Flood-mediated dispersal versus hatching: early recolonisation strategies of copepods in floodplain ponds

1. We explored the quantitative contribution of two alternative recolonisation strategies by a combination of experimental and field data from the Little Tallahatchie floodplain (Mississippi, U.S.A.). In this area, several floods of short duration occur in winter and spring. During floods, lentic and lotic waterbodies of different size are connected for a limited period. 2. Shortly before the onset of the flood season we collected sediment samples from six temporary ponds. The emergence of seven cyclopoid species was recorded from the experimentally flooded sediments and was quantified over a 4‐week period. Water samples were taken in the same ponds after flooding, from which eight further species were recorded and quantified. 3. The contribution of flood‐dispersed individuals was variable among flooded ponds (between 38 and 94%), but numbers of flood‐dispersed colonists did not differ significantly from those of hatching individuals in flooded ponds. Our results demonstrate the equal importance of both hydrological connections and hatching from dormant stages as pathways for recolonisation by copepods if ponds are hydrologically connected. 4. The significance of recolonisation by hatching individuals in disconnected waterbodies was apparent in an isolated temporary pond that was exclusively recolonised by cyclopoid copepods emerging from the sediment.     

Species assemblages of cyclopoid Copepoda from Mantecal,Venezuela

A 14 year monitoring of species composition and abundance of cyclopoid copepods throughout the rainy and dry seasons in a flooded savanna at Mantecal, Apure State, Venezuela, is presented. Two characteristic habitats within these water bodies were selected: the open center of small ponds free of emergent vegetation and the surrounding flooded grassland. In total, 13 species were registered, all of which appeared in the flooded grassland and 11 of which occurred in the open water. Ten species were common for the two habitats during the dry season. The highest abundance was found in flooded grassland during the dry season. Eight species showed abundances greater than 1.0 individual per liter in flooded grassland during both seasons; 3 and 7 species were this abundant in the open water during rainy and dry seasons, respectively. Mesocyclops meridianus and Microcyclops varicans were the most abundant species in both habitats and seasons, while Thermocyclops decipiens was the dominant species during rainy season in open water. Principal component analysis indicates that the important species were positively correlated between them (sharing habitat and season).     

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.     

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.     

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.     

Lotic meiofaunal community dynamics: colonisation, resilience and persistence in a spatially and temporally heterogeneous environment

1.  The spatial and temporal dynamics of lotic meiofaunal communities were examined with a focus on colonisation, maintenance of populations in lotic environments and persistence of meiofaunal communities.
2.  Lotic meiofaunal colonisation of new habitats may take place via a number of mechanisms and is rapid at both the patch scale (within hours–days) and the drainage basin scale (within 20 years). Successional patterns in lotic meiofaunal communities are evident although data are extremely limited.
3.  Lotic meiofaunal communities appear to be resilient to high flow disturbances. Resilience is moderated by the availability of in-stream refugia and habitat hydrology. Lotic meiofauna may also adopt a 'refuge as habitat' approach to such disturbances.
4.  Lotic cyclopoid copepods possess a common suite of life history characteristics that confers resilience to disturbances. Compared to pelagic planktonic species they have short generation times, many descendants per reproductive cycle and long lifespans. Females in 'source' populations are likely to survive disturbances and so could continuously reproduce over a long period of time producing large numbers of offspring which develop rapidly and recolonise 'sink' areas of the stream.
5.  Persistence of lotic meiofaunal communities is highly variable. Meiofaunal persistence does not increase with increasing proportions of in-stream flow refugia.     

Seasonal Dependent Effects of Flooding on Plant Species Survival and Zonation: a Comparative Study of 10 Terrestrial Grassland Species

Past research has provided compelling evidence that variation in flooding duration is the predominant factor underlying plant species distribution along elevation gradients in river floodplains. The role of seasonal variation in flooding, however, is far from clear. We addressed this seasonal effect for 10 grassland species by testing the hypothesis that all species can survive longer when flooded in winter than when flooded in summer. We carried out an inundation experiment under simulated conditions of summer and winter flooding in the greenhouse. The results showed that all species survived longer under winter floods than under summer floods. However, responses upon flooding were species-specific. All summer flood-tolerant species had high tolerance for winter floods as well, but summer flood sensitive species survived either a little longer, or dramatically longer when flooded under simulated winter conditions. Next, we examined whether winter or summer survival best predicted the lower distribution limits of the species as measured in a natural flooding gradient after an extremely long winter flood. We found a strong significant relationship between the lower distribution limits of species in the field and their tolerance to summer floods, although we measured the lower limits 14 years after the latest major summer flood. In contrast, no such significant relationship existed with species tolerance to winter floods. Some relatively intolerant species occurred at much higher floodplain elevations as was expected from their tolerance to winter inundation in the experiments. This suggests that zonation patterns as created by occasional summer floods may be maintained for a long time, probably due to the limited ability of species to re-colonise lower positions in the floodplain.     

Life cycles of two limnetic cyclopoid copepods, Cyclops vicinus and Thermocyclops crassus, in two different habitats

The life cycles of Cyclops vicinus and Thermocyclops crassusin two shallow eutrophic habitats, Junsainuma and Naganuma Ponds,Hokkaido, Japan, were investigated. Both ponds exhibited similarseasonal patterns of temperature, oxygen levels and pH duringice-free periods; however, oxygen levels were extremely lowerunder the ice in Naganuma Pond. Cyclops vicinus showed differentlife cycles in the two ponds; in Junsainuma Pond, it reproducedin winter and spring (January-May) and entered diapause duringsummer and autumn (June-October) as copepodite IV stage, whileit reproduced in autumn (October-November) and spring (April-May),and entered diapause in summer (June-September) and winter (Januaryand February) as copepodite V stage in Naganuma Pond. Thermocyclopscrassus entered diapause during winter (December-April) as copepoditeIV and V stages in both ponds, and egg-bearing females appearedonly during the warmseason, from early May to late October,when water temperatures were >10°C. Summer diapause inC.vicinus was suggested to be an adaptation against fish predation,whereas C.vicinus entered winter diapause in Naganuma Pond probablyto avoid low oxygen levels. Thermocyclops crassus entered diapausein both ponds to avoid low water temperature. These resultssuggest that biotic and abiotic factors are important for leadingto specific life cycles of cyclopoid copepods in small waterbodies.     

To fly or not to fly: response of water striders to drying out of habitat

Abstract .1. Adults of Aquarius paludum inhabit both temporary and permanent water surfaces; Gerris latiabdominis lives only in temporary habitats. To clarify whether adults of both species stay in position or fly when habitats dry up, overwintered adults of A. paludum and G. latiabdominis collected in spring were reared under one of the following four conditions: (A) on water with sufficient food, (B) on water, starved, (C) on wet paper with sufficient food, (D) on wet paper, starved. All rearings were at LD 15.5:8.5 h, 20 ± 2 °C, resembling natural conditions for April to June.
2. Females of A. paludum in group C had lower fecundity than the control group A and some stopped laying eggs. When a water surface was restored, females that had entered reproductive diapause began to lay eggs again. In contrast, females of G. latiabdominis continued to lay eggs even when reared on damp paper.
3. Adults of A. paludum lived longer and adopted diapause posture with high frequency when starved and reared without a water surface. There were, however, no significant differences in the longevity or in the number of adults showing diapause posture between groups A and D of G. latiabdominis.
4. Females of A. paludum collected in the middle of May had more mature oocytes (mean: 20.8) than females of G. latiabdominis (mean: 8.0), and most had histolysed their indirect flight muscles; most females of G. latiabdominis had retained their flight muscles and flew readily.
5. When water surfaces dry, with food shortage, adults of A. paludum may survive in place for a relatively long time until the water surface returns. Adults of G. latiabdominis may fly to other water surfaces and reproduce without delay.     

Resistance to complete submergence in Rumex species with different life histories: the influence of plant size and light

Resistance to complete submergence was tested in three Rumex species that occur in the Dutch river forelands. The species differ in both habitat and life history characteristics. The annual or biennial R. maritimus and the biennial or short lived perennial R. palustris grow on frequently flooded mud flats of low elevation, while the perennial R. thyrsiflorus can be found on dykes and river dunes that are seldom flooded. The flooding characteristics of the habitats of the three species were determined. These data were used to design experiments to determine the survival and biomass development of the three species during submergence and the influence of plant size and light level on these parameters. It was shown in all three species that plants submerged during daytime were much more resistant to flooding than those submerged at night. This is most probably due to the generation of oxygen or carbohydrates by underwater photosynthesis. Mature plants of the three species showed higher survival after submergence than juvenile plants, which might be caused by higher carbohydrate levels in the taproots of mature plants. In addition, the three species clearly differed in survival and biomass development during submergence. Rumex thyrsiflorus , the species least subjected to flooding, is least tolerant to complete submergence. Rumex maritimus , which can avoid the floods by having a short life cycle, is less tolerant to submergence than R. palustris , which has to survive the floods as a vegetative plant. It was noted that some plants that survived the flooding period itself, still died in the following period of drained conditions, possibly due to post-anoxic injury.     

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).     

Spatial and temporal variations in densities of small fishes across different temporary floodplain types of the lower Okavango Delta,Botswana

Small-sized and juvenile fishes as well as physicochemical water parameters in various primary, secondary and rarely flooded temporary floodplains of the Okavango Delta were assessed during the different hydrological phases of the 2009–2010 flooding season. Small fishes were sampled in the marginal zone of the floodplains using a throw-trap net and a suite of physicochemical properties were measured. Both physicochemical water quality parameters and densities of small fishes showed spatiotemporal variations across the temporary floodplain types. Turbidity, dissolved oxygen, conductivity, woody debris and chlorophyll a were all highest in rarely flooded floodplains compared to both primary and secondary floodplains (ANOVA, p < 0.05). Fish from 38 fish taxa belonging to 11 families were identified and classified during the study. The fish families Cichlidae (dominated by tilapias), Poeciliidae (Aplocheilichthys spp.) and Cyprinidae (Barbus spp.), were the most abundant across the study floodplains. The poeciliids were most abundant in frequently flooded primary and secondary floodplains, whereas juvenile cichlids dominated in rarely flooded floodplains. During high floods the rarely flooded portions of the delta function as important nursery habitats for juvenile cichlids, implying that a significant reduction in flooding may have negative effects on the delta's tilapia stocks due to reduced recruitment success.     

Diapause, a potent force in the evolution of freshwater crustaceans

After a brief historical review of the discovery of diapause in freshwater crustaceans, its dramatic nature in certain cyclopoid copepods, in which diapausing individuals may occur at densities of > 10⁶ per m², is used to illustrate the enormous ecological significance of the phenomenon. Some of the problems presented by dispause in cyclopoid copepods are noted, including the different behaviour in different lakes of what appears to be a single species. Different physiological cues or different genetic endowments are clearly involved.The wider incidence of diapause in freshwater copepods and ostracods is noted.Among freshwater crustaceans it it the Branchiopoda that have universally adopted diapause, always at the egg stage. Even such an ancient order as the Anostraca, perhaps the most primitive of all crustaceans, produces elaborately constructed resting eggs that are capable of cryptobiosis, can remain viable in a dry state for long periods, and can tolerate extreme conditions. The nature of branchiopod resting eggs is briefly reviewed. Of these, only those of the Anomopoda are protected by containers derived from the parental carapace. These are mechanically complex in the most advanced species but, as shown by fossils, are extremely ancient structures.Factors initiating the onset and termination of diapause in branchiopods are briefly noted, and the process of hatching of resting eggs is outlined.     

The role of light for fish–zooplankton–phytoplankton interactions during winter in shallow lakes – a climate change perspective

1. Variations in the light regime can affect the availability and quality of food for zooplankton grazers as well as their exposure to fish predation. In northern lakes light is particularly low in winter and, with increasing warming, the northern limit of some present-day plankton communities may move further north and the plankton will thus receive less winter light.
2. We followed the changes in the biomass and community structure of zooplankton and phytoplankton in a clear and a turbid shallow lake during winter (November–March) in enclosures both with and without fish and with four different light treatments (100%, 55%, 7% and <1% of incoming light).
3. In both lakes total zooplankton biomass and chlorophyll- a were influenced by light availability and the presence of fish. Presence of fish irrespective of the light level led to low crustacean biomass, high rotifer biomass and changes in the life history of copepods. The strength of the fish effect on zooplankton biomass diminished with declining light and the effect of light was strongest in the presence of fish.
4. When fish were present, reduced light led to a shift from rotifers to calanoid copepods in the clear lake and from rotifers to cyclopoid copepods in the turbid lake. Light affected the phytoplankton biomass and, to a lesser extent, the phytoplankton community composition and size. However, the fish effect on phytoplankton was overall weak.
5. Our results from typical Danish shallow eutrophic lakes suggest that major changes in winter light conditions are needed in order to have a significant effect on the plankton community. The change in light occurring when such plankton communities move northwards in response to global warming will mostly be of modest importance for this lake type, at least for the rest of this century in an IPCC A2 scenario, while stronger effects may be observed in deep lakes.     

Community structure in temporary freshwater pools: disentangling the effects of habitat size and hydroregime

1. Hydroregime (duration, frequency and predictability of the aquatic phase) is a key feature of temporary aquatic habitats that not only moulds community structure and diversity (species sorting) but also life history characteristics of the inhabitants (natural selection). However, since hydroregime is a complex multidimensional entity that cannot be estimated from short term observations, morphometric variables are commonly used as proxies for hydroregime, making it impossible to separate effects of habitat size and hydroregime on biota.
2. We have used a simple hydrological model, validated with recent (average r ² = 96%) and historic water level observations (average r ² = 81%), to accurately reconstruct hydroregime based on historical rainfall and evaporation data in a cluster of 36 temporary rock pools in central South Africa.
3. Using the model output, we demonstrated that both hydroregime and habitat size had unique and shared effects on temporary pool biota and that these effects depended on the dispersal modes of the taxa. Model-generated hydrological data explained more variation in community patterns than short-term observations of hydroperiod. Hydroregime was more important for passive dispersers than for active dispersers that can migrate when pools dry up. Basin morphometry was a relatively poor predictor of pool hydroregime. We concluded that simple modelling may greatly improve the resolution of studies linking hydroregime to biological variables.
4. An accurate account of hydrological variation provides a firm foundation to understand community and population structure and dynamics in temporary aquatic habitats. Since many of these habitats have been destroyed or degraded in recent decades, our findings and tools may contribute to the development of reliable conservation guidelines.     

Searching for mechanisms of synchrony in spatially structured gamebird populations

1.  Time series data on five species of gamebird from the Dolomitic Alps were used to examine the relative importance of dispersal and common stochastic events in causing synchrony between spatially structured populations.
2.  Cross-correlation analysis of detrended time series was used to describe the spatial pattern of fluctuations in abundance, while standardized time series were used to describe both fluctuations and the trend in abundance. There were large variations in synchrony both within and between species and only weak negative relationships with distance.
3.  Species in neighbouring habitats were more likely to be in synchrony than species separated by several habitats. Species with similar density-dependent structure were more likely to be in synchrony.
4.  In order to estimate the relative importance of dispersal and environmental stochasticity, we modelled the spatial dynamics of each species using two different approaches. First, we used estimating functions and bootstrapping of time series data to calculate the relative importance of dispersal and stochastic effects for each species. Second, we estimated the intensity of environmental stochasticity from climatic records during the breeding season and then modelled the dispersal rate and dispersal distance for each species. The two models exhibited similar results for rock ptarmigan, black grouse, hazel grouse and rock partridge, while contrasting patterns were observed for capercaillie.
5.  The results suggest that environmental stochasticity plays the dominant role in synchronizing the fluctuations of these galliform species, although there will also be some dispersal between populations.     

Myriapod (Arthropoda,Myriapoda) diversity and distribution in a floodplain forest of the Brazilian Pantanal

This study presents the diversity and distribution of myriapod species associated with different habitats in a seasonally flooded forest composed predominantly of Vochysia divergens Pohl. (Vochysiaceae) in the Brazilian Pantanal. We evaluated species richness distribution across habitats and over different seasonal periods, and also identified possible survival strategies employed during periodic flooding that occur within this forest. A total of 1505 myriapods were sampled, representing four classes and 15 species. Diplopoda and Chilopoda were the most abundant and rich taxa. Pauropoda and Symphyla were represented by only one species each. Species richness was distinctly distributed across habitats evaluated in this forest, as well as over the different seasonal periods. Survival strategies observed in Diplopoda and Chilopoda were primarily associated with vertical migration ranging from the edaphic environment to tree trunks and canopies, and horizontal migration associated with water level before and during the flood period. Species richness was considered low; however, the survival strategies, especially vertical and horizontal migration, demonstrate the adaptation of these species to the seasonal floods of this ecosystem.     

Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types

Aim   We analysed the variation of species richness in the European freshwater fauna across latitude. In particular, we compared latitudinal patterns in species richness and β-diversity among species adapted to different habitat types.
Location   Europe.
Methods   We compiled data on occurrence for 14,020 animal species across 25 pre-defined biogeographical regions of European freshwaters from the Limnofauna Europaea . Furthermore, we extracted information on the habitat preferences of species. We assigned species to three habitat types: species adapted to groundwater, lotic (running water) and lentic (standing water) habitats. We analysed latitudinal patterns of species richness, the proportion of lentic species and β-diversity.
Results   Only lentic species showed a significant species–area relationship. We found a monotonic decline of species richness with latitude for groundwater and lotic habitats, but a hump-shaped relationship for lentic habitats. The proportion of lentic species increased from southern to northern latitudes. β-Diversity declined from groundwater to lentic habitats and from southern to northern latitudes.
Main conclusions   The differences in the latitudinal variation of species richness among species adapted to different habitat types are in part due to differences in the propensity for dispersal. Since lentic habitats are less persistent than lotic or groundwater habitats, lentic species evolved more efficient strategies for dispersal. The dispersal propensity of lentic species facilitated the recolonization of central Europe after the last glaciation. Overall, we stress the importance of considering the history of regions and lineages as well as the ecological traits of species for understanding patterns of biodiversity.     

Recovery of zooplankton diversity in a restored Mediterranean temporary marsh in Doñana National Park (SW Spain)

Over 30 years after drainage for agriculture, a 2700 ha temporary marshland was recently restored in Doñana National Park. We describe the recovery of zooplankton communities (copepods, cladocerans and rotifers) in 47 new temporary ponds excavated as part of the restoration project during the first two hydroperiods (April 2006 and 2007), and compare them to those of eight reference sites in the surrounding marshland. Major changes in the species composition and abundance occurred in new ponds between years. While rotifers and cyclopoid copepods dominated in terms of number of individuals in 2006, calanoid copepods and cladocerans were the most abundant groups in 2007. Rotifer species richness was significantly lower in 2007, but there was an increase in Simpson and β-diversity in 2007 owing largely to a dramatic decline in the abundance of Hexarthra cf. fennica (rare in reference sites) from 93% of all rotifer individuals in new ponds in 2006 to only 32% in 2007. In contrast, species richness of copepods and cladocerans was significantly higher in new ponds in 2007, but there were no changes in Simpson diversity. β-Diversity of cladocerans was also significantly higher in 2007. In 2006, the species richness of cladocera and copepods was significantly lower in new ponds than in reference sites, but by 2007 there were no differences in richness or Simpson diversity. Overall, 7 copepod, 13 cladoceran and 26 rotifer taxa were recorded in new ponds, including 80% of taxa recorded in reference sites. These results indicate that zooplankton communities can be rapidly restored in Mediterranean temporary wetlands, at least when large source populations in the surrounding area reduce dispersal limitation. They also illustrate the importance of comparing different metrics of richness and diversity in studies of zooplankton restoration.     

Consequences of large-scale processes for the conservation of bird populations

1.  Detailed studies of population ecology are usually carried out in relatively restricted areas in which emigration and immigration play a role. We used a modelling approach to explore the population consequences of such dispersal and applied ideas from our simulations to the conservation of wild birds.
2.  Our spatial model incorporates empirically derived variation in breeding output between habitats, density dependence and dispersal. The outputs indicate that dispersal can have considerable consequences for population abundance and distribution. The abundance of a species within a patch can be markedly affected by the surrounding habitat matrix.
3.  Dispersal between habitats may result in lower population densities at the edge of good quality habitat blocks and could partially explain why some species are restricted to large habitat fragments.
4.  Habitat deterioration may not only lead to population declines within that habitat but also in adjacent habitats of good quality. This may confound studies attempting to diagnose population declines.
5.  Although mobile species have the advantages of colonizing sites within metapopulations, dispersal into poorer quality territories may markedly reduce total populations.
6.  There are two main approaches to conservation: one is to concentrate on establishing and maintaining protected areas, while the other involves conservation of the wider countryside. If dispersal is an important process then protecting only isolated areas may be insufficient to maintain the populations within them.