Scale-dependent topographic complexity underpins abundance and spatial distribution of ecosystem engineers on natural and artificial structures.

In response to ongoing coastal urbanization, it is critical to develop effective methods to improve the biodiversity and ecological sustainability of artificial shorelines. Enhancing the topographic complexity of coastal infrastructure through the mimicry of natural substrata may facilitate the establishment of ecosystem engineering species and associated biogenic habitat formation. However, interactions between ecosystem engineers and their substratum are likely determined by organismal size and resource needs, thus making responses to topography highly scale-dependent. Here, we assessed the topographic properties (rugosity, surface area, micro-surface orientations) that underpin the abundance and distribution of two ecosystem engineers (fucoids, limpets) across six spatial scales (1-500 mm). Furthermore, we assessed the 'biogenic' rugosity created by barnacle matrices across fine scales (1-20 mm). Field surveys and 3D scanning, conducted across natural and artificial substrata, showed major effects of rugosity and associated topographic variables on ecosystem engineer assemblages and spatial occupancy, while additional abiotic environmental factors (compass direction, wave exposure) and biotic associations only had weak influences. Natural substrata exhibited ≤67 % higher rugosity than artificial ones. Fucoid-covered patches were predominantly associated with high-rugosity substrata and horizontal micro-surfaces, while homescars of limpets (≥15 mm shell length) predominated on smoother substratum patches. Barnacle-driven rugosity homogenized substrata at scales ≤10 mm. Our findings suggest that scale-dependent rugosity is a key driver of fucoid habitat formation and limpet habitat use, with wider eco-engineering applications for mimicking ecologically impactful topography on coastal infrastructure.

Topography-based modulation of environmental factors as a mechanism for intertidal microhabitat formation: A basis for marine ecological design.

Topographic complexity is often considered to be closely associated with habitat complexity and niche diversity; however, complex topography per se does not imply habitat suitability. Rather, ecologically suitable habitats may emerge if topographic features interact with environmental factors and thereby alter their surrounding microenvironment to the benefit of local organisms (e.g., resource provisioning, stress mitigation). Topography may thus act as a key modulator of abiotic stressors and biotic pressures, particularly in environmentally challenging intertidal systems. Here, we review how topography can alter microhabitat conditions with respect to four resources required by intertidal organisms: a source of energy (light, suspended food particles, prey, detritus), water (hydration, buffering of light, temperature and hydrodynamics), shelter (temperature, wave exposure, predation), and habitat space (substratum area, propagule settlement, movement). We synthesize mechanisms and quantitative findings of how environmental factors can be altered through topography and suggest an organism-centered 'form-follows-ecological-function' approach to designing multifunctional marine infrastructure.

The north-south divide? Macroalgal functional trait diversity and redundancy varies with intertidal aspect.

BACKGROUND AND AIMS: Marine macroalgae ('seaweeds') are critical to coastal ecosystem structure and function, but also vulnerable to the many environmental changes associated with anthropogenic climate change (ACC). The local habitat conditions underpinning observed and predicted ACC-driven changes in intertidal macroalgal communities are complex and probably site-specific and operate in addition to more commonly reported regional factors such as sea surface temperatures. METHODS: We examined how the composition and functional trait expression of macroalgal communities in SW England varied with aspect (i.e. north-south orientation) at four sites with opposing Equator- (EF) and Pole-facing (PF) surfaces. Previous work at these sites had established that average annual (low tide) temperatures vary by 1.6 °C and that EF-surfaces experience six-fold more frequent extremes (i.e. >30 °C). KEY RESULTS: PF macroalgal communities were consistently more taxon rich; 11 taxa were unique to PF habitats, with only one restricted to EF. Likewise, functional richness and dispersion were greater on PF-surfaces (dominated by algae with traits linked to rapid resource capture and utilization, but low desiccation tolerance), although differences in both taxon and functional richness were probably driven by the fact that less diverse EF-surfaces were dominated by desiccation-tolerant fucoids. CONCLUSIONS: Although we cannot disentangle the influence of temperature variation on algal ecophysiology from the indirect effects of aspect on species interactions (niche pre-emption, competition, grazing, etc.), our study system provides an excellent model for understanding how environmental variation at local scales affects community composition and functioning. By virtue of enhanced taxonomic diversity, PF-aspects supported higher functional diversity and, consequently, greater effective functional redundancy. These differences may imbue PF-aspects with resilience against environmental perturbation, but if predicted increases in global temperatures are realized, some PF-sites may shift to a depauperate, desiccation-tolerant seaweed community with a concomitant loss of functional diversity and redundancy.

Victim of changes? Marine macroalgae in a changing world.

BACKGROUND: Marine macroalgae ('seaweeds') are a diverse and globally distributed group of photosynthetic organisms that together generate considerable primary productivity, provide an array of different habitats for other organisms, and contribute many important ecosystem functions and services. As a result of continued anthropogenic stress on marine systems, many macroalgal species and habitats face an uncertain future, risking their vital contribution to global productivity and ecosystem service provision. SCOPE: After briefly considering the remarkable taxonomy and ecological distribution of marine macroalgae, we review how the threats posed by a combination of anthropogenically induced stressors affect seaweed species and communities. From there we highlight five critical avenues for further research to explore (long-term monitoring, use of functional traits, focus on early ontogeny, biotic interactions and impact of marine litter on coastal vegetation). CONCLUSIONS: Although there are considerable parallels with terrestrial vascular plant responses to the many threats posed by anthropogenic stressors, we note that the impacts of some (e.g. habitat loss) are much less keenly felt in the oceans than on land. Nevertheless, and in common with terrestrial plant communities, the impact of climate change will inevitably be the most pernicious threat to the future persistence of seaweed species, communities and service provision. While understanding macroalgal responses to simultaneous environmental stressors is inevitably a complex exercise, our attempt to highlight synergies with terrestrial systems, and provide five future research priorities to elucidate some of the important trends and mechanisms of response, may yet offer some small contribution to this goal.

Metrics matter: Multiple diversity metrics at different spatial scales are needed to understand species diversity in urban environments.

Worldwide, natural habitats are being replaced by artificial structures due to urbanisation. Planning of such modifications should strive for environmental net gain that benefits biodiversity and ecosystems. Alpha (α) and gamma (γ) diversity are often used to assess 'impact' but are insensitive metrics. We test several diversity measures across two spatial scales to compare species diversity in natural and artificial habitats. We show γ-diversity indicates equivalency in biodiversity between natural and artificial habitats, but natural habitats support greater taxon (α) and functional richness. Within-site ß-diversity was also greater in natural habitats, but among-site ß-diversity was greater in artificial habitats, contradicting the commonly held view that urban ecosystems are more biologically homogenous than natural ecosystems. This study suggests artificial habitats may in fact provide novel habitat for biodiversity, challenges the applicability of the urban homogenisation concept and highlights a significant limitation of using just α-diversity (i.e., multiple metrics are needed and recommended) for assessing environmental net gain and attaining biodiversity conservation goals.

Occurrence and assemblage composition of intertidal non-native species may be influenced by shipping patterns and artificial structures.

Habitat modification coupled with the spread of non-native species (NNS) are among the top threats to marine biodiversity globally. Species are known to be transported to new locations via international shipping and secondarily spread via regional vessels and artificial structures. Rapid Assessment Surveys (RAS) combining quantitative and semi-quantitative methods compared NNS richness and assemblage composition on intertidal natural rocky shores and artificial structures in harbours in different regions along the south coast of England. Quantitative data showed that artificial habitats supported higher richness than natural habitats, while semi-quantitative data found no difference in richness among habitat types. This result was attributed to additional species found in rock pools during searches of complex microhabitats in natural habitats. Assemblages on artificial structures differed among regions, with regions and harbours with greater numbers of vessels supporting greater richness. Results highlight the importance of shipping and artificial structures for NNS introduction and spread.

Simulated seawater flooding reduces oilseed rape growth, yield and progeny performance.

BACKGROUND AND AIMS: Of the many threats to global food security, one of the most pressing is the increased incidence of extreme weather events. In addition to extreme rainfall, a combination of global sea level rise and storm surge is likely to result in frequent episodes of seawater flooding in arable systems along low-lying coasts. Our aim was to elucidate the effects of simulated seawater and freshwater flooding on the survival, growth and reproductive potential of four cultivars of the important seed crop, Brassica napus [canola, or oilseed rape (OSR)]. METHODS: Established plants were exposed to 24 or 96 h freshwater or seawater root zone immersion (with a no immersion 'control'). Initial post-treatment performance over 7 weeks was quantified using dry weight biomass. A second group of plants, cultivated until maturity, were used to quantify reproductive yield (siliqua and seed number, and seed size) and subsequent progeny performance (germination and seedling growth). RESULTS: Oilseed rape growth and reproductive responses were unaffected by freshwater, but seawater negatively affected growth and siliqua number for all cultivars, and seed mass for two ('Agatha' and 'Cubic'). In addition to impacts on crop yield, the growth of seedlings cultivated from seed collected from maternal plants subjected to seawater immersion was also reduced. CONCLUSIONS: Our results demonstrate the potential impact of seawater inundation on coastal cropping systems; although OSR may survive acute saline flooding, there are longer term impacts on growth and yield for some cultivars. The threat may necessitate changes in land-use practice and/or the development of salt-tolerant cultivars to maintain economically viable yields. In addition, by evidencing a hitherto unknown effect on reproductive performance (i.e. reduced seed yield) and subsequent seedling growth, our study highlights an important potential impact of coastal flooding on plant community dynamics for (semi-) natural habitats.

A pinch of salt: response of coastal grassland plants to simulated seawater inundation treatments.

BACKGROUND AND AIMS: The combination of rising sea levels and increased storm frequency and intensity is predicted to increase the severity of oceanic storm surge events and the impact of flooding on coastal ecosystems globally. Understanding how plant communities respond to this threat necessitates experiments involving plant immersion in saline water, but logistical issues and natural variation in seawater composition mean that pure NaCl solutions or marine aquarium salts (MS) are widely used. Nonetheless, their comparative impact on plant ecophysiology, and thus relevance to understanding real-world flooding scenarios, is unknown. METHODS: In the first of two experiments, we examined how six ecophysiological responses in white clover (Trifolium repens) varied when plants were subjected to five different inundation treatments: deionized water, natural seawater, an MS solution and two NaCl solutions. In a second experiment, we examined how immersion in deionized water, MS solution and natural seawater affected six European perennial herb species, three native to Spanish sand dunes, and three from British coastal grasslands. RESULTS: The two NaCl solutions induced exceptional Trifolium mortality, but responses varied little between MS and seawater treatments. In the second experiment, although leaf tissue necrosis and proline concentrations increased, and growth decreased compared with untreated controls, only one response in one species varied between MS and seawater treatments. Chemical speciation modelling revealed major variation in free Na+ and Cl- between NaCl solutions and seawater, but minor differences between MS and seawater. CONCLUSIONS: We show that NaCl solutions are unsuitable surrogates to investigate plant response to elevated environmental salinity. Although responses to natural seawater and MS were consistent within species, there was notable between-species variation. Consequently, the first steps to elucidating how these species-specific responses influence coastal plant community recovery following storm surge can likely be achieved using commercial marine aquarium salts as substitutes for natural seawater.

The gathering storm: optimizing management of coastal ecosystems in the face of a climate-driven threat.

BACKGROUND: The combination of rising sea levels and increased likelihood of extreme storm events poses a major threat to our coastlines and as a result, many ecosystems recognized and valued for their important contribution to coastal defence face increased damage from erosion and flooding. Nevertheless, only recently have we begun to examine how plant species and communities, respond to, and recover from, the many disturbances associated with storm events. SCOPE: We review how the threats posed by a combination of sea level rise and storms affects coastal sub-, inter- and supra-tidal plant communities. We consider ecophysiological impacts at the level of the individual plant, but also how ecological interactions at the community level, and responses at landscape scale, inform our understanding of how and why an increasing frequency and intensity of storm damage are vital to effective coastal management. While noting how research is centred on the impact of hurricanes in the US Gulf region, we take a global perspective and consider how ecosystems worldwide (e.g. seagrass, kelp forests, sand dunes, saltmarsh and mangroves) respond to storm damage and contribute to coastal defence. CONCLUSIONS: The threats posed by storms to coastal plant communities are undoubtedly severe, but, beyond this obvious conclusion, we highlight four research priority areas. These call for studies focusing on (1) how storm disturbance affects plant reproduction and recruitment; (2) plant response to the multiple stressors associated with anthropogenic climate change and storm events; (3) the role of ecosystem-level interactions in dictating post-disturbance recovery; and (4) models and long-term monitoring to better predict where and how storms and other climate change-driven phenomena impact coastal ecosystems and services. In so doing, we argue how plant scientists must work with geomorphologists and environmental agencies to protect the unique biodiversity and pivotal contribution to coastal defence delivered by maritime plant communities.

Climate variation, reproductive frequency and acorn yield in English Oaks.

AIMS -: Annually variable, but synchronous production of large seed crops ('masting') is a widespread phenomenon in temperate trees. Mounting concerns about the impacts of anthropogenic climate change (ACC) on plant reproduction, gives urgency to our need to understand better the role of climate on tree reproduction, and in particular, mast events. Unlike our understanding of reproductive phenology however, there is little consensus regarding how climate affects plant reproductive effort, or indeed the actual environmental triggers that underpin masting behaviour. METHODS -: We used a 27-year record of acorn yield from a population of 12 Quercus robur trees located in southern England to compare masting frequency and post-dispersal acorn yield each year for each tree, with long-term weather data over the same period. We focussed on discrete or sequential climate cues (temperature, precipitation, and frost days) as likely predictors of oak reproduction. IMPORTANT FINDINGS -: Annual post-dispersal acorn crop varied greatly; i.e. no acorns in 14 of the 27 years, but there was no sequential pattern of crop versus non-crop years indicating that weather, rather than resource limitation alone, dictated the timing of reproduction. Crop years were instead most closely associated with relatively cool late summer conditions in the preceding year, followed by anomalous summer warmth within crop year. Acorn yield increased following dry April and above average May and June temperatures within crop year. Although our results support a general association between warm late spring and summer conditions, and crop frequency and yield respectively, the influence of cooler later summer conditions in the year prior to masting highlights how a combination of weather cues may dictate the occurrence of mast years. Consequently, our results corroborate not only the hypothesis that temperature differentials between consecutive years, not absolute temperatures, may be the better predictor of mast seeding events, but lend support also to the suggestion that reproductive failure and resource accumulation resulting from a climate-linked environmental veto, drives future reproductive synchronization in temperate tree species.

Riding on the wind: volatile compounds dictate selection of grassland seedlings by snails.

Background and Aims: Seedling herbivory is an important selective filter in many plant communities. The removal of preferred food plants by both vertebrate and, more commonly, invertebrate herbivores can destroy entire seedling cohorts, and consequently dictate plant community assembly. Nevertheless, our understanding of how and why some seedlings are more prone to herbivore attack than their neighbours remains limited. For seedlings, where even minor tissue damage is fatal, avoiding contact with herbivores is probably advantageous and, on this basis, volatile organic compounds (VOCs) are strong candidates to fulfil a primary defensive role. Methods: We quantified seedling selection by snails (Cornu aspersum) for 14 common, European grassland species. Seedling acceptability was subsequently compared with species-specific expression of constitutive secondary defence metabolites (CSDMs), and VOCs to determine their relative influence on seedling selection. Results: We found no relationship between seedling acceptability and CSDMs, but seedling selection was strongly associated with VOC profiles. Monoterpenes (specifically ß-ocimene) were identified as likely attractants, while green leaf volatiles (GLVs) (3-hexen-1-ol acetate) were strongly associated with low seedling acceptability. Conclusions: By elucidating a relationship between VOCs and seedling acceptability, we contradict a long-held, but poorly tested, assumption that seedling selection by herbivores in (semi-)natural plant communities centres on CSDMs. Instead, our results corroborate recent work showing how GLVs, including 3-hexen-1-ol acetate, deter crop seedling selection by molluscs. Although our failure to establish any early-ontogenetic relationship between VOCs and CSDMs also suggests that the former do not 'advertise' possession of the latter, we nevertheless reveal the role that VOCs play in defending seedlings against herbivory before lethal damage occurs.

The city as a refuge for insect pollinators.

Research on urban insect pollinators is changing views on the biological value and ecological importance of cities. The abundance and diversity of native bee species in urban landscapes that are absent in nearby rural lands evidence the biological value and ecological importance of cities and have implications for biodiversity conservation. Lagging behind this revised image of the city are urban conservation programs that historically have invested in education and outreach rather than programs designed to achieve high-priority species conservation results. We synthesized research on urban bee species diversity and abundance to determine how urban conservation could be repositioned to better align with new views on the ecological importance of urban landscapes. Due to insect pollinators' relatively small functional requirements-habitat range, life cycle, and nesting behavior-relative to larger mammals, we argue that pollinators put high-priority and high-impact urban conservation within reach. In a rapidly urbanizing world, transforming how environmental managers view the city can improve citizen engagement and contribute to the development of more sustainable urbanization.

Predicting bee community responses to land-use changes: Effects of geographic and taxonomic biases.

Land-use change and intensification threaten bee populations worldwide, imperilling pollination services. Global models are needed to better characterise, project, and mitigate bees' responses to these human impacts. The available data are, however, geographically and taxonomically unrepresentative; most data are from North America and Western Europe, overrepresenting bumblebees and raising concerns that model results may not be generalizable to other regions and taxa. To assess whether the geographic and taxonomic biases of data could undermine effectiveness of models for conservation policy, we have collated from the published literature a global dataset of bee diversity at sites facing land-use change and intensification, and assess whether bee responses to these pressures vary across 11 regions (Western, Northern, Eastern and Southern Europe; North, Central and South America; Australia and New Zealand; South East Asia; Middle and Southern Africa) and between bumblebees and other bees. Our analyses highlight strong regionally-based responses of total abundance, species richness and Simpson's diversity to land use, caused by variation in the sensitivity of species and potentially in the nature of threats. These results suggest that global extrapolation of models based on geographically and taxonomically restricted data may underestimate the true uncertainty, increasing the risk of ecological surprises.

Something in the air? The impact of volatiles on mollusc attack of oilseed rape seedlings.

BACKGROUND AND AIMS: Mounting concerns about balancing food security with the environmental impacts of agro-chemical use underpin the need to better understand the mechanisms by which crop plants, particularly during the vulnerable seedling stage, attract or repel herbivores. METHODS: The feeding preferences of the mollusc Helix aspersa were determined for several oilseed rape (Brassica napus) cultivars and a rank order of acceptability was established. This was compared with glucosinolate concentrations and volatile organic compound (VOC) profiles to determine whether seedling acceptability to molluscs was linked to either form of defence. KEY RESULTS: While VOC profiles for each oilseed rape cultivar could be separated by canonical discriminant analysis and associated with mollusc feeding preferences, glucosinolate profiles were unrelated to snail feeding behaviour. A mixture of monoterpenes (α-pinene, ß-myrcene and δ-3-carene) was identified as a putative attractant, while a blend of the green leaf volatiles 3-hexen-1-ol, 3-hexen-1-ol acetate and the monoterpene α-terpinene was identified as a putative repellent mix. Added to the VOC profile of oilseed rape seedlings, the 'repellent' mix reduced mollusc selection, while the 'attractant' mix had no effect. CONCLUSIONS: Despite the widespread assumption that seedling selection by generalist herbivores is governed by chemical defence and taste, we show that olfactory cues may be more important. Oilseed rape may be atypical of wild plants, but our ability to identify repellent volatile organic compounds that can influence snail olfactory selection points to new methods for crop protection using modified VOC profiles during the vulnerable seedling stage.

Plants and climate change: complexities and surprises.

BACKGROUND: Anthropogenic climate change (ACC) will influence all aspects of plant biology over coming decades. Many changes in wild species have already been well-documented as a result of increased atmospheric CO2 concentrations, warming climate and changing precipitation regimes. A wealth of available data has allowed the use of meta-analyses to examine plant-climate interactions on more sophisticated levels than before. These analyses have revealed major differences in plant response among groups, e.g. with respect to functional traits, taxonomy, life-history and provenance. Interestingly, these meta-analyses have also exposed unexpected mismatches between theory, experimental, and observational studies. SCOPE: We reviewed the literature on species' responses to ACC, finding ∼42 % of 4000 species studied globally are plants (primarily terrestrial). We review impacts on phenology, distributions, ecophysiology, regeneration biology, plant-plant and plant-herbivore interactions, and the roles of plasticity and evolution. We focused on apparent deviations from expectation, and highlighted cases where more sophisticated analyses revealed that unexpected changes were, in fact, responses to ACC. CONCLUSIONS: We found that conventionally expected responses are generally well-understood, and that it is the aberrant responses that are now yielding greater insight into current and possible future impacts of ACC. We argue that inconclusive, unexpected, or counter-intuitive results should be embraced in order to understand apparent disconnects between theory, prediction, and observation. We highlight prime examples from the collection of papers in this Special Issue, as well as general literature. We found use of plant functional groupings/traits had mixed success, but that some underutilized approaches, such as Grime's C/S/R strategies, when incorporated, have improved understanding of observed responses. Despite inherent difficulties, we highlight the need for ecologists to conduct community-level experiments in systems that replicate multiple aspects of ACC. Specifically, we call for development of coordinating experiments across networks of field sites, both natural and man-made.

Home advantage? Decomposition across the freshwater-estuarine transition zone varies with litter origin and local salinity.

Expected increases in the frequency and intensity of storm surges and river flooding may greatly affect the relative salinity of estuarine environments over the coming decades. In this experiment we used detritus from three contrasting environments (marine Fucus vesiculosus; estuarine Spartina anglica; terrestrial Quercus robur) to test the prediction that the decomposition of the different types of litter would be highest in the environment with which they are associated. Patterns of decomposition broadly fitted our prediction: Quercus detritus decomposed more rapidly in freshwater compared with saline conditions while Fucus showed the opposite trend; Spartina showed an intermediate response. Variation in macro-invertebrate assemblages was detected along the salinity gradient but with different patterns between estuaries, suggesting that breakdown rates may be linked in part to local invertebrate assemblages. Nonetheless, our results suggest that perturbation of salinity gradients through climate change could affect the process of litter decomposition and thus alter nutrient cycling in estuarine transition zones. Understanding the vulnerability of estuaries to changes in local abiotic conditions is important given the need to better integrate coastal proceses into a wider management framework at a time when coastlines are increasingly threatened by human activities.

Variable response of three Trifolium repens ecotypes to soil flooding by seawater.

BACKGROUND AND AIMS: Despite concerns about the impact of rising sea levels and storm surge events on coastal ecosystems, there is remarkably little information on the response of terrestrial coastal plant species to seawater inundation. The aim of this study was to elucidate responses of a glycophyte (white clover, Trifolium repens) to short-duration soil flooding by seawater and recovery following leaching of salts. METHODS: Using plants cultivated from parent ecotypes collected from a natural soil salinity gradient, the impact of short-duration seawater soil flooding (8 or 24 h) on short-term changes in leaf salt ion and organic solute concentrations was examined, together with longer term impacts on plant growth (stolon elongation) and flowering. KEY RESULTS: There was substantial Cl(-) and Na(+) accumulation in leaves, especially for plants subjected to 24 h soil flooding with seawater, but no consistent variation linked to parent plant provenance. Proline and sucrose concentrations also increased in plants following seawater flooding of the soil. Plant growth and flowering were reduced by longer soil immersion times (seawater flooding followed by drainage and freshwater inputs), but plants originating from more saline soil responded less negatively than those from lower salinity soil. CONCLUSIONS: The accumulation of proline and sucrose indicates a potential for solute accumulation as a response to the osmotic imbalance caused by salt ions, while variation in growth and flowering responses between ecotypes points to a natural adaptive capacity for tolerance of short-duration seawater soil flooding in T. repens. Consequently, it is suggested that selection for tolerant ecotypes is possible should the predicted increase in frequency of storm surge flooding events occur.

Going native? Flower use by bumblebees in English urban gardens.

BACKGROUND AND AIMS: Although urban gardens provide opportunities for pollinators in an otherwise inhospitable environment, most garden plants are not native to the recipient biogeographical region and their value to local pollinators is disputed. This study tested the hypothesis that bumblebees foraging in English urban gardens preferentially visited sympatric Palaearctic-range plants over species originating outside their native range. METHODS: Twenty-seven surveys of flower availability and bumblebee visitation (Bombus spp.) were conducted over a 3-month summer period. Plants were categorized according to whether they were native British, Palaearctic or non-Palaearctic in origin. A phylogeny of the 119 plant species recorded was constructed and the relationship between floral abundance and the frequency of pollinator visits investigated by means of phylogenetically independent contrasts. Differentiation in utilization of plant species by the five bumblebee species encountered was investigated using niche overlap analyses. KEY RESULTS: There was conflicting evidence for preferential use of native-range Palaearctic plant species by bumblebees depending on which plants were included in the analysis. Evidence was also found for niche partitioning between species based on respective preferences for native and non-native biogeographical range plants. Two bumblebees (Bombus terrestris and B. pratorum) concentrated their foraging activity on non-Palaearctic plants, while two others (B. hortorum and B. pascourum) preferred Palaearctic species. CONCLUSIONS: The long-running debate about the value of native and non-native garden plants to pollinators probably stems from a failure to properly consider biogeographical overlap between plant and pollinator ranges. Gardeners can encourage pollinators without consideration of plant origin or bias towards 'local' biogeographical species. However, dietary specialist bumblebees seem to prefer plants sympatric with their own biogeographical range and, in addition to the cultivation of these species in gardens, provision of native non-horticultural ('weed') species may also be important for pollinator conservation.

Seedling-herbivore interactions: insights into plant defence and regeneration patterns.

BACKGROUND: Herbivores have the power to shape plant evolutionary trajectories, influence the structure and function of vegetation, devastate entire crops, or halt the spread of invasive weeds, and as a consequence, research into plant-herbivore interactions is pivotal to our understanding of plant ecology and evolution. However, the causes and consequences of seedling herbivory have received remarkably little attention, despite the fact that plants tend to be most susceptible to herbivory during establishment, and this damage can alter community composition and structure. SCOPE: In this Viewpoint article we review why herbivory during early plant ontogeny is important and in so doing introduce an Annals of Botany Special Issue that draws together the latest work on the topic. In a synthesis of the existing literature and a collection of new studies, we examine several linked issues. These include the development and expression of seedling defences and patterns of selection by herbivores, and how seedling selection affects plant establishment and community structure. We then examine how disruption of the seedling-herbivore interaction might affect normal patterns of plant community establishment and discuss how an understanding of patterns of seedling herbivory can aid our attempts to restore semi-natural vegetation. We finish by outlining a number of areas where more research is required. These include a need for a deeper consideration of how endogenous and exogenous factors determine investment in seedling defence, particularly for the very youngest plants, and a better understanding of the phylogenetic and biogeographical patterns of seedling defence. There is also much still be to be done on the mechanisms of seedling selection by herbivores, particularly with respect to the possible involvement of volatile cues. These inter-related issues together inform our understanding of how seedling herbivory affects plant regeneration at a time when anthropogenic change is likely to disrupt this long-established, but all-too-often ignored interaction.

Pollination and plant defence traits co-vary in Western Australian Hakeas.

* Despite the conflicting demands imposed by mutualistic (pollination) and antagonistic (florivory) processes, the relative importance of the key selective pressures influencing floral evolution are not readily apparent. In this study we quantified a range of floral and foliar traits within the genus Hakea to investigate how pollinator and herbivore selection might influence the evolution of floral attraction and defence attributes. * Plant material was collected from populations of 51 Australian Hakea species native to southwestern Australia, and measurements were taken of foliage and inflorescence morphology, inflorescence colour and floral chemical defence. Hakeas were separated into bird- vs insect-pollinated species on the basis of stigma-nectary distance. * Our results show how the evolution of insect vs bird pollination is closely linked to whether inflorescences are protected by physical (leaf spines, dense foliage) or chemical (floral cyanide) defences, respectively. * Rather than being constrained by the necessity to attract pollinators, we suggest that pre-existing adaptations to combat florivore and herbivore attack directed the evolution of floral characteristics employed to attract pollinators and deter florivores. The inter-correlation among bird pollination, red flower colour and floral cyanide indicates floral coloration may signal to vertebrate florivores that the inflorescences are unpalatable despite their high accessibility.