the earlier years.
6. Because direct and indirect temperature effects are species specific, we put forward the hypothesis that zooplankton species, rather than functional groups, are the nexus between environmental stress, such as rising air temperatures, and ecosystem changes.     

Why study diapause?

This review highlights a number of reasons why insect diapause merits attention. Knowledge of diapause is essential for understanding the seasonal biology of an insect species, and such information is also required for the development of effective pest management strategies; manipulating domesticated species used in pollination and silk production; developing accurate predictive models used to forecast periods of pest abundance; and increasing the shelf‐life of parasitoids and predatory mites used in the biological control industry. Mechanisms used by diapausing insects to survive low temperature may provide tips for cryopreserving insect stocks, a vital need within the research community. Diapause also presents an interesting model for probing fundamental questions in development, and we are indebted to diapause studies for early insights into insect hormones. In addition, I argue that insect diapause may provide insights into questions on aging, obesity and disease transmission, and diapausing insects offer a potentially rich source of pharmaceutical agents that may contribute to improvement of human health.     

Possible impact of mild winters on zooplankton succession in eutrophic lakes of the Atlantic European area

1. We report on a long-term study (1975–94) of water temperatures and plankton in a eutrophic lake (Heiligensee, Berlin, Germany). Using a phenomenological approach, we use historical data to infer how an increase in air temperature has influenced a natural zooplankton community.
2. Air temperatures in Berlin showed a significantly rising trend between 1975 and 1994. Mean winter air temperatures in the last 8 years always exceeded the long-term mean.
3. A rising trend was also found for April water temperature, which increased significantly beginning in 1988–89. An increase of 2.58°C in the last 21 years was recorded using a linear model. A significantly decreasing trend was found in June but no trend was noted for the other summer months.
4. Phytoplankton composition shifted from a dominance of diatoms and cryptophytes during winter and spring in the 1980s towards a dominance of cyanobacteria in 1990–94.
5. The dominant zooplankton species in spring shifted in recent years from the large-bodied Daphnia galeata to the smaller D. cucullata . Cyclops kolensis , previously the only invertebrate predator during winter, decreased in abundance while C. vicinus , usually present during spring and autumn, increased in abundance and was numerous during winter, a season passed in diapause in the earlier years.
6. Because direct and indirect temperature effects are species specific, we put forward the hypothesis that zooplankton species, rather than functional groups, are the nexus between environmental stress, such as rising air temperatures, and ecosystem changes.     

Seasonal dynamics and grazing rate of zooplankton in Yueqing Bay, China

The species composition, biomass, abundance, and species diversity of zooplankton were determined for samples collected from August 2002 to May 2003 from 14 stations in Yueqing Bay, China. Phytoplankton growth rate and microzooplankton grazing rate were obtained by using the dilution method developed by Landry and Hassett. The spatial and temporal variations of zooplankton and its relationship with environmental factors were also analyzed. The results showed that the zooplankton in the Yueqing Bay could be divided into four ecotypes, namely coastal low saline species, estuary brackish water species, offshore warm water species, and eurytopic species. A total of 75 species of zooplankton belonging to 56 genera and 17 groups of pelagic larva were identified in the Yueqing Bay. The coastal low saline species was the dominant ecotype in the study area, and the dominant species were Labidocera euchaeta, Acartia pacifica, Acrocalanus gibber, Pseudeuphausia sinica, and Sagitta bedoti among others. There was considerable seasonal variation in zooplankton biomass and abundance in the surveyed areas. The peak biomass appeared in August, descending in November and in May, and the lowest biomass appeared in February. Similarly, the highest abundance of zooplankton was observed in August, with the abundance descending in the following months: May, November, and February. There were similar horizontal distribution patterns for the biomass and the abundance of zooplankton. They both increased from the upper to the lower bay in February and May, but decreased from the upper to the lower bay in August. Biomass and abundance were evenly distributed in the Yueqing Bay in November. Moreover, there was marked seasonal variation in the species diversity of zooplankton, which conformed to the abundance of zooplankton. Results of the dilution experiments indicated that there was grazing pressure of microzooplankton on phytoplankton in the Yueqing Bay throughout the year though the rate of microzooplankton grazing on phytoplankton varied seasonally. Phytoplanktons were growing at 0.26–2.07/d and grazed by microzooplankton at a rate of 0.15–0.48/d in different seasons. __________ Translated from Acta Ecologica Sinica, 2005, 25(8): 1853–1862 [译自: 生态学报, 2005, 25(8): 1853–1862]