Recruitment limitation along disturbance gradients in river flood plains

Abstract. Question: Along river floodplains lower distribution limits of plant species seem largely determined by their tolerance to rarely occurring floods in the growing season. Such distribution patterns remain fixed for many years suggesting additional effects of winter floods at lower positions. Our objective was to investigate the direct and indirect effects of winter floods on colonization of floodplains in a series of field experiments. Location: River Rhine, The Netherlands. Methods: We measured the direct effects of winter floods on seedling survival and seed removal and survival at low and high floodplain elevation. Indirect effects of winter flooding through changes in the soil were investigated by measuring seedling emergence on soil transplants that were exchanged between high and low floodplain elevation. To investigate indirect effects of floods on the germination environment through changes in the vegetation structure, we measured the effects of vegetation removal on recruitment of sown species. Results: Recruitment was seed limited at both floodplain elevations. An additional effect of vegetation removal on seedling emergence was also observed. Soil types from both zones did not differently affect seedling emergence. Seeds were not removed from the soil surface by a single winter flood. Moreover, seeds remained viable in the soil for at least two years, while the experimental plots were flooded several times during the experimental period. During one of those floods a thick sand layer was deposited at the low zone and subsequently no seedlings were observed anymore. Conclusions: Colonization of low floodplain zones in years between subsequent summer floods is prevented by seed limitation while the direct effects of winter floods are limited except for irregularly occurring sand depositions.     

The effect of flooding and flood timing on leaf litter breakdown rates and nutrient dynamics in a river red gum (Eucalyptus camaldulensis) forest

Comparisons of litter standing-stocks in low-lying and higher areas of the floodplain and the effects of controlled flooding events on leaf litter decomposition and leaf litter nutrients were examined during autumn and winter in a southeastern Australian river red gum (Eucalyptus camaldulensis) floodplain forest. The mean mass of total litter and some litter components was significantly greater in autumn than in winter but there were few differences in litter mass between low-lying flood runners and higher sites (1.5 m) on the floodplain, regardless of season. Leaf decomposition was more rapid in flooded areas than in non-flooded areas and was significantly faster in autumn than in winter. In flooded leaves, concentrations of phosphorus and nitrogen dropped rapidly during the first 3 days of each experiment, increased to near original after 7–10 weeks and then decreased again. After 112 days of decomposition the C:N:P ratios of leaf litter increased, but this effect was most marked for flooded leaves. Simple models of leaf litter dynamics indicated that leaf litter standing-stocks in low-lying flood runners would be reduced by flooding, particularly during autumn. In contrast, models predicted a net gain in standing-stocks of leaf litter to be higher on the floodplain, particularly in autumn. Alteration to the seasonal timing of floods by river regulation has probably decreased litter standing-stocks and nutrients available in low-lying areas of the floodplain to support the production of macrophytes and biofilms during winter and spring floods.     

Summer floods shape meadow butterfly communities in a floodplain nature reserve in Central Europe

How flooding regimes shape temperate-zone butterfly communities has received little attention. At the river Danube in eastern Austria, a levee has largely interrupted natural river dynamics since the late nineteenth century. Only a fraction of the floodplain area still experiences annual summer inundations after snow-melt in the Alps. We surveyed meadow butterfly communities on either side of the levee in a year with an unusually strong flood (2013), and in a season with a weak flood typical for the region (2012). Altogether we observed 67 butterfly species. Butterfly abundance and species richness were lower on meadows with stronger flood impact, but differences were modest. In contrast, species composition differed prominently relative to flooding regime and nectar availability. Grass-feeding species tended to be rarer on flooded meadows, while Brassicaceae-feeding species were more prevalent on nutrient-rich flood-prone meadows. Highly dispersive butterflies made up a larger share on flooded meadows, whereas highly philopatric species were relatively more common at sites with little or no inundation. These results indicate that summer inundations at the river Danube act as filters for the local species composition of butterflies on floodplain meadows. Local resource availability and the differential potential of species to re-colonize meadows after catastrophic floods are likely drivers of these differences. Effects of inundations were not consistently stronger in a year of a catastrophic flood than in a normal season. Butterfly communities on non-flooded meadows had a higher regional conservation value.     

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.     

Conservation of Leafhoppers in Floodplain Grasslands – Trade-off between Diversity and Naturalness in a Northern German National Park

The impacts of mowing and of flooding on the leafhopper communities of a river flood plain were investigated. Samples were taken by a motor-driven suction apparatus. In 2001 leafhoppers were collected in a variety of sites differing in land use (fallows, mown sites) and in flood intensity (high, medium, low, none). In 2002 samples were only taken in fallows subject to different flooding regimes.In fallows, more species (43) were collected than in mown sites (33). Flooding had an effect only in fallows. Here, the most species-rich (29) communities occurred in sites not subject to flooding, whereas fewest species (21) were found in sites subject to regularly occurring long lasting winter floods. Mown sites were dominated by pioneer species. In fallows, the communities differed in respect to flood intensity. In fallows that were subject to summer and winter floods pioneer species prevailed. In contrast, in fallows that were flooded a long time during winter but not in summer, communities of very specialised species were found which were not very species-rich.For the conservation of the typical leafhopper communities of floodplain grassland, management by mowing should be at least reduced if not totally stopped and natural flooding dynamics should be restored.     

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.     

Seasonal flooding,topography, and organic debris interact to influence the emergence and distribution of seedlings in a tropical grassland

In seasonally flooded wetlands, inundation and associated organic debris deposition followed by a drawdown period can promote plant community diversity across space and time. Post‐flood regeneration might be influenced by the direct effect of flooding on seed dispersal and seedling emergence, as well as the indirect effect of organic debris on seed trapping and germination. Our objective was to examine the influence of seasonal flooding, topography, and organic debris cover on seedling distribution in a seasonally flooded grassland. We measured species richness, seedling abundance, and organic debris cover for 3 yr in a seasonally flooded grassland in the Pantanal, Brazil, at three topographic levels at the end of the flood season and during the dry season when there was no debris deposition. A total of 43 species were recorded, with no difference in species richness detected between seasons. However, the abundance of some species was higher post‐flood than during the dry period. The greatest seedling abundance and richness were found post‐flood at intermediate elevations, followed by high and the lowest elevations. Seed germination and seedling establishment were likely suppressed at low topographic positions due to shading from organic debris and poor drainage. Therefore, areas with predictable annual floods promote diversity by creating spatial and temporal variations in environmental conditions.     

A new approach to determining environmental flow requirements: Sustaining the natural values of floodplains of the southern Murray‐Darling Basin

Summary Large overbank flood events play an important role in maintaining large‐scale ecological processes and connectivity along and across the floodplains and between the rivers and their floodplains in the southern Murray‐Darling Basin. However, the regulation of rivers means that extensive overbank flooding can only occur in the rare circumstance of extreme flood events. Recent environmental water allocations have focussed on the largest floodplain blocks (‘icon’ sites) and a small set of specific values (e.g. colonial nesting waterbirds), as well as on trialling fine‐scale manipulation of infrastructure (e.g. pumping) to water relatively small areas. There has been no comprehensive systematic assessment of the entire floodplain and its wider set of flood‐dependent natural assets (such as ecosystems and species; herein referred to as ‘natural values’) to maximise the effectiveness of environmental water use and to catalogue values likely to be lost. This paper describes an assessment of some 220 000 ha found to support flood‐dependent natural values in Victoria. We mapped the geographic distribution and estimated components of the flooding requirements (natural flooding frequency, and maximum period without flooding and minimum duration of each flooding event before significant deterioration) for each natural value. Using an example of one stretch of the River Murray, we show how the resultant spatial data can be used with floodplain inundation modelling to compare the outcomes of real or planned environmental watering events; potentially providing tools for management agencies to conserve a wider range of floodplain values than is currently the case. That is, water managers and the public can see what ecosystems and threatened species are intended to be maintained by environmental watering and what values are intended to be abandoned across the whole floodplain, rather than just seeing the small subset of values and ‘icon’ sites that are intended to be maintained. Examples are provided to illustrate how information about the location, water requirements and extent covered by potential floods for specific values can be used to build adaptive watering strategies for areas as large as the whole floodplain.     

Is tolerance to summer flooding correlated with distribution patterns in river floodplains? A comparative study of 20 terrestrial grassland species

It is generally assumed that floods during the growing season have a strong impact on the distribution of grassland plant species in river floodplains but this proposition has never been tested. We examined the survival and growth responses of twenty species, originating from mid- and high-level floodplain grasslands along the River Rhine in the Netherlands, to total submergence for a maximum of two months in an outdoor flooding experiment. Plant survival and biomass reduction with flooding duration was determined as well as biomass recovery after de-submergence.
Our results indicate that species survival is the most prominent factor correlated with species distribution in floodplain areas. Relatively flood tolerant species occurred mainly at low elevations along the floodplain while more flood sensitive species were restricted to high parts of the floodplain gradient. Biomass reduction rates during submergence were only marginally significantly correlated with species lower distribution boundaries along the flooding gradient. Biomass recovery rate was significantly correlated with species distribution patterns in the field only after 2 weeks of complete submergence, but not after 4 and 8 weeks. Our results suggest that the more flood tolerant species can have various ways to survive and recover from flooding, ranging from low rates of biomass loss and low recovery to relatively high rates of biomass loss and quick recovery.
Our results are consistent with the notion that disturbance by floods during the growing season is an important determinant of species lower distribution boundaries in river floodplains. They also suggest that high survival under flooding may be achieved by different physiological mechanisms. Such mechanisms are discussed in this paper.     

Do terrestrial invertebrates experience floodplains as landscape mosaics? Immediate and longer-term effects of flooding on ant assemblages in a floodplain forest

Considering the floodplain landscape as a mosaic of habitat patches at different successional stages is useful for understanding (1) the processes associated with individual floods and (2) the legacy of flood history. Here, we investigate the applicability of the mosaic model to opportunistic ant species inhabiting the floodplain. Ground-active ant assemblages in river red gum (Eucalyptus camaldulensis) floodplain forest in south-eastern Australia were sampled before and for two years after a major flood in 2000–2001 at 24 sites with different inundation histories. Despite the mobility and opportunistic life history traits of floodplain ants, flood history appeared to impose a persistent mosaic structure on ant assemblages. Increasing duration of inundation of the forest floor was associated with decreasing species richness. β-diversity was low, with the ant species at the most inundation-prone sites being a subset of those at drier sites. Less extensive flooding occurred in 2002–2003, enabling the consistency of short-term responses to inundation to be assessed. Flooding acts as a resetting mechanism, creating a characteristic ant assemblage. After floodwaters receded, there was little evidence of convergence in the structure of ant assemblages through time between sites flooded for different durations. The persistence of dissimilarities in ant assemblages suggests that succession towards terrestrialization was either not occurring or that it was operating at a rate that was too slow to be detected.     

An assessment of the importance of water stress in seasonally floodedC. dactylon on the Pongolo River floodplain in the formulation of guidelines for controlled flooding

Daily variation in water potential was measured inCynodon dactylon in seasonally flooded areas of the Pongolo river floodplain. Soil moisture declined during the dry winter season when the plants were exposed, causing them to experience increasing water stress. Fog and mist appeared to relieve stress but it is concluded that water stress is sufficient to reduce production during exposure. The results are discussed in relation to the timing of flood releases from the upstream Pongolapoort dam and it is concluded that the period of exposure is of less significance than the duration and frequency of inundation in the formulation of guidelines for flood releases from the dam.     

Immediate and longer-term effects of managed flooding on floodplain invertebrate assemblages in south-eastern Australia: generation and maintenance of a mosaic landscape

1. We compared assemblages of ground‐active, terrestrial beetles and spiders from different areas of river red gum Eucalyptus camaldulensis floodplain forest in subhumid, south‐eastern Australia before and for 2 years following a managed flood to determine whether the Flood Pulse Concept is an appropriate ecological model for this regulated, lowland river‐floodplain system. 2. Immediately following flooding, the abundance, species richness and biomass of beetles were greatest at sites that had been inundated for the longest period (approximately 4 months). The abundance, species richness and biomass of spiders were not reduced at sites that were flooded for 4 months compared with unflooded or briefly flooded areas. Sites recently flooded for several months had high densities of predatory, hygrophilic beetles (Carabidae) and spiders (Lycosidae). 3. Over the 2 years following the flood, beetles generally were more abundant at sites that had been inundated for longer. At all sampling times, the species richness of beetles at sites increased with the length of time sites were inundated, even before the flood. Neither the abundance nor species richness of spiders was related to duration of flooding. 4. The structure of beetle and spider assemblages at sites that were flooded for different lengths of time did not appear to converge monotonically over the 2 years after the flood. 5. Managed flooding promotes diversity of beetles and spiders both by providing conditions that create a ‘pulse’ in populations of hygrophilic specialists in the short term, and by creating subtle, persistent changes in forest‐floor conditions. Despite its monotypic canopy, river red gum floodplain forest is a habitat mosaic generated by differing inundation histories.     

The impact of flooding regime on the soil seed bank of flood‐meadows

Abstract. We assessed the significance of flooding for the floristic composition of seed banks in flood‐meadows of the northern valley of the Upper Rhine. We compared three hydrological compartments of the alluvial plain, consisting of the regularly flooded land between the river and low summer dykes (functional flood‐plain), the occasionally flooded land between summer dykes and high winter dykes (hybrid floodplain) and the land behind the winter dykes, which is now only submerged by ascending groundwater (fossil flood‐plain). Due to their different flooding regime, the three compartments should differ with respect to the prevailing conditions of diaspore input. The seed density of soil samples increased with the duration of flooding in the three compartments, while species richness and the proportion of species not occurring in the vegetation was constant. The increase in seed density can be largely attributed to an increase of disturbance indicators, which are present in the above‐ground vegetation and capable of forming a long‐term persistent seed bank. No effects of flooding on the composition of seed banks in the three flood‐plain compartments were found. The differences in seed bank composition can be largely explained by corresponding differences in above‐ground vegetation and former and present‐day meadow management. Seeds of species absent from above‐ground vegetation can be attributed to the local species pool present in the immediate vicinity of the study plots. We discuss consequences of the results for the restoration of species‐rich flood‐plain meadows.     

Colonising Strategies and Diapause of Planktonic Rotifers (Monogononta,Rotifera) during Aquatic and Terrestrial Phases in a Floodplain (Lower Oder Valley,Germany)

The floodplains of the River Oder are inundated in winter and spring and are desiccated during summer. The plankton community developing towards the end of winter is dominated by monogonont rotifers. Some species maintain populations on the flooded areas as well as in the remaining permanent waters after flooding. Other species are found during the inundation period only. These species produce drought‐resistant diapause stages, beginning at low population densities between 5 to 36 females L^–1. Experiments showed that 9 out of 10 species, which were observed to produce resting eggs during the aquatic phase, were able to survive the terrestrial phase through dormancy in the desiccated floodplain soils. Other species colonise the temporarily inundated areas from the permanent waters again during each flooding period.     

Seasonal reversals of upland-riparian diversity gradients in the Sonoran Desert

Flood disturbance and water resource availability vary sharply over time and space along arid‐region rivers and can interact in complex fashion to shape diversity patterns. Plant diversity showed spatial patterning along a topogradient from the floodplain of the San Pedro River (Arizona, USA) to the arid upland, but the patterns shifted temporally as the suite of limiting factors changed. During two of three sampling times, spatial diversity patterns were shaped primarily by gradients of water availability, the regional limiting factor. In the summer dry season, microscale diversity (species richness per 1 m²) and mesoscale diversity (cumulative species and functional types in 20, 1‐m² plots) of herbaceous plants decreased along the topogradient from floodplain to upland, reflecting the greater water availability on the low surfaces. During a summer wet season with moderate rains and flooding, diversity increased in all hydrogeomorphic zones (floodplain, terrace, upland), but the spatial pattern along the topogradient persisted. Following a very wet winter, patterns along the topogradient reversed: scour from large floods limited diversity on the floodplain and competitive exclusion limited the diversity on undisturbed river terrace, while abundant rains allowed for high microscale diversity in the upland. Disturbance and resource availability thus interacted to influence plant species diversity in a fashion consistent with the dynamic‐equilibrium model of species diversity. In contrast to the microscale patterns, mesoscale diversity of species and functional types remained high in the floodplain during all sampling times, with 58% more plant species and 90% more functional types sampled in low floodplain than arid upland for the year as a whole. Species with a wide range of moisture and temperature affinities were present in the floodplain, and seasonal turnover of species was high in this zone. The floodplain zone of a perennial to intermittent‐flow river thus had greater plant diversity than arid Sonoran Desert upland, as measured at temporal scales that capture seasonal variance in resource and disturbance pulses and at spatial scales that capture the environmental heterogeneity of floodplains. Although periodically limited by intense flood disturbance, diversity remains high in the floodplain because of the combination of moderate resource levels (groundwater, seasonal flood water) and persistent effects of flood disturbance (high spatial heterogeneity, absence of competitive exclusion), in concert with the same climatic factors that produce seasonally high diversity in the region (temporally variable pulses of rainfall).     

Response of three arid zone floodplain plant species to inundation

For species to persist on floodplains and in temporary wetlands in arid climates, where large and unpredictable water level fluctuations are common, at least one life history stage must be able to survive inundation. We investigated the survival and performance (RGR, total biomass and above-to-belowground biomass (A:B)) of three common and often coexisting arid zone floodplain species: Xanthium strumarium, Cyperus gymnocaulos and Ludwigia peploides. Observations suggested the species had different responses to inundation, which was tested in a controlled pond experiment. Plants were held at three elevations (+ 10 cm, ? 20 and ? 70 cm) and subjected to three hydrological regimes (static 90 cm, 1 and 5 cm day^?1 inundation) for 16 weeks. Xanthium strumarium died when completely inundated for longer than 4 weeks but when partially flooded survived, showed lower growth rates, increased A:B and produced adventitious roots. C. gymnocaulos showed reduced growth rates when partially flooded and senesced to rhizomes when completely inundated for longer than 4 weeks, which re-sprouted after inundation pressure was removed. L. peploides responded positively to flooding with increased A:B and the production of adventitious roots. The species exhibited three contrasting responses to inundation, which do not necessarily fit neatly within existing water regime functional classification frameworks.     

Nutrient limitation of plant growth on the floodplain of the Narran River, Australia: growth experiments and a pilot soil survey

Water resource development has altered the hydrological regime on the Lower Balonne River in Queensland, Australia. Concerns have been raised about possible impacts to floodplain plant communities, which support a pastoral industry and a range of native fauna. Water and nutrients commonly limit plant growth in south central Queensland, where the climate is semi-arid and the soils are infertile. Floodplain plant productivity is boosted by inundation with water, but the role of flooding in nutrient provision is not known. Growth experiments and a pilot soil survey were conducted to help determine if soil nutrient deficiencies exist and if regular flooding is required to maintain floodplain soil fertility. Soils were sampled from areas representing three flood frequency classes: high, moderate, and low. Chemical extractions were performed as a surrogate for `bioavailable' nutrients. Soil nitrogen (N) but not phosphorus (P) limited the growth of seedlings of wheat (Triticum aestivum L. Gardia) based on responses to nutrient additions: plants supplied with N had greater shoot length and total biomass than plants without N. Clear evidence of an effect of flood frequency on fertility was not revealed. Neither soil P, soil N, nor plant growth varied significantly with flood frequency. However, this analysis had low statistical power. There were trends for greater biomass of seedlings grown on moderately flooded soils and thinner roots for seedlings grown on frequently flooded soils, but neither of these growth responses was clearly linked to nutrient limitation. Nevertheless, the possibility that flooding provides a nutrient subsidy to plants cannot be ruled out because of a number of factors, including the statistical power of this analysis and the possibility that short-term nutrient subsidies occur with floods.     

Fish recruitment in a large,temperate floodplain: the importance of annual flooding,temperature and habitat complexity

1. Large river floodplains are considered key nursery habitats for many species of riverine fish. The lower Volga River floodplains (Russian Federation) are still relatively undisturbed, serving as a suitable model for studying the influence of flooding and temperature on fish recruitment in floodplain rivers. 2. We examined the interannual variability in recruitment success of young‐of‐the‐year (YOY) fish in the lower Volga floodplain in relation to flood pulse characteristics and rising water temperatures in the spring. We sampled four areas with different flooding regimes, in three consecutive years (2006–2008). 3. Extensive areas with a long duration of flooding accommodated high densities of young fish. This suggests that extended inundation improves the recruitment success of river fish. In areas with extensive flooding, the biomass of YOY of most fish species was about three times higher in 2006 and 2007 than in 2008. We hypothesise that low spring temperatures in 2008 may have caused this reduced recruitment and that a flood synchronised with rising temperature enhances recruitment success. 4. Extensive flooding was particularly favourable for species characterised by large body size, delayed maturation, high fecundity and low parental investment, such as pike Esox lucius, roach Rutilus rutilus and ide Leuciscus idus. Gibel carp Carassius gibelio, a species tolerant of high temperature and hypoxia, did particularly well in small waterbodies in the driest parts of the floodplain. 5. Structural characteristics of floodplain waterbodies explained much of YOY fish density. These species–environment associations varied from year to year, but some species such as common bream Abramis brama, roach and gibel carp showed consistent relationships with structural habitat characteristics in all years, despite large interannual fluctuations in flood pulse and spring temperature.     

Frogs during the flood: Differential behaviours of two amphibian species in a dryland floodplain wetland

Flow regimes structure biotic processes and communities of floodplain wetlands. Most current understanding of the influence of the flood pulse on vertebrates is based on knowledge of relationships between flow regimes and fish and bird populations. Its influence on amphibian populations is poorly known, and possibly difficult to disentangle due to the strong influence of weather on amphibian movement and breeding. We predicted that the behaviour of two functionally different amphibian species would vary due to different physiological constraints and life histories driving habitat use and activity periods. We radio‐tracked 52 frogs, 27 barking marsh frogs (Limnodynastes fletcheri, Limnodynastidae) and 25 green tree frogs (Litoria caerulea, Hylidae), to investigate the influence of weather and flooding on habitat use, size of activity area and nightly movement. Activity area sizes and nightly distances of the species were highly variable. Limnodynastes fletcheri had a greater area of activity during floods (mean area: 5237 ± 2556 (SE) m² cf. 639 ± 247 m² post flood) while the tree frog had a greater area after flooding (207 ± 68 m² cf. 1946 ± 530 m²). Limnodynastes fletcheri movement was little influenced by weather or inundation, with distances slightly restricted by increasing wind speed. However, use of floodplain habitat and coincidental breeding behaviour indicated responsiveness by L. fletcheri to flooding. Litoria caerulea moved further with increasing rainfall and nightly temperatures, seldom used flooded areas and there was no evidence of breeding. Although the flood pulse is the dominant structuring force on the floodplain, its influence varies on amphibian species. Consequently for conservation, L. fletcheri populations may be negatively affected by river regulatory activities, but would respond to environmental flow management, whereas L. caerulea appears unlikely to be affected by river regulation, though the confounding effects of rain and preferred habitat inundation needs exploring.     

Base-line variations in stable isotope values in an Arctic marine ecosystem: effects of carbon and nitrogen uptake by phytoplankton

Floods are fundamental for the maintenance of floodplain biodiversity. As a result, well-functioning floodplains are characterized by a high spatio-temporal heterogeneity. Most floodplain-organisms need this shifting landscape mosaic to fulfil their environmental requirements and display a range of adaptations to survive floods. However, in temperate areas, where winter floods are common, extraordinary floods occurring in summer, a period of high physiological activity, may seriously impact the floodplain fauna. This is especially true for guilds characterized by low mobility, such as molluscs. Here we examined the immediate and longer-term response of Elbe grassland molluscs to the extreme 2002 Elbe summer flood in terms of abundance, diversity, and community composition by comparing pre- and post-flood data collected with identical methods. The flood favoured the colonization of aquatic species and led to a shift of the community towards a more hydrophilic composition. Both diversity and abundance increased significantly in the first year following the flood but decreased later gradually to the pre-flood levels. The high spatio-temporal habitat heterogeneity played an important part in the maintenance of mollusc diversity by increasing refuge opportunities and favouring the maintenance of various mollusc communities with different environmental requirements within the floodplain. Handling editor: S. I. Dodson