Response of bacterial extracellular enzymes to inundation of floodplain sediments

1. Bacterial extracellular enzymes provide a measure of microbial response to organic matter supply, pivotal to the recovery of riverine food webs after disturbances such as floods.
2. We examined the effects of flood duration on extracellular enzyme response from riverbank and floodplain wetland sediments from the Murrumbidgee River, south-east Australia.
3. There were strong temporal peaks in enzyme activity from both riverbank and billabong sites, peaking between 1 and 5 days following flooding and with a general decline by 21 days. A dominance of non-glucosidase and xylosidase enzymes resulted in no significant differences between billabong and riverbank sediments. This supports the hypothesis that regulated Australian river systems are driven by autochthonous carbon sources.
4. The short response time of the glucosidase after flooding suggests that even short pulses (24 h) in high flows may stimulate bacterial activity, as dissolved organic carbon (DOC) loads also peak at this time. However, a longer wetting time may be needed to drive hydrolysis of the proteins, fatty acids and longer chain polysaccharides, whether in the littoral zone of the river or connection with the floodplain.     

Seed growth suppression constrains the growth of seed parasites: premature acorn abscission reduces Curculio elephas larval size

Abstract.  1. The abscission of seeds infested by insects is common in many plants and has been proposed as a defensive mechanism, although its negative consequences for insects have rarely been assessed.
2. We assessed the consequences of seed abscission studying the interactions between the holm oak Quercus ilex and the chestnut weevil Curculio elephas , its main pre-dispersal seed predator. Female weevils oviposit into the acorns and the larvae must complete their development inside a single acorn feeding on the cotyledons. The growth of the infested acorns is suppressed because they are prematurely abscised.
3. Female weevils oviposit along the acorn growing period; hence, the size of the infested acorns increased with date. The growth of the larvae inside the smaller, early infested, acorns was constrained because food (i.e. cotyledons) was frequently depleted. Larval size increased with the date along with the size of the infested acorns, but it declined slightly in the latest dates as a result of the higher conspecific competition provoked by the larger number of larvae per acorn.
4. The present results demonstrate that premature seed abscission by Q. ilex had negative consequences for C. elephas , as a smaller acorn size reduced food availability and constrained larval size, a key insect life history trait. At the same time, it is suggested that the growth suppression of infested acorns may condition the oviposition phenology of these insects according to their body size. These results need to be considered in further research in the context of the evolutionary significance of premature seed abscission as a defensive mechanism.     

Digging deep for diversity: riparian seed bank abundance and species richness in relation to burial depth

相似文献   

生物结皮对5种不同形态的荒漠植物种子萌发的影响

生物结皮广泛分布于干旱、半干旱区, 强烈影响着土壤表层理化特性, 进而对种子散布、萌发和定居产生影响。目前关于生物结皮与植物种子萌发关系的研究结论存在争议。该文通过室内人工控制实验, 研究了生物结皮对古尔班通古特沙漠5种具不同种子形态特征的荒漠植物白梭梭(Haloxylon persicum)、蛇麻黄(Ephedra distachya)、角果藜(Ceratocarpus arenaarius)、涩芥(Malcolmia africana)和狭果鹤虱(Lappula semiglabra)的种子萌发的影响。结果表明, 在干燥和湿润两种条件下, 生物结皮对不同形态植物种子萌发均具有不同的作用。在干燥条件下, 生物结皮显著抑制了角果藜和涩芥种子的萌发(p<0.05), 对其它3种植物无显著影响; 而湿润条件下, 生物结皮显著抑制了白梭梭、角果藜和狭果鹤虱种子的萌发(p<0.05), 对蛇麻黄、涩芥则无显著影响。     

A fungal endosymbiont affects host plant recruitment through seed- and litter-mediated mechanisms

1.  Many grass species are associated with maternally transmitted fungal endophytes. Increasing evidence shows that endophytes enhance host plant success under varied conditions, yet studies have rarely considered alternative mechanisms whereby these mutualistic symbionts may affect regeneration from seed.
2.  We performed a microcosm experiment to evaluate whether infection with Neotyphodium occultans affects recruitment in the annual grass Lolium multiflorum either directly, by infecting the seeds, or indirectly, by altering the suitability of recruitment microsites through the litter shed by host plants. Endophyte effects on establishment were tested for different litter depths and watering regimes under natural herbivory by leaf-cutting ants.
3.  Seed infection increased seedling emergence through the litter as well as final recruitment, irrespective of microsite conditions. However, litter produced by infected plants delayed emergence and decreased density of both infected and non-infected grass populations.
4.  Individual plant biomass did not change with seed infection but was increased under deep litter from endophyte-infected plants. Although seed infection did not protect establishing plants from leaf-cutting ants, herbivory was reduced in the presence of deep litter shed by infected plants.
5.  We conclude that fungal endophytes may affect host plant recruitment across subsequent generations not only by infecting the seeds but also through the host's dead remains. While the former effect entailed an advantage to infected plants, litter-mediated effects did not discriminate by infection status, and generally promoted the establishment of fewer and larger plants. Thus hidden foliar symbionts may play an underappreciated role in maintaining host species dominance through the litter produced by prior patch occupants.     

Seed bank dynamics in abandoned meadows during a 20-year period in the Białowieża National Park

1 The relationship between size and floristic composition of the seed bank and vegetation dynamics was studied between 1976 and 1996 in 0.75 ha of an abandoned Cirsietum rivularis meadow. The plot was divided into 100-m² (10 × 10 m) quadrats and sampled 5-yearly to map the vegetation and determine the soil seed bank.
2 Densities of seeds in the soil fluctuated as succession proceeded. The initially small seed bank trebled by 15 years after abandonment, before falling, after 20 years, to approximately the same as in the initial stage.
3 The floristic richness of the seed bank decreased during succession, with the number of species falling from 38 to 25. The diversity of life forms, however, increased in later periods, with tall herbs, shrubs and trees appearing after 10 years.
4 Seed bank floristic composition is apparently both a product of the species composition of the current vegetation and a record of the long-term substitution of species. Other factors, including the structure of the vegetation, also influence the accumulation of seeds in the soil.
5 Although changes in number of species show a directional pattern, the seed bank size fluctuated in the course of succession on these fertile wet meadows.     

The influence of floods and precipitation on <Emphasis Type="Italic">Tamarix</Emphasis> establishment in Grand Canyon,Arizona: consequences for flow regime restoration

Decoupling of climate and hydrology combined with introduction of non-native species creates novel abiotic and biotic conditions along highly regulated rivers. Tamarix, a non-native shrub, dominates riparian assemblages along many waterways in the American Southwest, including the Colorado River through Grand Canyon. We conducted a tree-ring study to determine the relative influences of climate and hydrology on Tamarix establishment in Grand Canyon. Riparian vegetation was sparse and annually scoured by large floods until completion of Glen Canyon Dam, which allowed pioneer species, including Tamarix, to expand. Post-dam floods in the mid-1980s were associated with high Tamarix mortality but also initiated a large establishment event. Subsequent establishment has been low but continuous with some exceptions. From 1984 to 2006 establishment increased during years of high, late-summer flows followed by years of low precipitation. This combination provided moist surfaces for Tamarix establishment and may have caused reduced erosion of seedlings or reduced competition from native plants. Attempts to mimic pre-dam floods for ecosystem restoration through planned flood releases also have affected Tamarix establishment. Early (March 1996) and late (November 2004) restoration floods limited establishment, but a small restoration flood in May 2000 followed by steady summer flows permitted widespread establishment. Flood restoration is not expected to prevent Tamarix spread in this system because historic flood timing in May–July coincides with seed release. To decrease future Tamarix establishment, river managers should avoid floods during peak Tamarix seed release, which encompasses the historic spring and early summer flooding period. Tamarix dominance may be reduced by early spring floods that initiate asexual reproduction of clonal shrubs (e.g., Salix exigua, Pluchea sericea).     

Mast seeding in Abies balsamea, Acer saccharum and Betula alleghaniensis in an old growth, cold temperate forest of north-eastern North America

1 Several hypotheses have been put forward to explain the phenomenon of masting or mast seeding, i.e. the supra-annual, periodic production of a large number of seeds in long-lived plants. Some of these hypotheses deal with the proximate causes of masting (e.g. the climate hypothesis) but others are concerned mostly with ultimate, evolutionary explanations (e.g. the pollination efficiency hypothesis).
2 The seed production of three tree species, Abies balsamea , Acer saccharum and Betula alleghaniensis , was followed over a 7-year period in an old-growth, cold temperate forest of north-eastern North America. The main objectives were to determine the extent of interannual variations in seed production, to investigate the relationship between viable and potential seed crop and crop efficiency, and to explore the effects of climate on seed production.
3 Potential and viable seed production varied significantly among years for all three species. However, the timing of dispersal remained the same regardless of the level of seed production.
4 Seed rain was spatially less heterogeneous in years of high seed production, suggesting that most trees were reproducing in such years.
5 Over the 7-year period, there was a significant concordance among species in their viable seed crop and crop efficiency, but not in their potential seed crop. Crop efficiency was positively correlated to potential seed crop for Abies and Betula , but not for Acer .
6 High seed production was related to warm, dry conditions in the spring of the previous year (i.e. at reproductive bud initiation) but to a moist summer in the year of seed maturation.
7 Masting in these three species thus appears to be controlled by several factors, including climate and pollination efficiency.     

Carbon source for reproduction in a spring ephemeral herb, Corydalis ambigua (Papaveraceae)

1.  The carbon source for reproduction in plants may differ between flowering and fruiting stages. To clarify how spring ephemerals use current photosynthetic products for reproduction, the allocation patterns of photosynthate at flowering and fruiting and the effects of resource limitation on reproductive performance in Corydalis ambigua were assessed.
2.  A ¹³C tracing experiment revealed that about 20% of the current photosynthetic carbon was used for reproduction at both flowering and fruiting. The proportion of ¹³C allocated to fruits was constant irrespective of the light level. In contrast, ¹³C translocation to tubers increased at fruiting, and this trend was accelerated when plants were shaded.
3.  Defoliation treatment significantly reduced nectar production and tuber mass, while seed production was not affected. Therefore, when carbon assimilation was limited, carbon was preferentially allocated to current reproduction (seeds) rather than to pollinator attraction (nectar) or storage (tuber).
4.  If seed production is partly supported by carbohydrate reserved in the old tissue of tubers, nectar and seed production may not compete strongly for carbon sources. In contrast to the ability of high seed production, the susceptibility of nectar production to current photosynthesis indicates that seed production of this species is basically limited by pollen capture.
5.  Therefore, temporal separation of resource pool for reproduction may mitigate the joint limitation of seed production between pollinator attraction and resource availability. Temporal variation of the sink–source balance of storage organ is crucial to understand the cost of reproduction in perennial plants.     

生物土壤结皮对种子萌发和幼苗建植的影响研究进展

生物土壤结皮（BSC）在陆地生态系统中具有重要的生态地位,尤其是旱地生态系统中,BSC占据了种子植物之间的广阔地面。因此,BSC的发展必然影响种子植物更新过程与植被空间格局;但其作用方式、影响程度等因相关研究涉及多气候要素、土壤类型、BSC组成物种和种子植物物种的差异及其不同组合,导致目前的研究结论存在广泛争议。研究综合论述了BSC改变地表微形态对种子传播过程的影响;BSC改变土壤特性（物理、化学、生物学）对种子萌发和幼苗存活与建植等关键环节的影响;并结合种子形态特征及种子萌发、幼苗建植的性状等,综合分析了BSC对种子传播、种子萌发与幼苗建植等更新过程的潜在影响机理;探讨了目前研究矛盾性结论产生的原因。总体来说,深入研究并全面揭示BSC对种子植物更新过程的影响,应加强学科交叉,将分子生物学、植物生理学、生物化学等微观研究,与遥感、野外生态因子过程监测、控制实验等宏观、中观研究结合,从机理到过程方面动态研究BSC对种子植物更新过程的影响,并引入水文模型、气候模型、种群动态模型等模型预测方法,研究气候变化、各类干扰频发的情景下,BSC对种子传播、萌发及幼苗建植过程的潜在影响,以期促进对BSC与种子植物间相互关系的研究,加深对干旱脆弱生态系统植被发展规律的认识。     

Effects of grazing and ant/beetle interaction on seed production in the legume Vicia sepium in a seminatural grassland

Abstract.  1. Seed production can be affected by cattle grazing, insect herbivores, and seed predators. Bush vetch, Vicia sepium L. (Fabaceae), possess extra floral nectaries (EFNs), which can attract ants that act as plant defenders.
2. In this field study, four different manipulations were used to investigate interactions between V. sepium , ants, aphids, and seed predators, mainly Bruchus atomarius. The experiment was set up in grazed and non-grazed plots in a semi-natural grassland in south-central Sweden.
3. Grazing negatively affected number of V. sepium buds, flowers, and pods. Seed set, defined as the number of developed seeds per number of ovules in a pod, was higher in grazed plots. Ant abundance, however, was not affected by grazing. Ten different ant species were found on V. sepium and the number of ants was positively correlated with the presence of EFNs. Aphids were found only when ants were excluded from the plants. Pod and seed production of V. sepium plants was not affected by the presence of ants. The seed predator B. atomarius was not affected by the presence of ants. No support was found for the supposition that V. sepium benefited from attracting ants by producing extra floral nectar.
4. A small observational study of V. sepium plants in shrub ( Rosa dumalis ) plots compared with grassland plots was added to the experimental study . Vicia sepium plants were longer and had higher seed-set (seeds/ovules) in shrub plots than in grassland plots.     

Restoration strategies for river floodplains along large lowland rivers in Europe

1. Most temperate rivers are heavily regulated and characterised by incised channels, aggradated floodplains and modified hydroperiods. As a consequence, former extensive aquatic/terrestrial transition zones lack most of their basic ecological functions.
2. Along large rivers in Europe and North America, various floodplain restoration or rehabilitation projects have been planned or realised in recent years. However, restoration ecology is still in its infancy and the literature pertinent to river restoration is rather fragmented. (Semi-) aquatic components of floodplains, including secondary channels, disconnected and temporary waters as well as marshes, have received little attention, despite their significant contribution to biological diversity.
3. Many rehabilitation projects were planned or realised without prior knowledge of their potential for success or failure, although, these projects greatly contributed to our present understanding of river–floodplain systems.
4. River rehabilitation benefits from a consideration of river ecosystem concepts in quantitative terms, comparison with reference conditions, historical or others, and the establishment of interdisciplinary partnerships.
5. We present examples from two large European rivers, the Danube and the Rhine, in which the role of aquatic connectivity has been extensively studied. The Danube delta with its diversity of floodplain lakes across an immense transversal gradient (up to 10 km) serves as a reference system for restoration projects along lowland sections of large rivers such as the Rhine in the Netherlands.     

Adaptations to Flooding Stress: From Plant Community to Molecule

Abstract: This review highlights four major topics in plant flooding research: the processes underlying vegetation zonation in the floodplain, the challenges of using model species to reveal adaptive responses in shoots and roots, the role of micro-organisms in flooded soils in relation to plant growth, and the molecular regulation of the hormone ethylene which is heavily involved in the adaptation reaction of flood-resistant plants. Model species and vegetation strategies are used to unravel mechanisms of vegetation zonation in the river flood-plain. In the case of woodlands, hydrological conditions determine to a large extent their zonation patterns under natural conditions. For softwood species, such as Salicaceae, the interaction between water levels and timing of seed dispersal is the dominating process determining their establishment success on river banks. Their strategy is well adapted to irregular, high and prolonged floods. Hardwood species, Quercus, Fraxinus, UImus and Acer, are flood-sensitive and inhabit the higher sites. They mainly have heavy seeds and germinate under shaded conditions. The most shade-tolerant hardwood species are the least well adapted to flooding. Anthropogenically influenced parts of the floodplain are characterized by grasslands with elevation level and management practices determining the species composition. Low-lying grasslands have flood-tolerant species; elevated zones are seldom flooded and have flooding-sensitive species. Following Grime (1998^[59]), plant species of major vegetation types within the floodplain zone can be divided into three categories–dominants, subordinates and transients–illustrating the diversity in plant species in relation to environmental properties. Model species that are indicative of the different conditions in the various zones are chosen to help in the understanding of morphological and physiological adaptations at the plant level. The formation of aerenchymatous roots and the capacity to elongate shoot parts upon submergence are among the main responses of surviving plants. The role of hormones in the adaptation reaction is emphasized. Owing to high porosities in roots of flood-tolerant plants, radial oxygen loss greatly influences nitrification and denitrification processes in the flooded soil. Nutrient cycles are restored by root-derived oxygen and the oxygenated rhizosphere is detoxified. A new development in flooding ecology is the unravelling of the molecular regulation of hormonally controlled processes. The expression of an ethylene receptor gene in Rumex palustris is highlighted. This paper ends with some suggestions for future flooding research.     

Analysis of Naturalization Alternatives for the Recovery of Moist-soil Plants in the Floodplain of the Illinois River

The hydrologic regime of the Illinois River has been substantially altered by floodplain levees, navigation dams, and water diversion. Unnaturally frequent and untimely water level fluctuations, large and small, have decreased the productivity of many floodplain vegetation communities that provide important ecological services, including the moist-soil plant community. We simulated three scenarios, including two that were expected to benefit moist-soil plants: (1) existing conditions, with levees and navigation dams closed during the summer growing season; (2) levees opened to reconnect the river and its floodplain during the growing season; and (3) both the downstream navigation dam and the levees opened during the growing season. A 1-dimensional hydraulic model generated daily hydrographs of the river at three positions in the 135 km study reach: (1) near the downstream dam, (2) in the middle of the reach, and (3) near the upstream dam. These hydrographs then were used to run a model that predicts the growth of moist-soil plants at a range of floodplain elevations. As expected, the model predicted that plants would grow over a larger area with levees open during the growing season than under the existing conditions, but the outcomes showed a strong location dependency. Moist-soil plant production would increase in the upper and mid-reach locations, but there would be no change near the downstream dam despite opening the levees. Modelling revealed that the existing operation of the navigation dam permanently floods most of the floodplain zone where moist soil plants might grow for at least 15 km upstream of the dam. Trees currently grow all the way to the low water line and are likely to exclude moist soil plants from any restored portion of the floodplain. Sites for reconnecting the river with its floodplain should be carefully chosen to maximize the chances of recovering the important moist-soil plant community in this regulated river.     

The effect of water supply on seed-bank analysis using the seedling-emergence method

1. When using the seedling-emergence method to analyse a soil seed bank the greenhouse conditions should match the germination requirements of the species involved. Although the seedling-emergence method is common practice in ecological studies, the germination characteristics of many species are not known, or are only partly known.
2. Before carrying out a large-scale seed-bank study in a wet dune slack, we tested the water requirements of the species in the seed bank, comparing a waterlogged soil with a moist soil. Four species germinated in significantly larger numbers in a waterlogged soil, seven species in a moist soil and 11 species showed no significant difference. When a species was present in low densities, it was often missed using the less appropriate treatment.
3. These results emphasize the need for a preliminary study before carrying out a seed-bank analysis and show the danger of using a 'standard' method to analyse soil samples from different habitats.     

Geomorphic dynamics of floodplains: ecological implications and a potential modelling strategy

1.  The dynamics of channel migration and floodplain renewal constitute an important control of the ecological diversity of river corridors. Restoration initiatives should therefore assess whether these dynamics must be reinstated in order to address the cause rather than the symptoms of floodplain biodiversity decline.
2.  Restoration of reach-scale dynamism in rivers where this is a natural behavioural process will restore smaller-scale geomorphological and sedimentological processes that encourage vegetation regeneration, but may require catchment-scale management of material flows.
3.  Channel dynamics depend on the style of river–floodplain interaction, and this may be summarised in qualitative, classificatory, sedimentological models of floodplain architecture that have been somewhat neglected in the ecological literature.
4.  One approach to the assessment of floodplain biodiversity and its restoration would be through the development of simulation models based on specified channel styles, and involving simplified hydrodynamics and successional changes. Such models, currently the subject of research as a spin-off from modelling studies of landscape evolution, would permit evaluation of the consequences for ecological diversity of implementing various management options that may affect the dynamics of channel migration.     

Half a century of changes in the riverine landscape of Limay River: the origin of a riparian neoecosystem in Patagonia (Argentina)

Our study aims to determine the dynamic that led to the spread of exotic Salicaceae on the Limay River floodplain and its implications in shaping the current neoecosystem. We used images obtained by the HRG sensor on board the SPOT-5 satellite. We selected two images dates allowed a comparison of the floodplain under different flooding regimes programmed by the Interjuridiccional Basin Authority, at a rate of 1290 m³/s in Spring and a flow rate of less than 400 m³/s in Summer. To characterize the vegetation cover, the Normalized Difference Vegetation Index (NDVI) was used to compare images of September and December, also the Normalized Difference Water Index (NDWI) was applied to delineate the riverbed and floodplain in September. To evaluate the influence of flooding regime on the detected patches a Principal Component Analysis (PCA), was performed; and biotic and abiotic factors on the composition of the dominant tree species in each patch by Multiple Factor Analysis (MFA) were analized. A 58.4 % coverage of forest patches had developed on the Limay River floodplain. Patches of older trees grew on surfaces at 1.5 m above the water level. However, the surfaces not reached by floods >1.5 m have a very low rocky coverage (1 %). The analysis of the age of the trees downstream of the dam system showed that the vegetation, often exposed to high floods before the dams were built (1971), was composed of the native willow (Salix humboldtiana) and the exotic Salix alba. As the river regime was attenuated and extraordinary floods disappeared with the operation of the dam Arroyito in 1980, the first patches of Populus nigra spread.     

Plant Establishment from the Seed Bank of a Degraded Floodplain Wetland: A Comparison of Two Alternative Management Scenarios

There has been little research examining the soil seed banks of degraded floodplain wetlands and their contribution to wetland rehabilitation in Australia. Our aim was to assess the establishment of plants from the seed bank that may occur following the delivery of an environmental water allocation to Kanyapella Basin, a 2950 ha wetland located on the floodplain of the Goulburn and Murray Rivers in northern Victoria, Australia. Two hypothetical water regimes were investigated (flooded and dry) in a glasshouse experiment, where plants were left to establish from the seed bank over a period of 124 days. Differences in the establishment of plants from the seed bank indicated that the return of a flooding regime is likely to have a significant effect on the composition of the wetland vegetation. Mapping of the distribution of plant species indicated that propagules were highly dispersed across the wetland for the majority of taxa, in contrast to the localised distribution of many of the plant species represented in the extant vegetation. Inundation favoured the establishment of native wetland and floodplain plants, although many areas of Kanyapella Basin that are currently ‘weed-free’ have the potential to become colonised and potentially dominated by introduced plants if the wetland is not managed appropriately. Overall, results supported the aim of management to reestablish a wetting and drying regime through use of an environmental water allocation. This study presents a significant example of the application of seed bank investigations in wetland ecology and management.     

The interactive effects of climate, life history, and interspecific neighbours on mortality in a population of seed harvester ants

Abstract.  1. Fifteen years of census data from a population of the red harvester ant ( Pogonomyrmex barbatus ), 5 years of census data from populations of Aphaenogaster cockerelli and Myrmecocystus mimicus at the same site, and long-term precipitation data from a nearby weather station were used to examine how nearest interspecific neighbours, climate, and life history influence P. barbatus colony mortality.
2. It was found that more P. barbatus colonies die following dry summers than following wet summers, but there is no relationship between the establishment of new colonies and rainfall.
3.  Aphaenogaster cockerelli , but not M. mimicus , nearest neighbours influence P. barbatus colony mortality in some years and for some ages.
4. The effects of nearest A. cockerelli neighbours are usually most severe following dry summers, when resource availability is reduced.     

Effects of patch size and density on flower visitation and seed set of wild plants: a pan-European approach

1.  Habitat fragmentation can affect pollinator and plant population structure in terms of species composition, abundance, area covered and density of flowering plants. This, in turn, may affect pollinator visitation frequency, pollen deposition, seed set and plant fitness.
2.  A reduction in the quantity of flower visits can be coupled with a reduction in the quality of pollination service and hence the plants' overall reproductive success and long-term survival. Understanding the relationship between plant population size and/or isolation and pollination limitation is of fundamental importance for plant conservation.
3.  We examined flower visitation and seed set of 10 different plant species from five European countries to investigate the general effects of plant populations size and density, both within (patch level) and between populations (population level), on seed set and pollination limitation.
4.  We found evidence that the effects of area and density of flowering plant assemblages were generally more pronounced at the patch level than at the population level. We also found that patch and population level together influenced flower visitation and seed set, and the latter increased with increasing patch area and density, but this effect was only apparent in small populations.
5.   Synthesis. By using an extensive pan-European data set on flower visitation and seed set we have identified a general pattern in the interplay between the attractiveness of flowering plant patches for pollinators and density dependence of flower visitation, and also a strong plant species-specific response to habitat fragmentation effects. This can guide efforts to conserve plant–pollinator interactions, ecosystem functioning and plant fitness in fragmented habitats.