A simple method for maintaining a laboratory supply of limnetic cyclopoid copepods

The following method is used for maintaining a laboratory supply of freshwater cyclopoid copepods with intervening diapause phase. Animals are collected with a core sampler from the upper sediment surface just after initiation of diapause, i.e. during early summer for summer resting species and mid autumn for winter resting species. The method makes it possible at all times of the year to keep a variety of cyclopoid copepod species ready for experimental work. The following widespread species have been used: Mesocyclops leuckarti, Thermocyclops oithonoides, Diacyclops bicuspidatus, Cyclops vicinus, C. strenuus and C. abyssorum. Probably all cyclopoid copepods with a diapause phase could be stored according to the present method.     

Ecological and evolutionary significance of resting eggs in marine copepods: past, present, and future studies

The occurrence of a resting egg phase in the life cycle of marine and freshwater planktonic copepods is well documented and receiving increasing attention by investigators. The species generally occur in coastal marine waters, freshwater ponds and lakes in areas that undergo strong seasonal fluctuations, though examples have been reported for tropical and sub-tropical areas not subject to such extreme fluctuations. Typically such species disappear from the water column for portions of the year, but remain in the region as benthic resting eggs. Studies to date have focused on the conditions that promote the occurrence of resting eggs, the factors that affect their survival and hatching from sediments, the existence of egg banks in sediments, and the impact of resting eggs on plankton community structure. Benthic resting eggs of copepods include diapause eggs as well as subitaneous (non-diapause) eggs that are quiescent due to conditions in the sediments. As with other groups of organisms the resting egg phase is viewed as being critical for the perpetuation of species year after year, especially those that disappear from the water column for portions of the year. Some data indicate that eggs can survive for many years in sediments which would expand their influence to evolutionary time scales. This paper summarizes our understanding of embryonic dormancy in marine copepods.     

Coexistence with similar life cycles in two species of freshwater copepods (Crustacea)

Two species of freshwater cyclopoid copepods, Cyclops scutifer Sars and Mesocyclops leuckarti (Claus), with different ecological requirements, coexist in three humic ponds near the timberline in boreal forests in south-central Norway. The life cycle in both species is basically regulated by complete depletion of oxygen during the long winter. There is a short period of reproduction in spring followed by a rapid development during summer and autumn to advanced instars, capable of diapause when oxygen depletion starts in early winter. Both species have an exclusive 1-yr cycle.The long winter under anoxic conditions is passed in diapause as large copepodids (late developmental instars) with usually no individuals left in the plankton.Even if co-occurring in the same habitats with similar life cycles, there is a temporal and spatial segregation.     

Dormancy, dispersal and the survival of cyclopoid copepods (Cyclopoida, Copepoda) in a lowland floodplain

1. The survival of cyclopoid copepods was investigated in a floodplain for 2 years where flooding occurred during the cold season. The cyclopoid community was studied in three waterbodies with distinct hydroperiods: a permanent pond connected to the flooded area during inundation, a temporary pool that is part of the flooded area and an isolated temporary pool.
2. Field studies, including data obtained from samples of water, sediment and soil, showed the overall predominance of species with a summer diapause over those with a winter diapause or without diapause. Emergence of cyclopoid copepods at the onset of flooding, examined using emergence traps and data from recently filled or still isolated temporary pools, showed that some species can survive several months of drying.
3. The ability of the diapausing fourth copepodid stages of Cyclops strenuus and C. insignis , the two cyclopoids most abundant during winter and spring flooding, to survive terrestrial conditions was tested in laboratory experiments. Both species survived for several months, but rates differed among the species. A higher percentage of C. strenuus survived for a longer period, possibly explaining why this species was relatively more abundant in more temporary habitats.
4. Both dormancy and dispersal facilitated survival of cyclopoid copepods in transient habitats connected to each other during flood periods. Dormancy was the most important survival strategy, whereas dispersal could be more important following prolonged periods without flooding.     

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).     

Life‐history traits buffer against heat wave effects on predator–prey dynamics in zooplankton

In addition to an increase in mean temperature, extreme climatic events, such as heat waves, are predicted to increase in frequency and intensity with climate change, which are likely to affect organism interactions, seasonal succession, and resting stage recruitment patterns in terrestrial as well as in aquatic ecosystems. For example, freshwater zooplankton with different life‐history strategies, such as sexual or parthenogenetic reproduction, may respond differently to increased mean temperatures and rapid temperature fluctuations. Therefore, we conducted a long‐term (18 months) mesocosm experiment where we evaluated the effects of increased mean temperature (4°C) and an identical energy input but delivered through temperature fluctuations, i.e., as heat waves. We show that different rotifer prey species have specific temperature requirements and use limited and species‐specific temperature windows for recruiting from the sediment. On the contrary, co‐occurring predatory cyclopoid copepods recruit from adult or subadult resting stages and are therefore able to respond to short‐term temperature fluctuations. Hence, these different life‐history strategies affect the interactions between cyclopoid copepods and rotifers by reducing the risk of a temporal mismatch in predator–prey dynamics in a climate change scenario. Thus, we conclude that predatory cyclopoid copepods with long generation time are likely to benefit from heat waves since they rapidly “wake up” even at short temperature elevations and thereby suppress fast reproducing prey populations, such as rotifers. In a broader perspective, our findings suggest that differences in life‐history traits will affect predator–prey interactions, and thereby alter community dynamics, in a future climate change scenario.     

Diapause in copepods (Crustacea) from ephemeral habitats with different hydroperiods in Everglades National Park (Florida,U.S.A.)

Water management practices in the Everglades have severely stressed the natural system, particularly by reducing the hydroperiods of much of the region. During the dry season of 1999, we investigated the influence of hydroperiod on the species composition and dormancy patterns of freshwater copepod communities in seasonal wetlands of Everglades National Park, Florida, U.S.A. The habitats were characterized by an annual dry season, from December through June. We sampled at two locations: the Long Pine Key area of the Rocky Glades region (short hydroperiod, ca. 4–5 months), and western Taylor Slough (intermediate hydroperiod, ca. 8–10 months). Both areas have experienced a reduction in natural hydroperiods and an increase in the frequency of dry-down. We collected weekly plankton samples from Rocky Glades solution holes to assess the potential species pool of copepods. To document the taxa capable of surviving dry-down by resting, we performed three immersion trials in which we rehydrated, in laboratory aquaria, sediment patches from solution holes and surface soils from all stations. Only a subset of the planktonic species collected emerged from the dried sediments. The cyclopoids Microcyclops rubellus and Paracyclops poppei were dominant. This is the first record of diapause for P. poppei. Species distributions from the different hydroperiod soil patches indicated that more diapausing species occurred at the sites that dried for shorter periods. Emerging individuals of M. rubellus and P. poppei were mainly ovigerous females, demonstrating a resting strategy seldom before recorded. The cyclopoid Diacyclops nearcticushad not been previously reported to diapause, but they emerged from the dried sediments in our trials. Our collections included six new records for Florida: Diacyclops nearcticus, Megacyclops latipes, Orthocyclops modestus, Elaphoidella marjoryae, Bryocamptus sp. and Bryocamptuscf. newyorkensis. Paracyclops poppei, Macrocyclops fuscus and Arctodiaptomus floridanus are new records for Everglades National Park. Clearly, diapause is an important strategy for the persistence of copepods in short-hydroperiod wetlands. The duration of the dry period appears to be inversely related to the number of species that emerge from diapause.     

Patterns of life history among cyclopoid copepods of central Europe

• 1 Life history characters (body size of adults, egg diameter, egg sac length and breadth) of nineteen species of central European cyclopoid copepods were measured and sexual size dimorphism (adult female length x adult male length^?1), relative egg size (egg weight X body weight^?1), weight of adult females and of eggs, egg sac shape (egg sac length x egg sac breadth^?1), and reproductive effort (clutch weight produced per female weight per day) were calculated to detect trends in life history strategies.
• 2 Typical planktonic species exhibited the lowest reproductive effort. Among planktonic species, the value for egg sac shape increased with clutch size.
• 3 Large species and small species exhibited different trends in life history characters. Large species had larger clutches, larger eggs, and a greater sex size dimorphism than small species. However, small species had a greater relative egg size.
• 4 Large species live in cold water and reproduce during the spring bloom of phytoplankton where the production of large clutches with relatively small eggs is advantageous. Reserves are unnecessary for juveniles because food is abundant. Small species generally are most abundant during the warm season, when conditions are less predictable, and relatively large eggs, possibly provided with reserves, are advantageous.

     

Semiterrestrial meiofauna inhabiting a wet campo in central Brazil, with special reference to the Copepoda (Crustacea)

The wet campo (campo úmido) marsh type is widely distributed in the cerrado region of central Brazil. These marshes develop on slopes along margins of gallery forests where the water table persists at or near the soil surface year-round. Their grass and sedge vegetation covers spongy, highly organic (not peaty) soils. Ground and surface water seeping through the wet campos tends to be slightly acidic (pH about 5), ion-poor (conductivity less than 10 µS cm^–1) and well oxygenated.A typical freshwater meiofaunal community develops in those wet campos where soils remain moist throughout the year (moisture content more than about 60% of soil wet weight). Such a community was studied from 1979–1982 in a wet campo in a protected natural area on the Fazenda Água Limpa of the Universidade de Brasília. It was dominated by nematodes, rotifers and harpacticoid copepods, and included protozoans, turbellarians, cyclopoid copepods, cladocerans, ostracods, oligochetes, hydracarines and several families of aquatic insect larvae. This community was most fully developed in the wetter areas.Species richness of the copepod community is the highest yet recorded in a freshwater system. The 29 species of harpacticoid copepods and 4 species of cyclopoid copepods displayed pronounced zonation which seemed best correlated with soil moisture content and water regime.     

桡足类滞育规律研究

首次提出用滞育指数对桡足类滞育能力进行相对量化处理 ,并对 10 2种桡足类的滞育能力同迁移能力、体长、栖息地之间的关系进行了数学分析 ,初步得出了滞育在桡足类中的分布规律 :在桡足类中滞育与迁移呈负相关 ,两者之间存在一定程度的置换 ;个体较小的桡足类一般具有较强的滞育能力和较弱的迁移能力 ,个体较大的桡足类一般具有较弱的滞育能力和较强的迁移能力 ;淡水桡足类一般比海洋桡足类具有较强的滞育能力。最后探讨了这种分布模式的成因     

Food habits of small fishes in a common reed <Emphasis Type="Italic">Phragmites australis</Emphasis> belt in Lake Shinji,Shimane, Japan

To clarify the feeding habits of reed fishes, the gut contents of 13 fish species collected in a Phragmites australis belt in Lake Shinji were examined. Six species showed ontogenetic and/or seasonal changes in food use patterns. Smaller individuals generally preyed on small planktonic items (e.g., calanoid and cyclopoid copepods) or small crustaceans (gammaridean amphipods), subsequently changing to other prey items (e.g., mysids and filamentous algae) with growth. The most important dietary items for the reed fish assemblage comprised planktonic copepods, gammaridean amphipods and mysids. However, the relative importance of these changed seasonally, gammaridean amphipods being the most important in autumn and winter, and planktonic copepods and mysids the most important in spring and in summer. Cluster analysis based on dietary overlaps showed that the reed fish assemblage comprised five feeding guilds (planktonic-copepod, mysid, gammaridean-amphipod, filamentous-algae, and detritus feeders). Of these, the three former guilds were the most abundantly represented, whereas detritivores were represented by a single species.     

Diapause of copepods as an element for stabilizing the parasite system of some fish helminths

The analysis of infection dynamics of copepods, which are the intermediate hosts of three helminth species of freshwater fishes (Triaenophorus crassus, Proteocephalus exiguus and P. percae), has shown that diapause in the life cycle of the copepods is favourable for preserving the infection in the waterbody until physiological prerequisites for successful infection of the final host are acquired. The peculiarity of a copepod's life cycle may determine the strategy of a parasite in its preimaginal phase as a waiting stage, and the duration of the residence of helminth larvae in copepods that have an obligatory diapause, is one of the elements the provide stability in parasite systems.     

Algal Diet of Small-Bodied Crustacean Zooplankton in a Cyanobacteria-Dominated Eutrophic Lake

Small-bodied cladocerans and cyclopoid copepods are becoming increasingly dominant over large crustacean zooplankton in eutrophic waters where they often coexist with cyanobacterial blooms. However, relatively little is known about their algal diet preferences. We studied grazing selectivity of small crustaceans (the cyclopoid copepods Mesocyclops leuckarti, Thermocyclops oithonoides, Cyclops kolensis, and the cladocerans Daphnia cucullata, Chydorus sphaericus, Bosmina spp.) by liquid chromatographic analyses of phytoplankton marker pigments in the shallow, highly eutrophic Lake Võrtsjärv (Estonia) during a seasonal cycle. Copepods (mainly C. kolensis) preferably consumed cryptophytes (identified by the marker pigment alloxanthin in gut contents) during colder periods, while they preferred small non-filamentous diatoms and green algae (identified mainly by diatoxanthin and lutein, respectively) from May to September. All studied cladoceran species showed highest selectivity towards colonial cyanobacteria (identified by canthaxanthin). For small C. sphaericus, commonly occuring in the pelagic zone of eutrophic lakes, colonial cyanobacteria can be their major food source, supporting their coexistence with cyanobacterial blooms. Pigments characteristic of filamentous cyanobacteria and diatoms (zeaxanthin and fucoxanthin, respectively), algae dominating in Võrtsjärv, were also found in the grazers’ diet but were generally avoided by the crustaceans commonly dominating the zooplankton assemblage. Together these results suggest that the co-occurring small-bodied cyclopoid and cladoceran species have markedly different algal diets and that the cladocera represent the main trophic link transferring cyanobacterial carbon to the food web in a highly eutrophic lake.     

A simple and effective method for assessing chromatin diminution values in copepods using qPCR

The value of chromatin diminution (CD) in different species of freshwater cyclopoid copepods can differ significantly. The biological and evolutionary roles of these differences remain unclear. To expand the knowledge on CD distribution and magnitude in this group of copepods, a quick method for its evaluation was required. This study proposes a simple approach for CD assessment in copepods using quantitative realtime PCR (qPCR). The magnitude of changes in the genome size was assessed by comparing fluorescence curves of qPCR fragments of target genes for pre- and post-diminution materials. The method was tested on four cyclopoid copepods species. In Cyclops kolensis, CD was assessed as 95.3 ± 1.2; in Acanthocyclops vernalis it was assessed at 94.6 ± 0.8%; at C. insignis, it was 82.3 ± 5.2%; and for the first time, CD was found in Megacyclops viridis at 91.1 ± 2.6%. The advantages of our approach are its rapidity, simplicity and minimal requirements of materials studied.     

Copepod eggs survive a decade in the sediments of the Baltic Sea

Many planktonic calanoid copepod species have been proved to spend a part of their life cycle as benthic resting eggs. In addition to avoiding seasonally occurring unfavourable conditions resting stages may also be used as a long-term survival strategy. The aim of this study was to find out for how long eggs of calanoid copepods retain their viability in the sediments of the Baltic Sea. The occurrence of viable copepod eggs in sea bottom sediment was studied in Pojovik Bay, SW coast of Finland. Eggs were found throughout a 25 cm deep core but deeper than 20 cm they were very scarce. Eggs were incubated at 12 °C in order to check the viability, and their age was estimated by determining the sediment accumulation rate with ¹³⁷Cs-method. Viable eggs were estimated to be 10–13 yrs old, some possibly even 18–19 yrs. Most eggs in the top 8 cm were viable, their age being up to 7–8 yrs. Nauplii that hatched from the eggs belonged to Acartia bifilosa and Eurytemora affinis, A. bifilosa dominating the hatchers of the top sedment layers and E. affinis the deeper layers. Preliminary evidence is presented that E. affinis produces true diapause eggs in the Baltic Sea.     

Genome sizes of cyclopoid copepods (Crustacea): evidence of evolutionary constraint

Genome sizes for 36 species of cyclopoid copepods were determined by DNA-Feulgen cytophotometry of nuclei from adults collected from diverse habitats and locales in North America, South America, Europe, and Asia. Genome sizes are small, show a 20-fold range ( C  = 0.10–2.02 pg DNA), and vary in a discontinuous fashion. The genomes of cyclopoid copepods are remarkably small and constant within each species, unlike the large and variable genomes of marine calanoid species. These differences may reflect the evolutionary antiquity of marine copepods in relation to marine, brackish, and freshwater copepods, as well as differences in mechanisms used to modulate genome size. The small genome sizes of contemporary cyclopoids provide substantive evidence of evolutionary constraint, possibly favouring small genomes, rapid replication rates and accelerated development as adaptive strategies for survival in often fragmented, stressful, and changing habitats. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 625–635.     

Patterns in the distribution of Arctic freshwater zooplankton related to glaciation history

We analysed circumpolar samples from 68 lakes within the 10°C-July isotherm from Arctic Canada, Nunavut, Greenland, Svalbard, Eastern Siberia, the Beringia region, and Alaska. In total, we found 3 species of Anostraca, 17 of Diplostraca, 1 species of cyclopoid and 14 species of calanoid copepods. Our study identifies a wider distribution for some copepods—e.g. Eurytemora pacifica, Leptodiaptomus sicilis, Arctodiaptomus novosibiricus, Cyclops abyssorum—than previously known. Moreover, one anostracan species, Artemiopsis bungei, was recorded in North America for the first time; and one chydoriid, Chydorus gibbus, is a new species for Greenland. We observed that species richness of crustaceans is lower in lakes that were glaciated during the Quaternary period, compared to those not glaciated (e.g. Chukotski Peninsula, Siberia; Point Barrow, Alaska; and Disko Island, Greenland). This confirms the findings of classic studies: glaciation has strongly affected the biogeography of freshwater crustaceans in circumpolar areas. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Population differences in the timing of diapause: a test of hypotheses

Summary The reproductive phenology of the freshwater copepod Diaptomus sanguineus differs markedly between populations residing in two Rhode Island ponds. In a permanent pond the population switches abruptly from making subitaneous (immediately hatching) eggs to diapausing eggs at the end of March each year. In contrast, a temporary pond population switches egg types in May, returns to production of subitaneous eggs in June, and concludes the reproductive season by making diapausing eggs in July. An ESS model suggests that the pattern of diapause expected of a copepod population is a function of annual variation in the onset of harsh conditions (catastrophe date). When variation is relatively low, the superior strategy is for diapause to begin a constant period before the mean catastrophe date. When variation is high, females should make first subitaneous eggs and then diapausing eggs irrespective of the expected catastrophe date. With discrete generations, such a population would alternate between egg types. In the permanent pond, variation of catastrophe date the spring onset of planktivory by sunfish is low, whereas in the temporary pond variation of the catastrophe (pond drying) is high. The model predicts well the phenology of the two copepod populations.In the research reported here, we tested the hypothesis that copepods from the permanent pond, which switch to diapause at the same time every year, are cued by the environment to begin diapause (i.e. by photoperiod, temperature, or both), whereas those from the temporary pond make both egg types regardless of environmental conditions. In opposition to our hypothesis, experimental results indicate that diapause in both populations is cued by the environment. The distinct reproductive phenologies documented in the two populations apparently result from the copepods responding to different environmental cues, rather than one being responsive to the environment while the other is not.     

Control of Diapause by Acidic pH and Ammonium Accumulation in the Hemolymph of Antarctic Copepods

Life-cycles of polar herbivorous copepods are characterised by seasonal/ontogenetic vertical migrations and diapause to survive periods of food shortage during the long winter season. However, the triggers of vertical migration and diapause are still far from being understood. In this study, we test the hypothesis that acidic pH and the accumulation of ammonium (NH₄ ⁺) in the hemolymph contribute to the control of diapause in certain Antarctic copepod species. In a recent study, it was already hypothesized that the replacement of heavy ions by ammonium is necessary for diapausing copepods to achieve neutral buoyancy at overwintering depth. The current article extends the hypothesis of ammonium-aided buoyancy by highlighting recent findings of low pH values in the hemolymph of diapausing copepods with elevated ammonium concentrations. Since ammonia (NH₃) is toxic to most organisms, a low hemolymph pH is required to maintain ammonium in the less toxic ionized form (NH₄ ⁺). Recognizing that low pH values are a relevant factor reducing metabolic rate in other marine invertebrates, the low pH values found in overwintering copepods might not only be a precondition for ammonium accumulation, but in addition, it may insure metabolic depression throughout diapause.