Segundo-Ortin & Calvo provide a comprehensive overview of the many aspects of plant behavior examined to date. In our view, multiple lines of evidence make it difficult to deny plant sentience. We ...add further evidence to support the conclusion that plants are sentient organisms. As in animals, the behavior of plants can be seen and studied as an evolutionary trait, subject to and a consequence of increasing complexity in the interactions of plants with their environment. Our example is the evolution of floral behavior in Loasaceae, where complex patterns of stamen movement have co-evolved in interaction with specialized pollinators.
Floral traits are hypothesized to evolve primarily in response to selection by pollinators. However, selection can also be mediated by other environmental factors. To understand the relative ...importance of pollinator-mediated selection and its variation among trait and pollinator types, we analyzed directional selection gradients on floral traits from experiments that manipulated the environment to identify agents of selection. Pollinator-mediated selection was stronger than selection by other biotic factors (e.g., herbivores), but similar in strength to selection by abiotic factors (e.g., soil water), providing partial support for the hypothesis that floral traits evolve primarily in response to pollinators. Pollinator-mediated selection was stronger on pollination efficiency traits than on other trait types, as expected if efficiency traits affect fitness via interactions with pollinators, but other trait types also affect fitness via other environmental factors. In addition to varying among trait types, pollinator-mediated selection varied among pollinator taxa: selection was stronger when bees, long-tongued flies, or birds were the primary visitors than when the primary visitors were Lepidoptera or multiple animal taxa. Finally, reducing pollinator access to flowers had a relatively small effect on selection on floral traits, suggesting that anthropogenic declines in pollinator populations would initially have modest effects on floral evolution.
Conserving and restoring semi-natural habitat, i.e. enhancing landscape complexity, is one of the main strategies to mitigate pollinator decline in agricultural landscapes. However, we still have ...limited understanding of how landscape complexity shapes pollinator communities in both crop and non-crop habitat, and whether pollinator responses to landscape complexity vary with their association with mass-flowering crops. Here, we surveyed pollinator communities on mass-flowering leek crops and in nearby semi-natural habitat in landscapes of varying complexity. Surveys were done before and during crop bloom and distinguished between pollinators that visit the crop frequently (dominant), occasionally (opportunistic), or not at all (non-crop). Forty-seven per cent of the species in the wider landscape were also observed on leek flowers. Crop pollinator richness increased with local pollinator community size and increasing landscape complexity, but relationships were stronger for opportunistic than for dominant crop pollinators. Relationships between pollinator richness in semi-natural habitats and landscape complexity differed between groups with the most pronounced positive effects on non-crop pollinators. Our results indicate that while dominant crop pollinators are core components of crop pollinator communities in all agricultural landscapes, opportunistic crop pollinators largely determine species-richness responses and complex landscapes are local hotspots for both biodiversity conservation and potential ecosystem service provision.
Pollen plays a key role in plant reproductive biology. Despite the long history of research on pollen and pollination, recent advances in pollen‐tracking methods and statistical approaches to linking ...plant phenotype, pollination performance, and reproductive fitness yield a steady flow of exciting new insights. In this introduction to the Special Issue “Pollen as the Link Between Phenotype and Fitness,” we start by describing a general conceptual model linking functional classes of floral phenotypic traits to pollination‐related performance metrics and reproductive fitness. We use this model as a framework for synthesizing the relevant literature, highlighting the studies included in the Special Issue, and identifying gaps in our understanding and opportunities for further development of the field. The papers that follow in this Special Issue provide new insights into the relationships between pollen production, presentation, flower morphology, and pollination performance (e.g., pollen deposition onto stigmas), the role of pollinators in pollen transfer, and the consequences of heterospecific pollen deposition. Several of the studies demonstrate exciting experimental and analytical approaches that should pave the way for continued work addressing the intriguing role of pollen in linking plant phenotypes to reproductive fitness.
Status of pollinators in North America National Research Council; Division on Earth and Life Studies; Board on Agriculture and Natural Resources ...
2007, 2007-04-13
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Pollinators--insects, birds, bats, and other animals that carry pollen from the male to the female parts of flowers for plant reproduction--are an essential part of natural and agricultural ...ecosystems throughout North America. For example, most fruit, vegetable, and seed crops and some crops that provide fiber, drugs, and fuel depend on animals for pollination.This report provides evidence for the decline of some pollinator species in North America, including America's most important managed pollinator, the honey bee, as well as some butterflies, bats, and hummingbirds. For most managed and wild pollinator species, however, population trends have not been assessed because populations have not been monitored over time. In addition, for wild species with demonstrated declines, it is often difficult to determine the causes or consequences of their decline. This report outlines priorities for research and monitoring that are needed to improve information on the status of pollinators and establishes a framework for conservation and restoration of pollinator species and communities.
Pollinator service is essential for successful sexual reproduction and long-term population persistence of animal-pollinated plants, and innumerable studies have shown that insufficient service by ...pollinators results in impaired sexual reproduction ("pollen limitation"). Studies directly addressing the predictors of variation in pollinator service across species or habitats remain comparatively scarce, which limits our understanding of the primary causes of natural variation in pollen limitation. This paper evaluates the importance of pollination-related features, evolutionary history, and environment as predictors of pollinator service in a large sample of plant species from undisturbed montane habitats in southeastern Spain. Quantitative data on pollinator visitation were obtained for 191 insect-pollinated species belonging to 142 genera in 43 families, and the predictive values of simple floral traits (perianth type, class of pollinator visitation unit, and visitation unit dry mass), phylogeny, and habitat type were assessed. A total of 24,866 pollinator censuses accounting for 5,414,856 flower-minutes of observation were conducted on 510 different dates. Flowering patch and single flower visitation probabilities by all pollinators combined were significantly predicted by the combined effects of perianth type (open vs. restricted), class of visitation unit (single flower vs. flower packet), mass of visitation unit, phylogenetic relationships, and habitat type. Pollinator composition at insect order level varied extensively among plant species, largely reflecting the contrasting visitation responses of Coleoptera, Diptera, Hymenoptera, and Lepidoptera to variation in floral traits. Pollinator composition had a strong phylogenetic component, and the distribution of phylogenetic autocorrelation hotspots of visitation rates across the plant phylogeny differed widely among insect orders. Habitat type was a key predictor of pollinator composition, as major insect orders exhibited decoupled variation across habitat types in visitation rates. Comprehensive pollinator sampling of a regional plant community has shown that pollinator visitation and composition can be parsimoniously predicted by a combination of simple floral features, habitat type, and evolutionary history. Ambitious community-level studies can help to formulate novel hypotheses and questions, shed fresh light on long-standing controversies in pollination research (e.g., "pollination syndromes"), and identify methodological cautions that should be considered in pollination community studies dealing with small, phylogenetically biased plant species samples.
Current notions of "pollinator decline" and "pollination crisis" mainly arose from studies on pollinators of economic value in anthropogenic ecosystems of mid-latitude temperate regions. ...Comprehensive long-term pollinator data from biologically diverse, undisturbed communities are needed to evaluate the actual extent of the so-called "global pollination crisis." This paper analyzes the long-term dynamics of pollinator abundance in undisturbed Mediterranean montane habitats using pollinator visitation data for 65 plant species collected over two decades. Objectives are (1) to elucidate patterns of long-term changes in pollinator abundance from the perspectives of individual plant species, major pollinator groups, and the whole plant community and (2) to propose a novel methodological implementation based on combining a planned missing data design with the analytical strength of mixed effects models, which allows one to draw community-wide inferences on long-term pollinator trends in species-rich natural habitats. Probabilistic measurements ("patch visitation probability" and "flower visitation probability" per time unit) were used to assess pollinator functional abundance for each plant species on two separate, randomly chosen years. A total of 13,054 pollinator censuses accounting for a total watching effort of 2,877,039 flower-min were carried out on 299 different dates. Supra-annual unstability in pollinator functional abundance was the rule, with visitation probability to flowering patches and/or individual flowers exhibiting significant heterogeneity between years in the majority of plant species (83%). At the plantcommunity level, there was a significant linear increase in pollinator functional abundance over the study period. Probability of pollinator visitation to flowering patches and individual flowers increased due to increasing visitation by small solitary bees and, to a lesser extent, small beetles. Visitation to different plant species exhibited contrasting changes, and insect orders and genera differed widely in sign and magnitude of linear abundance trends, thus exemplifying the complex dynamics of community-wide changes in pollinator functional abundance. Results of this investigation indicate that pollinator declines are not universal beyond anthropogenic ecosystems; stress the need for considering broader ecological scenarios and comprehensive samples of plants and pollinators; and illustrate the crucial importance of combining ambitious sampling designs with powerful analytical schemes to draw reliable inferences on pollinator trends at the plant community level.
Nectar scents are thought to function as honest signals of reward used by pollinators, but this hypothesis has rarely been tested.
Using Penstemon digitalis, we examined honest signalling of the ...nectar volatile (S)‐(+)‐linalool and pollinator responses to linalool in both field and laboratory settings. Because our previous work showed that linalool emission was associated with higher female fitness and that nectar is scented with linalool, we hypothesized that linalool was an honest signal of nectar reward. To assess honesty, we measured linalool–nectar associations including nectar volume, sugar amount, concentration and production rate for inflorescences and flowers in several populations. We also assessed whether Bombus impatiens, the main pollinator of P. digitalis at our sites, can use linalool as a foraging signal. We supplemented real or artificial flowers in the field and laboratory with varying linalool–nectar combinations to measure pollinator behavioural responses.
We found that an inflorescence's linalool emissions could be used to predict nectar rewards in P. digitalis, but this was driven by indirect associations with display size rather than directly advertising more profitable flowers. For flowers within inflorescences there was also no evidence for an association between signal and reward. Field tests of bumblebee behaviour were inconclusive. However, in laboratory assays, bumblebees generally used variation in linalool emissions to choose more profitable flowers, demonstrating they can detect differences in linalool emitted by P. digitalis and associate them with reward profitability. These results suggest experiments that decouple display size, scent and reward are necessary to assess whether (and when) bees prefer higher linalool emissions. Bees preferred nectars with lower linalool concentrations when linalool flavoured the nectar solution, suggesting the potential for conflicting pressures on scent emission in the field.
Synthesis. Our results highlight the challenges of assessing function for traits important to fitness and suggest that the perception of floral signalling honesty may depend on whether pollinators use inflorescences or flowers within inflorescences when making foraging decisions. We conclude that future research on honest signalling in flowering plants, as well as its connection to phenotypic selection, should explicitly define honesty, in theoretical and experimental contexts.
Signal honesty in flowers is complex: we found that scented‐nectar is not always an honest signal in flowers of Penstemon digitalis. While inflorescences that had stronger linalool emission also produced more nectar with more sugar, we found that linalool is unlikely to predict which flowers within those inflorescences have the most nectar. The main pollinators, Bombus impatiens bumble bees, can use an association between linalool and nectar rewards to make foraging decisions, provided that the rewards are sufficiently motivating to support discriminating differences. Our results suggest that simple correlations between floral signals and rewards are unlikely to be common in nectar‐rewarding plants.
Selection leading to adaptation to interactions may generate rapid evolutionary feedbacks and drive diversification of species interactions. The challenge is to understand how the many traits of ...interacting species combine to shape local adaptation in ways directly or indirectly resulting in diversification. We used the well‐studied interactions between Lithophragma plants (Saxifragaceae) and Greya moths (Prodoxidae) to evaluate how plants and moths together contributed to local divergence in pollination efficacy. Specifically, we studied L. bolanderi and its two specialized Greya moth pollinators in two contrasting environments in the Sierra Nevada in California. Both moths pollinate L. bolanderi during nectaring, one of them–G. politella–also while ovipositing through the floral corolla into the ovary. First, field surveys of floral visitors and the presence of G. politella eggs and larvae in developing capsules showed that one population was visited only by G. politella and few other pollinators, whereas the other was visited by both Greya species and other pollinators. Second, L. bolanderi in these two natural populations differed in several floral traits putatively important for pollination efficacy. Third, laboratory experiments with greenhouse‐grown plants and field‐collected moths showed that L. bolanderi was more efficiently pollinated by local compared to nonlocal nectaring moths of both species. Pollination efficacy of ovipositing G. politella was also higher for local moths for the L. bolanderi population, which relies more heavily on this species in nature. Finally, time‐lapse photography in the laboratory showed that G. politella from different populations differed in oviposition behavior, suggesting the potential for local adaptation also among Greya populations. Collectively, our results are a rare example of components of local adaptation contributing to divergence in pollination efficacy in a coevolving interaction and, thus, provide insights into how geographic mosaics of coevolution may lead to coevolutionary diversification in species interactions.