Whether in ant-aphid mutualism the ants exert evolutionary selection pressure on aphid morphology has not yet been fully tested. Here, we tested whether the long proboscises of Stomaphis yanonis (Aphididae Lachninae) aphids confer an advantage in preventing predation by the tending ants. Specifically, we tested the hypothesis that aphids with a shorter proboscis would excrete less honeydew, making them more likely to be preyed upon by ants. Our results showed that aphid individuals with a shorter proboscis took up less phloem sap and excreted less honeydew than individuals with a longer proboscis. In addition, among aphids with a similar body size, those with a shorter proboscis were more susceptible to predation by ants than those with a longer proboscis. These results suggest that predation by tending ants, by exerting selection pressure on aphid proboscis morphology, has caused the aphids to evolve longer proboscises.

Morphological trait-matching and species abundance are thought to be the main factors affecting the frequency and strength of mutualistic interactions. However, the relative importance of trait-matching and species abundance in shaping species interactions across environmental gradients remains poorly understood, especially for plant-insect mutualisms involving generalist species. Here, we characterised variation in species and trait composition and the relative importance of trait-matching and species abundance in shaping plant-Hymenoptera and plant-Diptera mutualisms in four meadows across an elevational gradient (2,725-3,910 m) in Yulong Snow Mountain, Southwest China. We also evaluated the effects of morphological traits of flower visitors and plant composition on their foraging specialisation (d\' and normalised degree). There was a high degree of dissimilarity in the composition of Hymenoptera and Diptera visitors and their visited plants between communities. This variation was mainly driven by the spatial replacement of species. Both for plant-Hymenoptera and plant-Diptera networks, trait-matching between nectar tube depth and proboscis length was a stronger predictor of the interactions between temporally co-occurring plants and flower visitors than species abundance. Fourth-corner analyses revealed statistically significant trait-matching between nectar tube depth and proboscis length in plant-Hymenoptera networks at all sites, suggesting that Hymenoptera consistently foraged on plant species with nectar tube depths matching their proboscis lengths. By contrast, significant trait-matching in plant-Diptera networks was only observed at the two lower elevation sites. The species-level specialisation d\' of flower visitors increased significantly as the proboscis length and the difference in nectar tube depth between the plant community and the plants visited by flower visitors increased. Our results highlight that the importance of trait-matching in shaping pairwise interactions and niche partitioning depends on the specific features (e.g. species composition and trait availability) of the plant-pollinator system. For specialised plant-Hymenoptera systems, trait-matching is an important determinant of species interactions, whereas for generalist plant-Diptera systems, trait-matching is relatively unimportant.

Floral attributes often influence the foraging choices of nectar-feeding butterflies, given the close association between plants and these butterfly pollinators. The diversity of butterflies is known to a large extent in Nepal, but little information is available on the feeding habits of butterflies. This study was conducted along the periphery of Rupa Wetland from January to December 2019 to assess butterfly species diversity and to identify the factors influencing their foraging choices. In total, we recorded 1535 individuals of 138 species representing all six families. For our examination of butterfly-nectar plant interactions, we recorded a total of 298 individuals belonging to 31 species of butterfly visiting a total of 28 nectar plant species. Overall, total butterfly visitation was found to be significantly influenced by plant category (herbaceous preferred over woody), floral color (yellow white and purple preferred over pink), and corolla type (tubular preferred over nontubular). Moreover, there was a significant positive correlation between the proboscis length of butterflies and the corolla tube length of flowers. Examining each butterfly family separately revealed that, for four of the families (Lycaenidae, Nymphalidae, Papilionidae, and Pieridae), none of the tested factors (flower color, plant category, and corolla type) were shown to significantly influence butterfly abundance at flowers. However, Hesperidae abundance was found to be significantly influenced by both flower color (with more butterflies observed at yellow flowers than purple) and flower type (with more butterflies observed at tubular flowers than nontubular flowers). Our results reveal that Rupa Lake is a suitable habitat for butterflies, providing valuable floral resources. Hence, further detailed studies encompassing all seasons, a greater variety of plants, and other influential factors in different ecological regions are fundamental for creating favorable environments to sustain important butterfly pollinators and help create balanced wetland ecosystems.

A long-standing question in ecology is how species interactions are structured within communities. Although evolutionary theory predicts close size matching between floral nectar tube depth and pollinator proboscis length of interacting species, such size matching has seldom been shown and explained in multispecies assemblages. Here, we investigated the degree of size matching among Asteraceae and their pollinators and its relationship with foraging efficiency. The majority of pollinators, especially Hymenoptera, choose plant species on which they had high foraging efficiencies. When proboscides were shorter than nectar tubes, foraging efficiency rapidly decreased because of increased handling time. When proboscides were longer than nectar tubes, a decreased nectar reward rather than an increased handling time made shallow flowers more inefficient to visit. Altogether, this led to close size matching. Overall, our results show the importance of nectar reward and handling time as drivers of plant-pollinator network structure.

Understanding how ecological communities are structured is a major goal in ecology. Ecological networks representing interaction patterns among species have become a powerful tool to capture the mechanisms underlying plant-animal assemblages. However, these networks largely do not account for inter-individual variability and thus may be limiting our development of a clear mechanistic understanding of community structure. In this study, we develop a new individual-trait based approach to examine the importance of individual plant and pollinator functional size traits (pollinator thorax width and plant nectar holder depth) in mutualistic networks. We performed hierarchical cluster analyses to group interacting individuals into classes, according to their similarity in functional size. We then compared the structure of bee-flower networks where nodes represented either species identity or trait sets. The individual trait-based network was almost twice as nested as its species-based equivalent and it had a more symmetric linkage pattern resulting from of a high degree of size-matching. In conclusion, we show that by constructing individual trait-based networks we can reveal important patterns otherwise difficult to observe in species-based networks and thus improve our understanding of community structure. We therefore recommend using both trait-based and species-based approaches together to develop a clearer understanding of the properties of ecological networks.

Relationships between species distribution and abundance, the influence of proboscis length on species-packing, and species associations within the local assemblages were studied in local communities of bumblebees in northern Spain along an altitudinal gradient. Local species abundance and altitudinal range occupied accounted for much of the variation in species distribution. Altitudinal range occupied by species was related to species distribution, but the most important variable accounting for species distribution was the local percentage abundance. Despite this, there was no evidence for age abundance. Despite this, there was no evidence for bimodality in the distributions of species incidence. A general trends for mean proboscis length in each locality to be greater in lowland localities exists, but this variable was not related to species distribution or abundance. Proboscis length spacings were studied among species in local assemblages and in most of the cases observed spacing did not differ from random expectations. The same patterns were demonstrated calculating spacings for \'core\' species in each local assemblage. Furthermore, species showed little tendency towards associations, so it may be concluded that bumblebee assemblages were irregularly structured and no clear patterns emerged from the present study.

OBJECTIVE: Unrelated organisms that share similar niches often exhibit patterns of convergent evolution in functional traits. Based on bimodal distributions of hawkmoth tongue lengths and tubular white flowers in Africa, this study hypothesized that long-tongued hawkmoths comprise a pollination niche (ecological opportunity) that is distinct from that of shorter-tongued hawkmoths.
METHODS: Field observations, light trapping, camera surveillance and pollen load analysis were used to identify pollinators of plant species with very long-tubed (>8 cm) flowers. The nectar properties and spectral reflectance of these flowers were also measured. The frequency distributions of proboscis length for all captured hawkmoths and floral tube length for a representative sample of night-blooming plant species were determined. The geographical distributions of both native and introduced plant species with very long floral tubes were mapped.
RESULTS: The convolvulus hawkmoth Agrius convolvuli is identified as the most important pollinator of African plants with very long-tubed flowers. Plants pollinated by this hawkmoth species tend to have a very long (approx. 10 cm) and narrow flower tube or spur, white flowers and large volumes of dilute nectar. It is estimated that >70 grassland and savanna plant species in Africa belong to the Agrius pollination guild. In South Africa, at least 23 native species have very long floral tubes, and pollination by A. convolvuli or, rarely, by the closely related hawkmoth Coelonia fulvinotata, has been confirmed for 11 of these species. The guild is strikingly absent from the species-rich Cape floral region and now includes at least four non-native invasive species with long-tubed flowers that are pre-adapted for pollination by A. convolvuli.
CONCLUSIONS: This study highlights the value of a niche perspective on pollination, which provides a framework for making predictions about the ecological importance of keystone pollinators, and for understanding patterns of convergent evolution and the role of floral traits in plant colonization.

Mutualistic relationships between plants and their pollinators have played a major role in the evolution of biodiversity. While the vulnerability of these relationships to environmental change is a major concern, studies often lack a framework for predicting impacts from emerging threats (e.g. biological invasions). The objective of this study was to determine the reliance of Platanthera ciliaris (orange-fringed orchid) on Papilio palamedes (Palamedes swallowtail butterfly) for pollination and the relative availability of alternative pollinators. Recent declines of P. palamedes larval host plants due to laurel wilt disease (LWD) could endanger P. ciliaris populations that rely heavily on this butterfly for pollination. We monitored pollinator visitation and fruit set and measured nectar spur lengths of P. ciliaris flowers and proboscis lengths of its floral visitors in Jackson County, MS, USA. Papilio palamedes was the primary visitor with minimal visitation by Phoebis sennae (cloudless sulfur butterfly). Lengths of P. ciliaris nectar spurs were similar to proboscis lengths of both pollinator species. Fruit set was moderate with access to pollinators (55 ± 10.8 %), yet failed (0 %) when pollinators were excluded. Visitation increased with inflorescence size, but there was no such pattern in fruit set, indicating that fruit set was not limited by pollinator visitation within the range of visitation rates we observed. Our results are supported by historical data that suggest P. palamedes and P. sennae are important pollinators of P. ciliaris. Although P. sennae may provide supplemental pollination service, this is likely constrained by habitat preferences that do not always overlap with those of P. cilaris. Observed declines of P. palamedes due to LWD could severely limit the reproductive success and persistence of P. ciliaris and similar orchid species populations. This empirical-based prediction is among the first to document exotic forest pests and pathogens as an indirect threat to plant-pollinator interactions.

OBJECTIVE: Plant-pollinator interactions are thought to have shaped much of floral evolution. Yet the relative importance of pollinator shifts and coevolutionary interactions for among-population variation in floral traits in animal-pollinated species is poorly known. This study examined the adaptive significance of spur length in the moth-pollinated orchid Platanthera bifolia.
METHODS: Geographical variation in the length of the floral spur of P. bifolia was documented in relation to variation in the pollinator fauna across Scandinavia, and a reciprocal translocation experiment was conducted in south-east Sweden between a long-spurred woodland population and a short-spurred grassland population.
RESULTS: Spur length and pollinator fauna varied among regions and habitats, and spur length was positively correlated with the proboscis length of local pollinators. In the reciprocal translocation experiment, long-spurred woodland plants had higher pollination success than short-spurred grassland plants at the woodland site, while no significant difference was observed at the grassland site.
CONCLUSIONS: The results are consistent with the hypothesis that optimal floral phenotype varies with the morphology of the local pollinators, and that the evolution of spur length in P. bifolia has been largely driven by pollinator shifts.

OBJECTIVE: Pollinator landscapes, as determined by pollinator morphology/behaviour, can vary inter- or intraspecifically, imposing divergent selective pressures and leading to geographically divergent floral ecotypes. Assemblages of plants pollinated by the same pollinator (pollinator guilds) should exhibit convergence of floral traits because they are exposed to similar selective pressures. Both convergence and the formation of pollination ecotypes should lead to matching of traits among plants and their pollinators.
METHODS: We examined 17 floral guild members pollinated in all or part of their range by Prosoeca longipennis, a long-proboscid fly with geographic variation in tongue length. Attractive floral traits such as colour, and nectar properties were recorded in populations across the range of each species. The length of floral reproductive parts, a mechanical fit trait, was recorded in each population to assess possible correlation with the mouthparts of the local pollinator. A multiple regression analysis was used to determine whether pollinators or abiotic factors provided the best explanation for variation in floral traits, and pollinator shifts were recorded in extralimital guild member populations.
RESULTS: Nine of the 17 species were visited by alternative pollinator species in other parts of their ranges, and these displayed differences in mechanical fit and attractive traits, suggesting putative pollination ecotypes. Plants pollinated by P. longipennis were similar in colour throughout the pollinator range. Tube length of floral guild members co-varied with the proboscis length of P. longipennis.
CONCLUSIONS: Pollinator shifts have resulted in geographically divergent pollinator ecotypes across the ranges of several guild members. However, within sites, unrelated plants pollinated by P. longipennis are similar in the length of their floral parts, most probably as a result of convergent evolution in response to pollinator morphology. Both of these lines of evidence suggest that pollinators play an important role in selecting for certain floral traits.