Habitat fragmentation leads to reduced pollinator visitation,fruit production and recruitment in urban mangrove forests

Mangrove forests worldwide undergo anthropogenic fragmentation that may threaten their existence, and yet there have been few tests of the effects of fragmentation on demographic processes critical for mangrove regeneration. Predicting the effects of habitat fragmentation on mangroves is problematic as pollinators may move more freely across water than terrestrial habitat, and propagules can be widely dispersed by water. Here, within each of two estuaries, we compared pollinator diversity and activity, reproductive effort and output, and rates of recruitment for sets of three large (>1500 trees), medium (300–500) and small (<50) stands. As predicted, most measures of reproductive activity and success were inversely related to stand size with large stands typically producing significantly more and larger fruit, and significantly more seedlings. Most strikingly, we found the effect of fragmentation on the abundance of pollinators (honeybees), the production and quality of fruit and the survival rate of seedlings to be similar, showing significant reduction of recruitment in small stands. This study provides the first rigorous evidence that recruitment of mangroves, like for many terrestrial plants, is negatively impacted by habitat fragmentation. From a management perspective, we argue that in the short term our data imply the importance of conserving the largest possible stands. However, additional work is needed to determine (1) the proportion of recruits within small stands that originate within large stands, (2) how seedling performance varies with fruit size and genotype, and (3) how seedling size and performance vary with the abundance and diversity of pollen.     

Pollinator shift to managed honeybees enhances reproductive output in a bumblebee-pollinated plant

Introduced honeybees have had a large impact on native ecosystems by disrupting native plant–pollinator interactions. However, little is known of the effect of honeybees on reproduction of bumblebee-pollinated plants. Seasonal displacement of native bumblebees by introduced honeybees (Apis mellifera and A. cerana) was observed in Pedicularis densispica, endemic to Hengduan Mountains, China, providing an opportunity for honeybee presence/absence comparisons. Five-year field surveys were conducted in one frequently disturbed population at Yila Pasture (YP). We compared pollination effectiveness (combinations of visitation rate, efficiency in pollen transfer, and potential geitonogamy) between native and introduced managed bees. The total visitation rate of native bees and subsequent reproductive output decreased progressively, but honeybee introduction resulted in at least twofold increase in visitation and 70 % increase in seed set. In general, native bumblebees, which have larger bodies and longer proboscises and spent more time probing single flowers, were more efficient than honeybees in terms of pollen removal and pollen deposition during first visits to virgin flowers. Compared with bumblebees, honeybees visited markedly fewer flowers in sequence within individual plants, potentially reducing geitonogamous pollination. Our data highlight that introduced honeybees can provide pollination service in terms of both quantity and quality for P. densispica. We suggest honeybee introduction as an effective way to augment pollination of P. densispica at disturbed and isolated sites.     

Pollination of invasive Eichhornia crassipes (Pontederiaceae) by the introduced honeybee (Apis mellifera L.) in South China

Heterostyly functions as an outcrossing mechanism facilitating accurate pollen transfer from anthers to stigmas of particular heights as a result of the behavior of specialist pollinators. However, heterostylous plants are also visited by generalist pollinators, which may affect the plant–pollinator mutualism. Eichhornia crassipes is a tristylous invasive species, with only the mid- and long-styled morphs (M and L) found in China. We recorded flower-visiting insects in Zhuhai, Zhongshan and Nanning in South China. We hand-pollinated the two morphs to determine their compatibility. In addition, by allowing controlled insect pollination in artificial isoplethically monomorphic and bimorphic populations, we undertook a detailed analysis of pollen deposition between the floral morphs, and fruit and seed set. Ranked by relative abundance, the flower-visiting insects were: Apis mellifera, A. cerana, Lasioglossum sp. and Eristalis arvorum. Hand pollination showed that both the M and L morphs were self-compatible, but the former was probably more so than the latter. Intra-morph pollen transfer by A. mellifera within a population was significantly greater than legitimate pollen transfer between populations, suggesting that the pollen exchange between populations was limited. Seed set of the L morph was significantly greater than that of the M morph in monomorphic populations, indicating intra-morph pollen deposition in the former was higher than in the latter. The results showed that A. mellifera was the major pollinator in South China and able to pollinate E. crassipes legitimately and to promote its fruit and seed set, even though high levels of intra-morph pollination occurred.     

Does the surrounding matrix influence corridor effectiveness for pollen dispersal in farmland?

Recent decades have seen a shift in agricultural land use from pasture to arable combined with increased use of fertilisers and pesticides. In this quite hostile landscape matrix, pollinator movements between native vegetation remnants may be impeded. Linear landscape elements (LLEs) in farmland can function as biological corridors by facilitating pollinator movements and pollen flow between fragmented plant populations. The type of landscape matrix surrounding LLEs and plant populations, and LLE habitat quality may influence the effectiveness of LLEs as corridors for pollen dispersal through the availability of floral resources and nesting opportunities for pollinators. Using fluorescent dyes as pollen analogues, we investigated dye dispersal patterns between fragmented populations of the rare insect-pollinated Primula vulgaris connected by existing LLEs. We examined how dye deposition on P. vulgaris and within LLEs (on other co-flowering insect-pollinated species) could be influenced by the surrounding matrix type (pasture, arable field), the recipient population traits of P. vulgaris (flowering population size, flower display, flowering plant density and co-flowering floral resources) and by LLE traits (LLE length and co-flowering floral resources). Dye dispersal through corridors was significantly higher when the landscape matrix surrounding P. vulgaris recipient populations and LLEs consisted of pastures rather than arable fields (or a mix of both), even after accounting for differences in floral resources. A higher cover and diversity of insect-pollinated plants increased dye deposition when co-flowering within small P. vulgaris populations, but led to dye loss within LLEs. Large P. vulgaris populations appeared more attractive thereby increasing heterospecific dye deposition in the LLEs. Our study shows that farming practices shifting from pastures to maize arable fields have a negative impact on dye dispersal patterns, and so possibly affect pollen dispersal of P. vulgaris, likely through a reduced pollinator service. Corridor effectiveness depends on the type of surrounding matrix, and the abundance of floral resources directly influences dye dispersal patterns. Preserving the remaining pastureland is essential, but restoring rich flowering vegetation at field boundaries and along LLEs may also promote corridor effectiveness for pollen dispersal.     

Infectivity of DWV Associated to Flower Pollen: Experimental Evidence of a Horizontal Transmission Route

Deformed wing virus (DWV) is a honeybee pathogen whose presence is generally associated with infestation of the colony by the mite Varroa destructor, leading to the onset of infections responsible for the collapse of the bee colony. DWV contaminates bee products such as royal jelly, bee-bread and honey stored within the infected hive. Outside the hive, DWV has been found in pollen loads collected directly from infected as well as uninfected forager bees. It has been shown that the introduction of virus-contaminated pollen into a DWV-free hive results in the production of virus-contaminated food, whose role in the development of infected bees from virus-free eggs has been experimentally demonstrated. The aim of this study was twofold: (i) to ascertain the presence of DWV on pollen collected directly from flowers visited by honeybees and then quantify the viral load and (ii) determine whether the virus associated with pollen is infective. The results of our investigation provide evidence that DWV is present on pollen sampled directly from visited flowers and that, following injection in individuals belonging to the pollinator species Apis mellifera, it is able to establish an active infection, as indicated by the presence of replicating virus in the head of the injected bees. We also provide the first indication that the pollinator species Osmia cornuta is susceptible to DWV infection.     

Floral neighborhood influences pollinator assemblages and effective pollination in a native plant

Pollinators represent an important intermediary by which different plant species can influence each other’s reproductive fitness. Floral neighbors can modify the quantity of pollinator visits to a focal species but may also influence the composition of visitor assemblages that plants receive leading to potential changes in the average effectiveness of floral visits. We explored how the heterospecific floral neighborhood (abundance of native and non-native heterospecific plants within 2 m × 2 m) affects pollinator visitation and composition of pollinator assemblages for a native plant, Phacelia parryi. The relative effectiveness of different insect visitors was also assessed to interpret the potential effects on plant fitness of shifts in pollinator assemblage composition. Although the common non-native Brassica nigra did not have a significant effect on overall pollinator visitation rate to P. parryi, the proportion of flower visits that were made by native pollinators increased with increasing abundance of heterospecific plant species in the floral neighborhood other than B. nigra. Furthermore, native pollinators deposited twice as many P. parryi pollen grains per visit as did the nonnative Apis mellifera, and visits by native bees also resulted in more seeds than visits by A. mellifera. These results indicate that the floral neighborhood can influence the composition of pollinator assemblages that visit a native plant and that changes in local flower communities have the potential to affect plant reproductive success through shifts in these assemblages towards less effective pollinators.     

The presence of the invasive plant Solanum elaeagnifolium deters honeybees and increases pollen limitation in the native co-flowering species Glaucium flavum

Invasive plants can impact biodiversity and ecosystem functioning by displacing native plants and crop species due to competition for space, nutrients, water and light. The presence of co-flowering invasives has also been shown to affect some native plants through the reduction in pollinator visitation or through the deposition of heterospecific pollen on the native’s stigmas leading to stigma clogging. We examined the impact of the invasive plant Solanum elaeagnifolium Cavanilles (silver-leafed nightshade), native to South and Central America and South-western parts of North America, on the seed set of the native Glaucium flavum Crantz (yellow-horned poppy) on Lesvos Island, Greece. To do this we measured seed set and visitation rates to G. flavum before and after the placement of potted individuals of the invasive near the native plants. In addition, we hand-crossed G. flavum flowers with super-optimal amounts of conspecific pollen, bagged flowers to measure the rate of spontaneous selfing, and applied self-pollen to measure self-compatibility of G. flavum. The hand-selfing treatment resulted in very low seed set, which indicates that G. flavum is to a large degree self-incompatible and highlights the plant’s need for insect-mediated outcrossing. We show that the presence of the invasive significantly enhanced pollen limitation, although the overall visitation rates were not reduced and that this increase is due to a reduction in honeybee visitation in the presence of the invasive resulting in reduced pollination.     

High invasive pollen transfer, yet low deposition on native stigmas in a Carpobrotus-invaded community

Background and Aims

Invasive plants are potential agents of disruption in plant–pollinator interactions. They may affect pollinator visitation rates to native plants and modify the plant–pollinator interaction network. However, there is little information about the extent to which invasive pollen is incorporated into the pollination network and about the rates of invasive pollen deposition on the stigmas of native plants.

Methods

The degree of pollinator sharing between the invasive plant Carpobrotus affine acinaciformis and the main co-flowering native plants was tested in a Mediterranean coastal shrubland. Pollen loads were identified from the bodies of the ten most common pollinator species and stigmatic pollen deposition in the five most common native plant species.

Key Results

It was found that pollinators visited Carpobrotus extensively. Seventy-three per cent of pollinator specimens collected on native plants carried Carpobrotus pollen. On average 23 % of the pollen on the bodies of pollinators visiting native plants was Carpobrotus. However, most of the pollen found on the body of pollinators belonged to the species on which they were collected. Similarly, most pollen on native plant stigmas was conspecific. Invasive pollen was present on native plant stigmas, but in low quantity.

Conclusions

Carpobrotus is highly integrated in the pollen transport network. However, the plant-pollination network in the invaded community seems to be sufficiently robust to withstand the impacts of the presence of alien pollen on native plant pollination, as shown by the low levels of heterospecific pollen deposition on native stigmas. Several mechanisms are discussed for the low invasive pollen deposition on native stigmas.Key words: Alien plant, Carpobrotus aff. acinaciformis, competition for pollinators, invasion, Mediterranean shrubland, plant-pollinator network, pollen loads, pollinator visits, stigma     

Pollination ecology of a plant in its native and introduced areas

Entomophilous and obligate out-crossing non-native plants need to become well integrated in the resident plant–pollinator network to set seeds and become established. However, it is largely unknown how pollination patterns differ between native ranges and those where plants have been introduced.We compared the identity, abundance and visitation rates of pollinators, insect pollen loads, pollen deposition on stigmas, and fruit and seed sets of Hedysarum coronarium, an entomophilous short lived N-fixing perennial, in populations from native and introduced ranges in Spain (South of mainland Spain and Menorca Island, respectively).In both areas, Hedysarum was visited by a similar number of species, mainly hymenopterans; seven species were common between native and introduced areas. However, pollinator richness, abundance, and visits per flower were greater in the native than in the introduced range, as were fruit and seed sets. Hedysarum pollen loads on stigmas and on Apis mellifera, the most common pollinator, did not differ between areas. Lower abundance of pollinators might be causing lower visitation rates, and to some extent reducing Hedysarum fruit and seed sets in the introduced area.Our biogeographical approach shows that integration of a non-native plant in a resident pollinator network does not prevent pollen limitation in the introduced area. Therefore, despite being necessary, pollination mutualistic relationships might not be the key for non-native plant establishment success in the introduced area.     

Can the pollination biology and floral ontogeny of the threatened Acacia carneorum explain its lack of reproductive success?

Many Acacia species in arid areas of eastern Australia have been severely impacted by grazing, habitat degradation and fragmentation. These factors have been at the core of proposed explanations for the reproductive failure and numerical decline of Acacia carneorum and other threatened acacias. Paradoxically, the sympatric Acacia ligulata is thriving and highly fecund. Although these species have superficially similar floral displays, differences in sexual reproductive success may reflect interactions between flower and inflorescence ontogeny and pollinator assemblages. We compared the floral biology and flower visitor assemblages of A. carneorum and A. ligulata at four sites per species. Both species displayed similar floral ontogeny and synchronicity of display, with inflorescences simultaneously hermaphroditic for 4–5 days. However, A. ligulata displayed a higher density of flowers than A. carneorum and, while both species received a range of flower visitors, A. ligulata was visited by relatively few species and was serviced primarily by the non-native honeybee Apis mellifera, which typically made many within-plant movements during foraging bouts. In contrast, A. carneorum was visited by a diverse suite of native insects that carried little pollen and made fewer within plant movements. On average, Apis mellifera carried 98.4 % A. ligulata pollen, whereas the native insect visitors of A. carneorum carried only 45 % A. carneorum pollen. Differing floral ontogeny or lack of native pollinators does not explain the reproductive failure of A. carneorum. The success of A. ligulata may reflect pollination services provided by A. mellifera and interactions with plant mating systems.     

The excavation and cultivation in containers of mature grey mangroves,Avicennia marina

The grey mangrove, Avicennia marina, grows in intertidal habitat that is under pressure from expanding human settlement and industry along coastlines in the tropics and subtropics. Inappropriate clearing, and also the dieback of large stands of A. marina associated with pollution, have created an interest in methods for revegetation and also the need for a better understanding of the species physiological tolerances including to herbicide residue. Eco-toxicological studies have so far been exclusively on seedlings because of perceived difficulties cultivating mature plants in controlled environments. Reforestation projects are often associated with low rates of establishment, because of the susceptibility of propagules, seedlings and saplings to wind and wave erosion, as well as flooding and desiccation associated with the natural inter-tidal environment. We report on the successful excavation, and cultivation in containers with an automated irrigation system of eight mature A. marina plants. It is possible to transplant mature mangroves and achieve significant vegetative growth and also flowering and fruiting within the first year. It is difficult to compare the likely costs of transplanting a more limited number of mature plants to many propagules. However, we suggest that in situations where reforestation with young propagules, seedlings or saplings has failed because of excessive wash, drowning or smothering, the transplanting of mature mangroves could be a better option.     

The reproductive assurance benefit of selfing: importance of flower size and population size

Autonomous selfing can provide reproductive assurance (RA) for flowering plants that are unattractive to pollinators or in environments that are pollen limited. Pollen limitation may result from the breakdown of once-continuous habitat into smaller, more isolated patches (habitat fragmentation) if fragmentation negatively impacts pollinator populations. Here we quantify the levels of pollen limitation and RA among large and small populations of Collinsia parviflora, a wildflower with inter-population variation in flower size. We found that none of the populations were pollen limited, as pollen-supplemented and intact flowers did not differ in seed production. There was a significant effect of flower size on RA; intact flowers (can self) produced significantly more seeds than emasculated flowers (require pollen delivery) in small-flowered plants but not large-flowered plants. Population size nested within flower size did not significantly affect RA, but there was a large difference between our two replicate populations for large-flowered, small populations and small-flowered, large populations that appears related to a more variable pollination environment under these conditions. In fact, levels of RA were strongly negatively correlated with rates of pollinator visitation, whereby infrequent visitation by pollinators yielded high levels of RA via autonomous selfing, but there was no benefit of autonomous selfing when visitation rates were high. These results suggest that autonomous selfing may be adaptive in fragmented habitats or other ecological circumstances that affect pollinator visitation rates.     

Mechanisms of pollen deposition by insect pollinators

Summary Studies of pollen dispersal in insect-pollinated plants have often documented highly leptokurtic patterns of pollen deposition that can increase the likelihood of long-distance mating. To examine potential causes of highly leptokurtic deposition, we introduce four functions that arise when (1) the duration of pollinator visits to pollen sources is limited, (2) the rate of pollen deposition varies randomly among pollinators and/or among visits, (3) the rate of pollen deposition changes monotonically over time or (4) pollen is carried in layers or compartments on the pollinator's body that differ in deposition rate. Maximum likelihood techniques were used to fit deposition functions to data obtained from honey bees (Apis mellifera L.) visiting mustard plants (Brassica campestris L.) that contained a marker gene. Each of the alternative leptokurtic functions fit the experimental data better than a simple exponential function and the best-fit function predicted a mean pollen dispersal distance more than three times greater than the exponential. We argue that studies of pollen deposition need to test a broader range of deposition models to assess outcrossing distance in plant populations accurately.     

Importance of Conserving Alternative Pollinators: Assessing the Pollination Efficiency of the Squash Bee, Peponapis limitaris in Cucurbita moschata (Cucurbitaceae)

Although the honey bee, Apis mellifera, has been considered the best pollinator for crops needing insect pollination, the current pandemic of varroatosis among honeybees highlights the need to find additional or alternative species as managed crop pollinators. Moreover, there is evidence that A. mellifera may not always be the most efficient pollinator. Introduction of A. mellifera into crops may be unnecessary, and even detrimental to non-Apis bee populations, which should be considered as an alternative for crop production improvement. Evaluating the pollination efficiency of non-Apis bees is one of the first steps in planning successful strategies for their conservation. In this study, we evaluated the pollination efficiency of Peponapis limitaris and A. mellifera in plots of Cucurbita moschata: pollen removal and deposition; pollinator visit frequency; and the pollinator visit–nectar production relationship. The results show P. limitaris to be the most efficient pollinator as: (1) both males and females remove and deposit almost four times as much pollen as A. mellifera; (2) they make significantly more floral visits than A. mellifera; and (3) their visit frequency shows a strong relationship to C. moschata nectar production during anthesis. Recommendations arising from this study are: (1) the introduction of A. mellifera be avoided in C. moschata crops; and (2) basic research be done on the biology of P. limitaris that contribute to its conservation and greater exploitation.     

Fly pollination of Linum lewish (Linaceae)

This study examines the reproductive biology of Linum lewisii Pursh. (Linaceae), a polyphilic species visited by small bees and generalist flies in montane Colorado. L. lewisii plants growing at different sites experience large temporal and spatial variations in pollinator visits. Their ability to attract both dipteran and hymenopteran pollinators allows pollination under varying conditions as pollinator pool composition changes. Although L. lewisii is self-compatible, hand-pollination studies indicate that insects are required for seed production. The relative effectiveness of fly and bee pollinators is assessed in terms of per-visit pollen deposition. Insect visitation patterns are combined with per-visit effectiveness data to evaluate the relative importance of different pollinator groups. Overall, bees tend to be more effective than flies in depositing pollen. However, in many instances flies appear to be responsible for more pollen deposition due to their higher visitation rates.     

Within-population spatial variation in pollinator visitation rates,pollen limitation on seed set,and flower longevity in an alpine species

Pollen limitation through insufficient pollen deposition on stigmas caused by too infrequent pollinator visitation may influence the reproductive outcome of plants. In this study we investigated how pollinator visitation rate, the degree of pollen limitation, and flower longevity varied spatially among three sites at different altitudes within a population of the dwarf shrub Dryas octopetala L. in alpine southern Norway. Significant pollen limitation on seed set only occurred at the mid-elevation site, while seed set at the other sites appeared to be mainly resource limited, thus indicating a spatial variation in pollen limitation. There was no association between the spatial variation in the extent of pollen limitation and pollinator visitation rate to flowers. However, pollinator visitation rates were related to flower longevity of Dryas; sites with low visitation rates had long-lived flowers and vice versa. Thus, our results suggest within-population spatial co-variation between pollinator visitation rates, pollen limitation, and a developmental response to these factors, flower longevity.     

Non-native plants affect generalist pollinator diet overlap and foraging behavior indirectly,via impacts on native plant abundance

Flowering invasive plants can have dramatic effects on the resource landscape available to pollinators. Because many pollinators exhibit behavioral plasticity in response to competitor or resource density, this in turn can result in impacts on ecological processes such as pollination and plant reproduction. We examine how interactions between five common generalist eusocial bees change across an invasion gradient by examining how bee abundance and diet overlap changed with variation in both invasive plant abundance and competitor abundance in a temperate oak-savannah ecosystem. Specifically we focus on the bumblebees Bombus bifarius, B. mixtus, B. melanopygus and B. vosnesenskii, as well as the non-native honeybee Apis mellifera, and their interactions with the native flowering plants Camassia quamash, Camassia liechtlinii, and the invasive shrub Cytisus scoparius. We further examine whether changes in pollinator visits to the invasive and two common native plants can explain changes in diet overlap. Abundance of the invasive plant and other common floral resources had strong impacts on focal bee abundance, with certain species more likely to be present at highly invaded sites. This may be because highly invaded sites tended to be embedded in forested landscapes where those bees are common. Diet overlap was most affected by abundance of a common native plant, rather than the invasive plant, with diet overlap increasing non-linearly with abundance of the native plant. Furthermore, Apis mellifera, did not appear to have direct competitive effects on native bumblebees in this habitat. However, visit patterns suggest that bees most abundant at highly invaded sites may compete for access to native resources. Thus the impacts of this invasive plant on our focal bee species may be primarily indirect, via its’ competitive effects on native plants.     

The most effective pollinator revisited: pollen dynamics in a spring-flowering herb

Most flowers are visited by a wide range of potential pollinators. However, their efficiency in pollen removal and deposition, and other behavioural factors affecting pollination effectiveness may greatly differ among taxa, and even individuals. Fritillary (Fritillaria meleagris L., Liliaceae) is a spring-flowering, critically endangered plant in the Polish flora, red-listed in most of the European countries of its range. Based on indirect evidence, that is, body pollen loads, visitation frequency and seasonal abundance, it is estimated that its key pollinators are queen bumblebees, but, as shown in the literature, the largest Fritillaria pollen loads are carried by solitary bees. To study pollinator effectiveness for floral visitors to F. meleagris, we performed a garden experiment, where we analysed pollen deposition and assessed pollen removal per single flower-visit in the plant. Similarly to field conditions reported in the literature, our experimental plants were serviced by nectar-seeking bumblebee queens and two taxa of solitary bees, small pollen-collecting Andrena and large, nectar-seeking Anthophora males. When “quality” component was addressed, despite the character of visits, insects from all groups deposited more pollen than was found on unvisited flowers, but they did not differ significantly from each other in pollen deposition on virgin stigmas. We also found some differences in pollen removal both within- and among-visitor species and control flowers, unfortunately due to extremely high variation of the results they were all statistically insignificant. However, when “quantity” component of insect performance was concerned, we observed that over 81 % of visits were by bumblebees. Bombus queens stayed on flowers significantly less time than small Andrena individuals (13 % of recorded visits) and equally long as Anthophora males (only 6 % of visits). We conclude that although all the visitor groups can pollinate the flowers of F. meleagris, bumblebee queens indeed proved to be the most effective pollinators of the plant, when both quality and quantity components of pollination are concerned.     

The Abundance and Pollen Foraging Behaviour of Bumble Bees in Relation to Population Size of Whortleberry (Vaccinium uliginosum)

Habitat fragmentation can have severe effects on plant pollinator interactions, for example changing the foraging behaviour of pollinators. To date, the impact of plant population size on pollen collection by pollinators has not yet been investigated. From 2008 to 2010, we monitored nine bumble bee species (Bombus campestris, Bombus hortorum s.l., Bombus hypnorum, Bombus lapidarius, Bombus pascuorum, Bombus pratorum, Bombus soroensis, Bombus terrestris s.l., Bombus vestalis s.l.) on Vaccinium uliginosum (Ericaceae) in up to nine populations in Belgium ranging in size from 80 m² to over 3.1 ha. Bumble bee abundance declined with decreasing plant population size, and especially the proportion of individuals of large bumble bee species diminished in smaller populations. The most remarkable and novel observation was that bumble bees seemed to switch foraging behaviour according to population size: while they collected both pollen and nectar in large populations, they largely neglected pollen collection in small populations. This pattern was due to large bumble bee species, which seem thus to be more likely to suffer from pollen shortages in smaller habitat fragments. Comparing pollen loads of bumble bees we found that fidelity to V. uliginosum pollen did not depend on plant population size but rather on the extent shrub cover and/or openness of the site. Bumble bees collected pollen only from three plant species (V. uliginosum, Sorbus aucuparia and Cytisus scoparius). We also did not discover any pollination limitation of V. uliginosum in small populations. We conclude that habitat fragmentation might not immediately threaten the pollination of V. uliginosum, nevertheless, it provides important nectar and pollen resources for bumble bees and declining populations of this plant could have negative effects for its pollinators. The finding that large bumble bee species abandon pollen collection when plant populations become small is of interest when considering plant and bumble bee conservation.     

Climate and human influence shape the interactive role of the honeybee in pollination networks beyond its native distributional range

Empirical evidence has shown that introduced honeybees, Apis mellifera L., can change the structural organization of ecological networks involving pollinators and flowering plants. In this case, studies have shown that A. mellifera is highly connected within networks (i.e., high interactive role) mainly due to its high abundances, long colony lifetime, generalist diet, and great capacity of spread and adaptation to new environments. However, there is little knowledge on how the interactive role of A. mellifera in these flower-visitor networks changes spatially and the potential correlates of such variation across ecological and geographical gradients worldwide. In this study, we evaluated how native and exotic distribution ranges, local climate, human influence (i.e., anthropogenic disturbance), and latitude shape the interactive role of the honeybee in plant-floral visitor networks globally. To achieve that, we compiled 95 weighted plant-floral visitor networks encompassing ca. 87 degrees of latitude and 227 degrees of longitude across the globe. We found that the interactive role of A. mellifera is higher in its exotic range than in the native distributional range and positively associated with mean annual temperature and precipitation. Further, the interactive role of A. mellifera was high at intermediate values of global human influence and increased with latitude. Our findings help us to understand how different drivers related to rapid global change could affect the capacity of A. mellifera to colonize and increase its impact on other pollinator species in different environments.