Community studies in pollination ecology in the high temperate Andes of central Chile II. effect of temperature on visitation rates and pollination possibilities

Visitation rates and mean numbers of visits per flower per day are determined at three altitudinal levels (2 200–3 600 m) in the high Andes of central Chile from quantified observations of flowers visitors to a total of 134 species of plants, studied over three flowering seasons. Significant altitudinal decreases in the mean no. visits/flower/minute and per day were recorded, with Level III flowers, on the average, being pollinated only about 1/2 as frequently and to as low as 1/4 as frequently in certain months, as Level I flowers. Visitation rates are generally highest in early and mid-summer at all altitudes. — The lower visitation rates at the higher elevations are due to lower insect abundance relative to plant resources and lower levels of activity for the insects present, stemming from the generally lower ambient temperatures. Seasonal differences in visitation rates may also be related to differences in insect abundance levels. The total probability of ovule pollination cannot be estimated directly from the pollination rates, because stigmas are probably receptive for longer periods at the higher elevations. When differences in the duration of stigma receptivity are estimated from differences in flower lifelength, the probability of pollination in Level III is not very different from that expected in Level I. However, for some months, the deficit in Level III may still be substantial. — These results underline the inherent dangers of predicting pollination possibilities directly from pollination rates. They also suggest that the impoverished pollination conditions assumed in hypotheses predicting higher amounts of self-compatibility at high altitudes might not be justified for all high temperate mountains.First part of this series:Arroyo & al. (1982).     

Latitudinal patterns in the size of European butterflies

The importance of seasonality in shaping latitudinal size patterns was investigated in 16 species of lycaenine and satyrine butterflies with distributions in Sweden and continental Europe. Species without shifts in voltinism within the study area generally displayed clear size clines, increasing in size from north to south within Sweden. These species were also larger in continental Europe than in Sweden. In northern temperate areas a linear model for mean size at different latitudes seems to be a reasonable approximation for most species in this category. The size patterns for three species with shifts from univoltinism to bivoltinism at latitudes within Sweden were more reminiscent of the "saw-tooth"-pattern predicted by theory for such cases. The results suggest that size patterns in European butterflies to a large extent is governed by the length of the favorable season at different latitudes, affecting selection for development time and size. Altitude and drought also affect the effective season length and may amplify or complicate latitudinal patterns.     

Climatic warming increases voltinism in European butterflies and moths

Climate change is altering geographical ranges, population dynamics and phenologies of many organisms. For ectotherms, increased ambient temperatures frequently have direct consequences for metabolic rates, activity patterns and developmental rates. Consequently, in many insect species both an earlier beginning and prolongation of seasonal duration occurred in parallel with recent global warming. However, from an ecological and evolutionary perspective, the number of generations (voltinism) and investment into each generation may be even more important than seasonality, since an additional generation per unit time may accelerate population growth or adaptation. Using a dataset extending back to the mid-nineteenth century, I report changes in the voltinism of butterfly and moth species of Central Europe. A significant proportion of 263 multi-voltine species showed augmented frequency of second and subsequent generations relative to the first generation in a warm period since 1980, and 44 species even increased the number of generations after 1980. Expected ecological consequences are diverse. Since multi-voltinism has been linked to insect outbreaks they include an increase in the abundance of herbivorous pests of agriculture and forestry. However, disruption of the developmental synchrony associated with multi-voltinism and host plant phenology may also reduce fitness, potentially having unexpected consequences for species of conservation concern. The ability of species to adapt evolutionarily to a changing environment may be facilitated by increased voltinism.     

Comparing the efficiency of pollination mechanisms in Papilionoideae

The pump pollination mechanism is typical of basal clades within Papilionoideae and might be associated with less efficient pollen transfer systems, while the explosive tripping mechanism is considered more advanced and might represent the highest expression of the trend in pollen economy. Crotalaria pumila, C. stipularia, Desmodium incanum and D. subsericeum present secondary pollen presentation with pump and explosive pollination mechanisms, respectively. In the present study, we evaluate and compare (1) pollen removal, (2) pollen deposition and (3) pollen transfer efficiency of both mechanisms, considering single visits by Megachile spp., common pollinators of the four plant species in Salta Province, Argentina. Comparisons of visit durations are made in relation to the type of mechanism and rewards offered. We detected significant differences between both mechanisms in the proportion of pollen grains removed and deposited in a flower after a single visit of Megachile. We found that efficiency in pollen transfer was significantly higher for explosive mechanism (2.13?±?0.42 pollen grains deposited per 100 removed) than for pump mechanism (0.33?±?0.17 pollen grains deposited per 100 removed). This is the first study that compares efficiency between pollination mechanisms in this group of plants.     

Flower heating following anthesis and the evolution of gall midge pollination in Schisandraceae

? Premise of the study: Flower heating is known from a few species in 11 of the c. 450 families of flowering plants. Flowers in these families produce heat metabolically and are adapted to beetles or flies as pollinators. Here, we focus on the Schisandraceae, an American/Asian plant family known to exhibit flower heating in some species, but not others, raising the question of the adaptive function of heat production. ? Methods: We used field observations, experiments, and ancestral trait reconstruction on a molecular phylogeny for Schisandraceae that includes the investigated species. ? Key results: At least two Chinese species of Illicium are exclusively pollinated by gall midges that use the flowers as brood sites (not for pollen feeding). Continuous monitoring of flower temperatures revealed that the highest temperatures were attained after the flowers' sexual functions were over, and experiments showed that post-anthetic warming benefited larval development, not fruit development. Midge larvae in flowers with trimmed tepals (and hence a lower temperature) died, but fruit set ratios remained unchanged. Based on the DNA phylogeny, gall midge pollination evolved from general fly/beetle pollination several times in Schisandraceae, with some species adapted to flower-breeding midges, others to pollen-feeding midges. ? Conclusions: Flower heating may be an ancestral trait in Schisandraceae that became co-opted in species pollinated by flower-breeding midges requiring long-persistent warm chambers for larval development.     

Flower visitation and oviposition behavior of Frankliniella occidentalis (Tysan., Thripidae) on cucumber plants

Oviposition behaviour of western flower thrips, Frankliniella occidentalis (Pergande) on greenhouse cucumber, Cucumis satifus (L.) was investigated. Most eggs were laid in the leaves, along veins and under leaf hairs, with only a few on plant stems and flowers. Oviposition rate was higher during the day than during the night. During the day, more adult thrips were found in the flowers than during the night. The number of adult thrips per flower increased rapidly after sunrise with the highest densities occurring around noon and thereafter the number of thrips in flowers decreased during the afternoon. No differences were found in the number of larvae (first and second instars) in flowers during the same period. The number of adult thrips on male and female cucumber flowers was not different, indicating that pollen is not the only attraction in flowers for thrips.     

Adult longevity and its relationship with conservation status in European butterflies

Many European butterfly species are currently experiencing serious declines, and may be threatened with extinction. Nevertheless, due to limited knowledge on the species biology and ecology, detailed assessments of endangerment level are not possible, and instead identifying species of conservation concern has to rely on proxies. Earlier studies suggested several characteristics, including host plant specificity, overwintering stage, patch size requirements or mobility, as potentially useful indicators of butterfly species vulnerability, but the usefulness of adult longevity in this respect has not been considered so far. Based on the information gathered through an extensive literature search we investigated the relationship between adult life span, flight period length or the temporal fragmentation index calculated as the ratio of the two parameters, and conservation status of European butterflies. We found that the species classified in one of the IUCN conservation concern categories (i.e. Endangered, Near Threatened, or Vulnerable) lived shorter as adults and were characterised by higher values of the temporal fragmentation index, while there was no particular pattern concerning flight period length. We believe that the apparent effects detected reflect the fact that shorter adult life span, and thus increased temporal fragmentation, in combination with protandry, i.e. earlier emergence of males, decrease individual chances of finding mating partners. Such a situation leads to lower effective population size and reduced viability, especially in the case of small populations. All concerned, the investigated parameters reflecting adult longevity may serve as ‘early warning’ indicators, helping to flag-up butterfly species possibly at risk.     

Observation of nest visitation by multiple males in the European Nightjar Caprimulgus europeaus

ABSTRACT

We report observations of nest visitation of multiple males to three nests of European Nightjars Caprimulgus europeaus. In one case, we report the direct observation of two males apparently exhibiting parental behaviour at the same nest. In two other cases, second males visited nests. We consider the potential functions of nest visits by extra-pair males in the Nightjar.     

Similarity and specialization of the larval versus adult diet of European butterflies and moths

Many herbivorous insects feed on plant tissues as larvae but use other resources as adults. Adult nectar feeding is an important component of the diet of many adult herbivores, but few studies have compared adult and larval feeding for broad groups of insects. We compiled a data set of larval host use and adult nectar sources for 995 butterfly and moth species (Lepidoptera) in central Europe. Using a phylogenetic generalized least squares approach, we found that those Lepidoptera that fed on a wide range of plant species as larvae were also nectar feeding on a wide range of plant species as adults. Lepidoptera that lack functional mouthparts as adults used more plant species as larval hosts, on average, than did Lepidoptera with adult mouthparts. We found that 54% of Lepidoptera include their larval host as a nectar source. By creating null models that described the similarity between larval and adult nectar sources, we furthermore showed that Lepidoptera nectar feed on their larval host more than would be expected if they fed at random on available nectar sources. Despite nutritional differences between plant tissue and nectar, we show that there are similarities between adult and larval feeding in Lepidoptera. This suggests that either behavioral or digestive constraints are retained throughout the life cycle of holometabolous herbivores, which affects host breadth and identity.     

Direct costs of forest reproduction,bee-cycling and the efficiency of pollination modes

The bee guild represents direct primary costs of angiosperm reproduction. Tropical flower visitors take an amount comparable to herbivores, exceeding 3% of net primary production energy. Therefore herbivory and aboveground net primary production have been underestimated. Comparing pollinators to other herbivores, harvest in mature forest by tropical bees is greater than leafcutter ants, game animals, frugivores, vertebrate folivores, insect defoliators excluding ants, flower-feeding birds and bats, but not soil organisms. The ratio of total aboveground net primary production to investment in pollen, nectar and resin used by pollinators suggests wind pollination is several times more efficient in temperate forests than is animal pollination in neotropical moist forest. Animal pollination may be favoured by habitat mosaics and an unpredictable or sparse dispersion of conspecifics — consequences of fluctuating abiotic and biotic environments. Natural selection evidently favours diminished direct reproductive costs in forests, for example by wind pollination, regardless of latitude and disturbance regime. An example is “wind pollination by proxy” of dominant trees in seasonal southeast Asian forests. They flower only occasionally and their pollen is dispersed by tiny winged insects that are primarily carried by the wind — rather than the nectar-hungry bees, bats, birds and moths used by most tropical flora. Increasing evapotranspiration is associated with greater net primary production; I show its correlation with species richness of social tropical bees across the isthmus of Panama, which may indicate increasing forest reproductive effort devoted to flowering, and its monopolization by unspecialized flower visitors in wetter and less seasonal lowland forests.     

Factors governing flower visitation patterns and quality of pollination services delivered by social and solitary bee species to coffee in central Uganda

In Uganda, information on visitation frequency of pollinator species visiting coffee flowers is absent, although such information is critical for the stability of coffee yield through the enhancement of pollination services. This study was conducted to understand the role played by local and environmental factors on the visitation intensity of coffee flowers by different bee species. Stepwise multiple regressions were used to investigate the effects of light intensity, distance to forest, foraging time of the day, coffee blooming season and abundance of coffee floral resources on the flower visitation frequency of different bee species. Results indicated contrasting responses of different bee species. The most important factors for social bees included forest distance, light intensity and the time of the day, whereas most determinant factors for solitary bee species were the length of the flowering season and the abundance of coffee floral resources. There is a need for developing habitat and landscape management strategies for the conservation of frequent native species in the vicinity of coffee fields to increase the delivery of pollination services to coffee. It is recommended to farmers to grow their coffee farms in the adjacent of forest habitats and related semi‐natural habitats to receive high bee visitations.     

Alpine biogeography of Parnassian butterflies during Quaternary climate cycles in North America

Growth of alpine glaciers during the Pleistocene had profound effects on montane landscapes in North America and the organisms now inhabiting alpine ecosystems. Biogeography of this region has often been viewed as a system of sky islands despite the fact that species richness patterns deviate from a strict island biogeographic model. One explanation is that alpine species are not in equilibrium because of late Quaternary geographic range shifts. Genetic data can provide evidence of nonequilibrium dynamics and the distributional shifts that occur during glaciation events in alpine landscapes. Using mitochondrial and nuclear sequence data, we examine the evolutionary history of butterflies in the Parnassius phoebus complex. We test explicit, alternative models of the biogeographic history of Parnassius smintheus and Parnassius behrii , including an equilibrium island model, ancestral radiation and fragmentation, an expanding alpine archipelago and an alpine archipelago refuge model. Our results support the alpine archipelago refuge model, in which alpine butterflies undergo population contraction during glacial climates followed by population expansion during interglacial phases. While butterflies can disperse between distant mountain ranges during glacial periods, gene flow is rare. We find evidence of recent connectivity between California and Colorado, population expansion events following deglaciation ∼20 000 years B.P., and small population sizes during the last glacial period. An analysis of lineage splitting suggests that morphological differences in P. smintheus and P. behrii are the result of late Pleistocene divergence (∼48 000 years B.P.) with limited gene flow. Our results demonstrate that spatially complex and nonequilibrium population dynamics influence alpine diversity patterns.     

Flower herbivory and seed predation in Silene vulgaris (Caryophyllaceae): effects of pollination and phenology

Pollination and seed predation were studied in Silene vulgaris populations during two seasons, one with much lower pollinator abundance than the other. Among the pollinators, noctuid moths of the genus Hadena also acted as seed predators. Nectar-foraging female moths oviposited in flowers, and their larvae consumed flowers and seed capsules.
Despite a lower percentage of pollinated flowers in the year of low pollinator abundance, similar numbers of flowers set fruit in both years, because fewer flower buds and flowers were eaten by Hadena larvae during the year of low pollinator visitation. The number of seed capsules preyed upon was also lower in the year with low pollinator abundance, resulting in a higher seed set. The positive correlation between the percentage of pollinated flowers and the percentage of seed capsules destroyed was also observed when comparing flowers opening in different parts of the season.
Early flowering plant individuals had the same pollination success but suffered higher seed predation than late flowering ones. Selection for maximized pollination success through synchronous flowering, is probably the main reason for the compressed flowering period in 5. vulgaris , but the high level of predation early in the season may further increase the reproductive success of synchronous flowering individuals.     

Assessing the vulnerability of European butterflies to climate change using multiple criteria

Vulnerability of 100 European butterfly species to climate change was assessed using 13 different criteria and data on species distributions, climate, land cover and topography from 1,608 grid squares 30′ × 60′ in size, and species characteristics increasing the susceptibility to climate change. Four bioclimatic model-based criteria were developed for each species by comparing the present-day distribution and climatic suitability of the occupied grid cells with projected distribution and suitability in the future using the HadCM3-A2 climate scenario for 2051–2080. The proportions of disadvantageous land cover types (bare areas, water, snow and ice, artificial surfaces) and cultivated and managed land in the occupied grid squares and their surroundings were measured to indicate the amount of unfavourable land cover and dispersal barriers for butterflies, and topographical heterogeneity to indicate the availability of potential climatic refugia. Vulnerability was also assessed based on species dispersal ability, geographical localization and habitat specialization. Northern European species appeared to be amongst the most vulnerable European butterflies. However, there is much species-to-species variation, and species appear to be threatened due to different combinations of critical characteristics. Inclusion of additional criteria, such as life-history species characteristics, topography and land cover to complement the bioclimatic model-based species vulnerability measures can significantly deepen the assessments of species susceptibility to climate change.     

The evolutionary significance of dry and wet season forms in some tropical butterflies

An explanation is developed for the adaptive significance of the forms characteristic of the dry season and wet season adult generations of some tropical butterflies. The divergent phenotypes are accounted for as adaptive responses to a shift in the optimum balance between dependence on the alternative (but interdependent) strategies of active anti-predator devices and of crypsis. The seasonal polyphenisms exhibited by the satyrines Melanitis leda (Fabricius) and Orsotrioena medus (Fabricius) and the nymphalid Junonia almana (Linné) are examined in detail. The wet season forms show prominent marginal eyespot patterns which are displayed at rest and function principally in the deflection of attacks by vertebrate predators. In contrast, the dry season forms show very small or no spots and are wholly cryptic. Wet season butterflies are more active in general than those in the dry season when aestivation behaviour is often observed. The alternative phenotypes represent responses to the differences in behaviour, environment and nature of predation. Reproductive success is optimized in each season by an interaction of phenotype and behaviour. The hypothesis must be tested in detail by an investigation of the behavioural ecology and population biology of particular species. It is argued that these seasonal polyphenisms provide examples of adaptations to a repeating pattern of changing environments where the mode of selection is, in broad terms, understood.