Prey capture by the crab spider Misumena calycina (Araneae: Thomisidae)

Summary Crab spiders Misumena calycina (L.) in pasture rose Rosa carolina flowers regularly attacked bumble bees, smaller bees, and syrphid flies that visited these flowers. Attacks reached a maximum rate of over 20/h during mid morning, but only 1.6% of the most important prey item, bumble bees, were captured. The next most important food source, the most frequently taken item, syrphid flies Toxomerus marginatus (Say), were captured in 39% of the attempts. Since these flies have a biomass only 1/60th that of bumble bees, they comprised a much less important food source than did bumble bees. Spiders would obtain over 7% more food by specializing on bumble bees than by attacking all insect visitors, and as much as 20% more food at certain times of the day. However, they did not show a tendency to specialize at any time.     

Mechanisms that limit pollinator range in Ericaceae

Studies were conducted in 2001-2003 at Valdai National Park (Novgorod region) and at the Zvenigorod biological station of Moscow State University. The morphology of flowers, flowering dynamics and composition of insect visiting flowers of Ericaceae species: Andromeda polifolia, Chamaedaphne calyculata, Ledum palustre, Oxycoccus palustris, Vaccinium myrtillus, V. uliginosum, and V. vitis-idaea L. were studied. Some species of insects visiting flowers were excluded from the list of pollinators on the basis of observation on their behavior. L. palustre was visited mainly by flies where as other investigated species were visited mainly by bumblebees. In some cases bumblebees were the only visitors of the investigated plants. Mechanisms that protect flowers from flies and short-tongued solitary bees visits and ensure a best pollination by bumblebees are various among different species of Ericaceae. Efficiency of nectary protection also differs among different plant species and is defined by particularities of their habitats and flowering phenology. As far as all species of this family during the flowering are dominants in typical habitats, a competition for the pollination with species of other families in most cases is megligible. Flowering periods of V. vitis-idaea and V. myrtillus in forest ecosystems overlapped weakly. Moreover, V. myrtillus is pollinated mainly by bumblebee queens where as pollinators of V. vitis-idaea are bumblebee workers, solitary bees and horse flies. The other investigated plant species inhabit only oligotrophic peat bogs. Thery are pollinated by bumblebees but periods of flowering are not overlapped and consequently follow one after another. L. palustre and V. uliginosum flower simultaneosly but they are pollinated by different pollinators.     

Pollination systems in the cool temperate mixed coniferous and broad-leaved forest zone of Yakushima Island

Animal pollination was observed in a cool temperate mixed coniferous and broadleaved forest, and in shrubby vegetation on a mountain summit, on Yakushima Island (30.2°N, 130.3°E), to the south of Kyushu, Japan. In the mixed forest, two groups of plants were recognized: exclusively canopy-flowering species, and understory-flowering species. All of the canopy-flowering species had dish-shaped flowers or flowers without petals, were visited by opportunist insects, and most of them showed a mass-flowering pattern. Each segregated its flowering time from those of the others. On the other hand, most of the understory-flowering species had bell-or funnel-shaped flowers which were pollinated by birds or bumble bees, and showed an extended flowering pattern. Their phenological flowering series (exceptCamellia japonica that was pollinated by birds), without a break, coincided very well with the active period of a bumble bee species,Bombus ardens. In the shrubby vegetation on the mountain summit, two types of species were recognized: one type also grew in the forest, whereas the other type only grew in the shrubby vegetation. The former type of species in this vegetation was visited by a more diverse range of insects than that in the forest. In particular, species visited mainly by bumble bees in the forest attracted many opportunist insects. All but one of the species that only grew in the shrubby vegetation were visited only by opportunist insects and never by bumble bees.     

The relation between oilseed rape and pollination of later flowering plants varies across plant species and landscape contexts

Increasing cultivation of oilseed rape may have consequences for pollinators and wild plant pollination. By providing pollinating insects with pollen and nectar, oilseed rape benefits short-tongued, generalist insect species. Long-tongued bumble bee species, specialized to other flower types, may instead be negatively affected by increased competition from the generalists (e.g. due to nectar-robbing of long-tubed flowers) after oilseed rape flowering has ceased. We expected that the increased abundance of short-tongued pollinators and reduced abundance of long-tongued bumble bees in landscapes with a high proportion of oilseed rape would impact the pollination of later flowering wild plant species. In addition, we expected contrasting effects on plants pollinated by short-tongued pollinators and those pollinated by long-tongued bumble bees. We predicted that semi-natural grasslands, which provide insects with alternative floral resources, would reduce both negative and positive effects on pollination by mitigating competition between pollinators.In 16 semi-natural grasslands, surrounded by agricultural landscapes, with a variation in both the proportion of oilseed rape and the proportion of semi-natural grassland within 1 km, we studied reproductive output in two species of potted plants with different pollination strategies: the woodland strawberry (Fragaria vesca) and red clover (Trifolium pratense). The first species is mainly pollinated by short-tongued pollinators, e.g. hoverflies and solitary bees, and the latter by long-tongued bumble bees. Both species flowered after oilseed rape.Strawberry weight was higher in landscapes with a high proportion of oilseed rape, but only in landscapes with a low proportion of semi-natural grassland. The proportion of developed achenes was also positively related to the proportion of oilseed rape, but only during the latest flowering period. In contrast, red clover seed set was unrelated to the proportion of oilseed rape. Whereas the discrepancy between the two strawberry measurements calls for further research, this study suggests that oilseed rape can affect later flowering plants and that the impact differs among species.     

Pollination biology in the Snowy Mountains of Australia: Comparisons with montane Colorado,USA

Various aspects of the pollination biology of the alpine flora of Kosciusko National Park, NSW, were examined from late December 1983 until the end of March 1984, including flowering phenology, corolla tube lengths, flower colour, ultraviolet reflectance patterns, visitation rates to the flowers and proboscis lengths of the flower-visiting insects. An average of 5.3 species flowered in each of 13, 2 m×2 m montane plots and 5.6 species in the 13 alpine plots. The maximum number in flower simultaneously averaged 4.1 species in the montane and 3.3 in the alpine plots; flowering peaked in mid-January, Corolla tube lengths of the flora averaged 1.73 mm. The most common floral colour was white or predominantly white (40 species), followed by yellow (14 species). Only six of the 38 species (16%) examined had some type of reflectance pattern; the remaining species all absobed ultraviolet. Flies appeared to be the major pollinators. The insects collected in the study area comprised 60 species of Diptera, 33 species of Hymenoptera, and several species each of Lepidoptera and Coleoptera. On average, 14.4% of flowers watched during 379 observation periods (10 min each) were visited. On average, each plant species was visited by 6.4 species of flies, 2.4 species of bees, wasps or sawflies, one species of butterfly or moth and 0.3 species of beetles. Visitation rates increased over the growing season, and were significantly affected by ambient temperature (positively), light levels (positively) and wind speed (negatively). The maximum proboscis length for the 25 most common species of bees was 2.76 mm, but 18 of 51 species of flies had proboscis lengths longer than this. The mean proboscis length for all 25 species of bees was 1.68 mm, and for 51 species of flies was 2.31 mm. Proboscis lengths for flies were positively correlated with the average corolla length for the plant species they visited. For bees, however, the range in proboscis lengths was relatively small and did not show this pattern. There appear to be significant differences between the plant-pollinator community of alpine Australia and other alpine areas where bumblebees are common pollinators. These differences include shorter proboscis and corolla tube lengths, and perhaps an increased diversity and significance of flies as pollinators.     

The Potential Influence of Bumble Bee Visitation on Foraging Behaviors and Assemblages of Honey Bees on Squash Flowers in Highland Agricultural Ecosystems

Bee species interactions can benefit plant pollination through synergistic effects and complementary effects, or can be of detriment to plant pollination through competition effects by reducing visitation by effective pollinators. Since specific bee interactions influence the foraging performance of bees on flowers, they also act as drivers to regulate the assemblage of flower visitors. We selected squash (Cucurbita pepo L.) and its pollinators as a model system to study the foraging response of honey bees to the occurrence of bumble bees at two types of sites surrounded by a high amount of natural habitats (≥ 58% of land cover) and a low amount of natural habitats (≤ 12% of land cover) in a highland agricultural ecosystem in China. At the individual level, we measured the elapsed time from the departure of prior pollinator(s) to the arrival of another pollinator, the selection of honey bees for flowers occupied by bumble bees, and the length of time used by honey bees to explore floral resources at the two types of sites. At the community level, we explored the effect of bumble bee visitation on the distribution patterns of honey bees on squash flowers. Conclusively, bumble bee visitation caused an increase in elapsed time before flowers were visited again by a honey bee, a behavioral avoidance by a newly-arriving honey bee to select flowers occupied by bumble bees, and a shortened length of time the honey bee takes to examine and collect floral resources. The number of overall bumble bees on squash flowers was the most important factor explaining the difference in the distribution patterns of honey bees at the community level. Furthermore, decline in the number of overall bumble bees on the squash flowers resulted in an increase in the number of overall honey bees. Therefore, our study suggests that bee interactions provide an opportunity to enhance the resilience of ecosystem pollination services against the decline in pollinator diversity.     

Floral color change in Weigela middendorffiana (Caprifoliaceae): reduction of geitonogamous pollination by bumble bees

We examined the significance of retaining color-changed flowers in pollination success of Weigela middendorffiana through a single visit of bumble bees. Inner parts of flowers changed color with age from yellow to red. In an investigation of the mating system, duration of each color phase, reproductive ability of each of the color-phase flowers, and the effects of color-changed flowers on bumble bee behavior (1) flowers of this species were self-incompatible, (2) color-changed flowers provided little reward to pollinators and little residual reproductive ability, (3) the timing of floral color change was delayed with the progress of flowering season within individual plants, while the duration of the red phase shortened with the progress of flowering season, and (4) red-phase flowers did not attract bumble bees at a distance but did contribute to reducing the number of successive flower visits during a single stay within the plants. Red-phase flowers seemed to indicate the low reward level of old flowers and functioned as a cue to discourage pollinators from staying longer on the same plant. Our results predict that the retention of color-changed flowers without sexual function can enhance the pollination success of a whole plant through male function by reducing successive flower visits during a single stay of pollinators, i.e., geitonogamous pollination.     

A theoretical basis for the study of predatory syrphid fly ecology

Predacious syrphid fly species, also known as flower flies or hover flies, are cosmopolitan diptera that play two important ecological roles: predator and pollinator. In decades past, syrphid flies were studied by agricultural researchers due to their larvae’s ability to function as a biological control agent. In recent years, the global decline in both honey bees and various important wild bee species has led ecological researchers to investigate the role of syrphid fly pollination in both natural systems and agriculture. While these two roles have often been considered separately, they are rarely considered together in single studies. Syrphid fly population fluctuations in natural and agricultural systems are understudied, prompting calls for further study into the fundamental drivers of population dynamics of syrphid communities. In order to develop a deeper understanding of the fundamental dynamics of syrphid ecology, the present study offers a community model where both syrphid predation and pollination are incorporated into a single dynamic model. Using populations of predacious syrphid flies, herbivorous insects, and a shared resource flowering plant, the model is used to investigate community dynamics and persistence across different levels of plant reproductive dependence on syrphid pollination. Results indicate distinct levels of community viability across different pollination relationships as well as a tendency toward chaotic dynamics inherent to the trophic interactions of the community.     

The influence of distinct pollinators on female and male reproductive success in the Rocky Mountain columbine

Although there are many reasons to expect distinct pollinator types to differentially affect a plant's reproductive success, few studies have directly examined this question. Here, we contrast the impact of two kinds of pollinators on reproductive success via male and female functions in the Rocky Mountain columbine, Aquilegia coerulea . We set up pollinator exclusion treatments in each of three patches where Aquilegia plants were visited by either day pollinators (majority bumble bees), by evening pollinators (hawkmoths), or by both (control). Day pollinators collected pollen and groomed, whereas evening pollinators collected nectar but did not groom. Maternal parents, potential fathers and progeny arrays were genotyped at five microsatellite loci. We estimated female outcrossing rate and counted seeds to measure female reproductive success and used paternity analysis to determine male reproductive success. Our results document that bumble bees frequently moved pollen among patches of plants and that, unlike hawkmoths, pollen moved by bumble bees sired more outcrossed seeds when it remained within a patch as opposed to moving between patches. Pollinator type differentially affected the outcrossing rate but not seed set, the number of outcrossed seeds or overall male reproductive success. Multiple visits to a plant and more frequent visits by bumble bees could help to explain the lack of impact of pollinator type on overall reproductive success. The increase in selfing rate with hawkmoths likely resulted from the abundant pollen available in experimental flowers. Our findings highlighted a new type of pollinator interactions that can benefit a plant species.     

Pollination of Euphorbia dendroides by lizards and insects: Spatio-temporal variation in patterns of flower visitation

The patterns of flower visitation by lizards (Podarcis lilfordi, Lacertidae) and insects (mainly flies, bees and wasps) on the shrub Euphorbia dendroides, were studied in the island of Cabrera (Balearic Islands) during the flowering seasons of 1995 and 1996. Lizards act as true pollinators of the plant, moving large quantities of pollen within and among shrubs. To our knowledge, this is the first time that pollination by lizards has been empirically demonstrated. Variation in the quantitative component of pollination (frequency of visits × flower visitation rate) by the two groups of pollinators (lizards and insects) is documented at both spatial (within a plant population) and temporal scales (throughout the flowering season and between seasons). Variation in lizard density on a small spatial scale (within c. 200 m), presumably due to differences in vegetation cover, strongly affected their frequency of flower visitation. Insects were rather scarce, mainly because the plant flowers at a time (mid-March) when temperatures are still low. At the site where lizards were abundant, their frequency of flower visits was more than 3 times that of insects, they stayed on the shrubs about 3 times longer and visited about 8 times more cyathia per minute than did insects. Fruit and seed set were greater at this site, and this is attributed to the different frequency of flower visits by lizards, as shrubs are similar in size and produce similar amounts of cyathia in the two sites compared. Both, lizards and insects went more frequently to plants with large flower crops. However, flower crop was not associated with seed viability. We found no evidence for pollinator-mediated selection on plant traits related to fitness. Received: 28 August 1996 / Accepted: 26 February 1997     

Activity and abundance of bumble bees near Crested Butte,Colorado: diel,seasonal, and elevation effects

1. We revisited bumble bee survey data collected by Pyke in 1974 (Pyke, Ecology, 63 , 555–573, 1982) to evaluate seasonal changes in abundances of bumble bees and their floral resources, diel patterns of bumble bee activity, and elevation effects on plant and bumble bee phenology. 2. Bumble bee abundance increased during summer as spring queens founded colonies that produced workers, followed by males and autumn queens. The number of plant species visited by bumble bees increased to a peak in midsummer, then declined. 3. The number of bumble bees recorded per person‐hour peaked later than the number of flowering plant species used by the bees. Few autumn queens were observed. 4. Despite species differences in emergence times of spring queens, there were no apparent phenological differences among species in worker abundances. 5. Because flowering commences later at higher elevation, abundances of workers and males are also shifted later; therefore elevational comparisons must be seasonally adjusted. 6. These analyses provide basic information about important pollinating insects, and permit future investigations of elevational shifts over time to be properly adjusted for phenological and elevation effects in survey data.     

Fruit set in perennial vetch (Vicia unijuga): reproductive system and insect role in pollination

Perennial vetch (Vicia unijuga) is a wild plant found in parts of East Asia and potentially valuable as a forage species for more extreme environments. Information on its reproductive system and pollination biology is needed for progress in domestication of the species. We characterized the reproductive system of perennial vetch as facultative xenogamy (i.e. it is largely cross‐pollinated by insects but is also self‐compatible and can self‐pollinate). There was no significant difference (P > 0.05) in the fruit set ratio between insect cross‐pollination and artificial cross‐pollination at the seed maturation stage, but natural self‐pollination was inefficient. In our study conducted on the Tibetan plateau, eight insect species (especially certain bumble bees) are identified as potential pollinators, and four other insect species belonging to the Lepidoptera and Diptera visited flowers but are unlikely to be pollinators. The flower visitation rate of wild bumble bees was 1.6–3.3 times higher than domestic honeybees, with Bombus lepidus having the highest visitation rate of 15.7 florets/min. The diurnal floret opening rhythm of perennial vetch was synchronized with diurnal activity of potential pollinators. Optimal pollination of perennial vetch would likely be achieved using wild bees, as they have behaviour characteristics and flower tripping ability necessary, and are present in sufficient numbers to be efficient pollinators of this crop. However, even with sufficiency of pollination, there remains a fundamental problem with low fruit set which requires further investigation from a plant biology perspective.     

Three wild Lithuanian strawberry species and their pollinators

The European wild strawberry species Fragaria vesca L., Fragaria viridis Weston, and Fragaria moschata Weston are sympatric and bloom concurrently, introducing the possibilities of cooperation and competition for pollinators. We studied the pollination biology of strawberries that were grown in rectangular plots at a field site near Vilnius city in Lithuania. The number of open flowers, the number of insects visiting flowers, and the plot area covered by strawberry plants were recorded. Wild strawberries were visited by solitary bees (38.5%), flies (31.4%), and other hymenopterans (21%; ants accounted for approximately 98% of this group). Coleopterans, lepidopterans and honey bees together represented less than 10% of the insects recorded on strawberries. Fragaria viridis and F. moschata flowers were more attractive to pollinators than those of F. vesca. The main visitors of F. viridis were small solitary bees; those of F. moschata were ants, while F. vesca flowers were visited by solitary bees, ants and flies in comparable numbers. We discuss competition for pollinators and the possible impact of climate fluctuation on strawberry-pollinator interactions.     

Plant interactions for pollinator visits: a test of the magnet species effect

Summary From 1985–1987, patterns of fruit and seed set were studied in a population of mayapple (Podophyllum peltatum), a clonal, self-incompatible herb found in deciduous woods in eastern North America. Mayapple flowers do not produce nectar, but depend on infrequent visits by nectar-seeking queen bumble bees for pollination. In all years female reproductive success in mayapple colonies was influenced by colony size (number of flowers), by the distance to neighbouring colonies and by proximity to lousewort plants (Pedicularis canadensis), a prolific nectar producer heavily visited by bumble bees. In all years fruit and seed set were greater in mayapple colonies <25 m from lousewort flowers than in matched colonies which were >50 m from lousewort. In 1985 and 1987 the frequency of queen bumble bee visits to flowers in colonies close to lousewort was about four times greater than to distant colonies. In 1986 I removed about 80% of lousewort flowers to test whether the enhanced fruit and seed set in mayapples close to lousewort was pollinator mediated. Mayapple colonies close to flowerless lousewort patches did not differ in fruit or seed set from matched colonies >50 m from lousewort. In contrast, mayapples close to flowering lousewort patches had greater fruit and seed set compared with distant colonies. Over all years, a larger proportion of mayapples close to flowering lousewort patches had enhanced fruit and seed set compared with colonies close to louseworts without flowers. Though rarely documented, this type of facilitative interaction between plants that are highly attractive to pollinators (magnet species), and co-flowering species that are rarely visited by pollinators, may be widespread in plant communities.     

Leaf damage by herbivores affects attractiveness to pollinators in wild radish, Raphanus raphanistrum

 We carried out two experiments to determine the effect of leaf damage on plant attractiveness to pollinators using wild radish, Raphanus raphanistrum (Brassicaceae), a self-incompatible annual herb. Pairs of plants from 36 full-sib families were grown in pots in the greenhouse. One member of each pair was damaged by Pieris rapae larvae that were allowed to remove half of the leaf area of each of the first four rosette leaves. The plants were subsequently taken out for pollinator observations once a week from the beginning of flowering in late June until the end of August. We conducted two experiments to examine how foliar damage affected visitation by pollinators. In the first experiment, numbers of pollinators visiting plants were compared between damaged and control sibling plants. In the second experiment, the number of open flowers during observations was controlled to be the same for both damaged and undamaged sibs. Damage significantly decreased the number and size of flowers during the first observations in late June. Damaged plants received fewer visits by native bees during the first week of observations. Since damage did not affect native bee visits when the number of open flowers was equalized between treatments, flower number was probably the main cue attracting native bees to plants. In the experiment without flower number control, syrphid flies, the other abundant pollinator taxon, spent more time per flower on the undamaged than on the damaged plants. When flower number was controlled, flies probed significantly more flowers during each visit on the undamaged than on the damaged plants and had higher visitation rates to undamaged plants early in the season. Since syrphid flies preferred undamaged plants both with and without flower number control, they apparently used cues apart from flower number for visitation. The difference between undamaged and damaged plants in floral characteristics and pollinator visitation vanished within a few weeks after the start of flowering. This result suggests that early damage may not have a strong fitness effect through reduction in mating success. However, poor weather conditions can cause early mortality of plants in the field, and nutrient depletion and competition decrease fruit set of later flowers. Therefore, conditions exist under which visitation to early flowers may affect plant fitness. Received: 30 July 1996 / Accepted: 10 February 1997     

Pollination ecology and breeding system of the very narrow coastal endemic Seseli farrenyi (Apiaceae). Effects of population fragmentation

The reproductive biology of Seseli farrenyi (Apiaceae), a very narrow endemic to Cape Creus (Catalonia, Spain), including flowering timing patterns, quantity and quality of pollination services (type and frequency of pollinators, pollen carryover, pollen deposition on stigmas and reproductive success measured as fruit set), and breeding system was studied. Given the decline of population size detected in the last twenty years, we also analyzed the effects of fragmentation on pollination mechanisms. Protandry along with strong synchrony of floral development within umbels and sequential inflorescence emission within individual stalks, produces sexual phase alternation that promotes a strong outcrossing despite its non-specific pollination system and its (at least partial) self-compatibility. This pronounced xenogamy is supported by results of the insect exclusion test, hand-pollination experiments, and high P/O ratio. S. farrenyi flowers received visits from at least 28 species of insects, including wasps, small bees, ants, flies, syrphid flies, beetles and stink bugs, with different pollen carry-overs. Heterospecific pollen on stigmas decreased notably during the season (50% to 2.5%), averaging 12%. In the small population the stigmatic pollen loads and seed set decreased, but there was no effect of pollinator visitation rates. It was more affected by the composition of pollinators and their efficiency. The wind had a considerable effect on the plant. Some conservation measures are proposed.     

Are pollination syndromes useful predictors of floral visitors in Tasmania?

Abstract Diurnal visitors to the flowers of many native plant species were identified in a wide range of Tasmanian sclerophyllous vegetation between September 1996 and April 1997. These foraging profiles were analysed to determine whether they were characteristic of various floral morphologies in predictable ways. It was found that although visitor profiles were sometimes consistent with classic pollination syndromes, these syndromes were unreliable predictors of floral visitors. Very few flowers were exclusively bird‐pollinated, and none were strictly fly‐, beetle‐, wasp‐, or butterfly‐pollinated. The majority of flowering plants were unspecialized in their morphology, and consequently hosted a diverse array of visitors. In addition, visitor profiles to congeners with similar floral morphologies, and even to conspecifics, differed between habitats. Altitude was a major factor in determining visitors, with flies being the most abundant visitors above 700 m. However, congeners in several genera of Epacridaceae, as well as the genus Correa, which differed in floral morphology also differed in visitor profiles. Tubular flowers were associated with birds, while flowers with more accessible nectar were visited by insects. The only taxa exhibiting a bee‐pollination syndrome that were largely visited by bees were the Fabaceae and Goodenia ovata Sm. Several species with purple or pink flowers were also predominantly visited by bees, but did not strictly conform to the melittophilous syndrome. In contrast, other flowers exhibiting an ostensibly mellitophilous syndrome hosted very few bees. Of these, species that occurred at high altitude were mainly visited by flies, while others received very few potential pollen vectors.     

Seasonal complementary in pollinators of soft-fruit crops

Understanding the relative contributions of wild and managed pollinators, and the functional contributions made by a diverse pollinator community, is essential to the maintenance of yields in the 75% of our crops that benefit from insect pollination. We describe a field study and pollinator exclusion experiments conducted on two soft-fruit crops in a system with both wild and managed pollinators. We test whether fruit quality and quantity is limited by pollination, and whether different pollinating insects respond differently to varying weather conditions. Both strawberries and raspberries produced fewer marketable fruits when insects were excluded, demonstrating dependence on insect pollinators. Raspberries had a short flowering season which coincided with peak abundance of bees, and attracted many bees per flower. In contrast, strawberries had a much longer flowering season and appeared to be much less attractive to pollinators, so that ensuring adequate pollination is likely to be more challenging. The proportion of high-quality strawberries was positively related to pollinator abundance, suggesting that yield was limited by inadequate pollination on some farms. The relative abundance of different pollinator taxa visiting strawberries changed markedly through the season, demonstrating seasonal complementarity. Insect visitors responded differently to changing weather conditions suggesting that diversity can reduce the risk of pollination service shortfalls. For example, flies visited the crop flowers in poor weather and at the end of the flowering season when other pollinators were scarce, and so may provide a unique functional contribution. Understanding how differences between pollinator groups can enhance pollination services to crops strengthens the case for multiple species management. We provide evidence for the link between increased diversity and function in real crop systems, highlighting the risks of replacing all pollinators with managed alternatives.     

Pollinator aggregative and functional responses to flower density: does pollinator response to patches of plants accelerate at low‐densities?

Plant reproduction is often reduced at low densities, due to reduced pollinator visitation rates. Recent theory suggests that a disproportionate increase in pollinator visits to patches of plants as heterospecific plant density increases (i.e. if visitation is a sigmoid function of patch density) can rescue sparse populations of a focal plant species from reduced reproductive success or population decline. A field experiment was performed to determine the shape of the pollinator visitation response to patches of differing density of the common weed Brassica rapa . Both the aggregative and functional response for the entire pollinator community were saturating rather than sigmoid, indicating that pollinator response does not accelerate when density increases. The results for the entire pollinator community were consistent among temporal and spatial replicates. Aggregative response curves for specific pollinator taxa were either linear (bombyliid flies) or saturating (syrphid flies, solitary bees, and Lepidoptera). Functional responses for these taxa were saturating (syrphid flies and solitary bees) or flat (bombyliid flies and Lepidoptera). Individual pollinators visited more plants during foraging bouts in high-density patches, but visits per plant decreased. Seeds per fruit and seeds per flower increased with increasing density. There is no evidence that pollinators disproportionately visit denser patches, or that the conditions for this mechanism of pollination facilitation are likely to be met in this generalist pollinator system.     

Effectiveness of ants as pollinators of<Emphasis Type="Italic"> Lobularia maritima</Emphasis>: effects on main sequential fitness components of the host plant

The effectiveness of ants as pollinators of Lobularia maritima (Cruciferae) was experimentally analyzed by assessing (1) their quantitative importance at flowers; (2) their effect on host plant seed production; (3) their effect on the performance of host plant progeny, estimated as seed germination, seedling emergence, seedling survival to flowering, and (4) the overall effect of ants on a cumulative, more realistic measure of plant fitness related to the recruitment probability. Flowers of L. maritima were visited during the 2 years of study (1996 and 1997) by more than 50 pollinator species belonging to about 30 families of disparate taxonomic affiliation, notably ants and flies. There was significant seasonal variability in insect abundance and type. Ants, especially Camponotus micans (Formicidae), visited the flowers of L. maritima in summer, representing 81.2% of the visits during this season. This ant species acted as a pollinator of L. maritima, with flowers visited exclusively by ants producing significantly more seeds than flowers from which all pollinators were excluded, whereas flowers visited by only winged insects did not differ from self-pollination. Ant-pollinated flowers produced seeds with a germination rate comparable to the other treatments. Moreover, seedlings from these seeds emerged as fast, and survived at the same rate as controls. Consequently, both ants and winged insects had similar overall effects on host plant recruitment probability (0.6 and 0.7% of initial ovules produced flowering offspring, respectively), a result similar to that of open-pollinated flowers (1.0%). This study demonstrated that the overall effectiveness of the ant C. micans as a summer pollinator of L. maritima was as high as that of other winged insects, contributing not only to the seed production of this crucifer but also to the recruitment of new flowering offspring. Received: 17 February 1999 / Accepted: 26 August 1999