POLLINATORS POLLINATION AND FOOD PRODUCTION

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THE ASSESSMENT REPORT ON POLLINATORS, POLLINATION AND FOOD PRODUCTION TABLE 2.5.1 Main sources (meta-analyses, reviews) and scope of evidence used in assessment of the impact of invasive alien plants, pollinators, herbivores and predators on native pollinator species, networks and pollination. 90 2. DRIVERS OF CHANGE OF POLLINATORS, POLLINATION NETWORKS AND POLLINATION Citation Stout J.C. & Morales C.L. (2009). Apidologie, 40, 388-409 Traveset & Richardson (2006) Trends in Ecology & Evolution 21, 208-216. Traveset & Richardson (2014) Annual Review of Ecology, Evolution and Systematics 45, 89–113. Bezemer, et al. (2014) Annual Review of Entomology 59, 119-141. Dafni, et al. (2010) Applied Entomology and Zoology 45, 101-113. Dohzono & Yokoyama (2010) Applied Entomology and Zoology 45, 37-47. Monceau, et al (2014) Journal of Pest Science 87, 1-16. Morales & Traveset (2008). Critical Reviews in Plant Sciences, 27, 221-238. Paini D.R. (2004). Austral Ecology, 29, 399-407. Moritz, R.F.A., Hartel, S., Neumann, P. (2005) Ecoscience 12, 289-301. Study type Review Northern America (021) Northern Europe (154) Western Europe (155), Eastern Europe (151), Southern Europe (039), Asia (142) Impact of alien invasive species (plants & pollinators) on pollinators and pollination Review World (001) Impact of alien invasive insect & plant species on pollinators and pollination Review World (001) Effect of invasive alien species on mutualisms, including pollinators and pollination Review World (001) Response of native insect pollinators and plants to invasive alien plants. Review World (001) Impact of commercial Bombus terrestris introductions on native bumble bees Review South America-Argentina-Brazil (005-032-076)/Eastern Africa- Mauritius (014-480)/Australia (036)/ Southern Europe–Spain (039-724)/ Asia-Japan (142-392) Impacts of alien honey bees and bumble bees on native plant-pollinator relationships Review Europe (150) Potential impacts of invasive Asian hornet (Vespa velutina) on European honey bees Review World (001) Impact of invasive alien plants for native plant reproduction Review World (001) Impact of the introduced honey bee (Apis mellifera) on native bees Review World (001) Global invasions of the western honeybee (Apis mellifera) and the consequences for pollinators and pollination. plant species. Introduced mutualists may either fail or succeed in establishing within a novel ecological community according to the strength of interaction with the native species, for instance, if an introduced pollinator fails to obtain sufficient resources from the resident plant species then establishment is unlikely (Jones and Gomulkiewicz, 2012). Moreover, genetic diversity in introduced and resident species may, contingent on interaction strength, lead to rapid evolutionary selection for integration of the invader into the recipient community (Bossdorf et al., 2005; Jones and Gomulkiewicz, 2012; Vandepitte et al., 2014). Insect-pollinated plant species often dominate lists of invasive alien plants, but at least in the early stages of colonization the ability of these plants to self-pollinate enables establishment and spread (Chrobock et al., 2013; Pysek et al., 2011; Traveset and Richardson, 2014). Over time, other plant traits (e.g., flower morphology, copious nectar or pollen rewards, large floral or long duration displays) lure and co-opt UN geographical regions (numeric code) Topic Effect of invasive Invasive plants: +/-/=; Invasive pollinators: =/- Invasive pollinators:+/-/= Invasive herbivores: - Invasive predators: - Invasive plants: +/- Invasive plants: +/- Invasive plants +/-/= - Apis mellifera: +/- Bombus - - (Predicted) =/- Some - interactions, but impacts on fitness or population size either equivocal or = Mostly =, but a few examples of - impacts pollinator species whose phenotypes are pre-adapted to the floral resources the invasive alien plant offers (Chrobock et al., 2013; Kleijn and Raemakers, 2008; Morales and Traveset, 2009; Naug and Arathi, 2007; Pysek et al., 2011; Stout and Morales, 2009). Invasive alien plant species can thus become integrated into the ecosystem and dominate plant-pollinator interactions (Pysek et al., 2011; Traveset et al., 2013; Traveset and Richardson, 2006; Vilà et al., 2009). For example, pollen loads carried by insects may become dominated by alien pollen and hence potentially reduce conspecific pollen transfer among native plant species (e.g., Kleijn and Raemakers, 2008; Lopezaraiza-Mikel et al., 2007). Invasive flowering plants can also affect pollinators’ nutrition. Indeed, nutritional requirements differ among bee species and honey bee worker castes, and the growth and survival of social and solitary bee species is sensitive to the composition of the pollen diet (Paoli et al., 2014; Praz et al., 2008; Sedivy et al., 2011; Tasei and Aupinel, 2008). Therefore, while alien pollen and nectar may provide an additional food source
THE ASSESSMENT REPORT ON POLLINATORS, POLLINATION AND FOOD PRODUCTION for pollinators adapted to exploit them, there may also be a potential risk to pollinator health if invasive alien plant pollen is nutritionally poor compared to that from native plants (Stout and Morales, 2009). Invasive plants are expected to affect pollinators adversely if they either ill-adapted to exploit the alien food resource or dependent on native plants outcompeted by the invader (Bjerknes et al., 2007; Palladini and Maron, 2014; Stout and Morales, 2009). There is, however, little evidence from meta-analyses or reviews (Bjerknes et al., 2007; Montero- Castaño and Vilà, 2012; Stout and Morales, 2009), and only very few individual examples (Lopezaraiza-Mikel et al., 2007; Moroń et al., 2009; Nienhuis et al., 2009) of alien plant invasions consistently lowering overall pollinator diversity or abundance. There is more evidence, however, that alien plant invasions can influence the assembly of pollinator communities. Plant-pollinator community networks are permeable to plant invaders (Traveset et al., 2013; Traveset and Richardson, 2014), which according to the species involved can rewire plant-pollinator interactions (e.g., Bartomeus et al., 2008). Network architecture can often be relatively unaltered by alien plant invasions, for instance, a pan-European analysis showed network nestedness, a property thought to confer a degree of stability on the community, was unaffected by the integration of alien plants into the network (Vilà et al., 2009). Although recent global meta-analyses have demonstrated changes in network properties following integration of alien invasive plants, the attractiveness of these invasive plants to native pollinators altered their behaviour, which led to changes in network properties (e.g. modularity, interaction strength) that are thought to enhance community stability (Aizen et al., 2008; Albrecht et al., 2014). For example, invasive plant species increased connectivity between network modules (subsets of tightly connected species) (Albrecht et al., 2014), which potentially increased the network’s robustness to species losses arising from future environmental changes. Furthermore, highly invaded networks are, on average, characterised by weaker mutualism strength (i.e. weak or asymmetric mutual dependences between interacting species), a property that may reduce the probability of secondary extinctions should a partner species in the network be lost (Aizen et al., 2008). It should be noted, however, that many of these predictions around network robustness are derived from simulation models of empirical network data (frequency of pairwise species interactions at the community level). The challenges of collecting such data typically preclude greater biological realism (temporal network dynamics, species competition) being built into these simulations. Therefore, due caution is required in interpreting these insights from simulation models for community stability. Nonetheless, while invasive plant species do not generally alter diversity or abundance (Montero-Castaño and Vilà, 2012) through usurpation and domination of pollinator interaction networks, they often hold a key role in community organisation (Aizen et al., 2008; Albrecht et al., 2014). This key role of invasive alien plants (and invasive alien pollinators – see section 2.5.3), once integrated into pollinator networks, has potential ramifications for individual native plant species. If the native plant becomes overly reliant on the invader for facilitation of pollination, then there is a potential risk to the native species should those connections become eroded or lost due to further environmental changes (Aizen et al., 2008). Invasive alien plants may alter interactions between native plants and their pollinators either through competition for pollinator visitation (Bjerknes et al., 2007; Dietzsch et al., 2011) or by elevation of pollinator activity to the level where co-flowering native plant pollination is facilitated (Bjerknes et al., 2007; Cawoy et al., 2012; McKinney and Goodell, 2011). Primary and meta-analyses suggest that pollinator visitation rates to native plant species tend to decrease with plant invasion, suggesting that competition for pollinators may be the prevailing process (Brown et al., 2002; Montero- Castaño and Vilà, 2012; Morales and Traveset, 2009). Whether this translates into reduced reproductive output of native plant species is less certain, potentially because of plant compensatory mechanisms (i.e., self-reproduction, recruitment of alternative pollinators) (Bjerknes et al., 2007; Dietzsch et al., 2011; Morales and Traveset, 2009; Traveset and Richardson, 2014), but see examples where negative effects are reported (Brown et al., 2002; Chittka and Schurkens, 2001; Thijs et al., 2012). Furthermore, the level of impact on flower visitation may be contingent on the composition of the pollinator community because of differential responses of pollinator groups (e.g., flies versus bees) to the invasive plant (Carvalheiro et al., 2014; Montero-Castaño and Vilà, 2012). The negative impact that alien plant invasions can have on native plant pollination and reproductive success is increased at high relative densities of alien flowers and/or when alien and native plants are related or have similar floral traits (i.e., flower anatomy, color, phenology large floral displays) (Bjerknes et al., 2007; Brown et al., 2002; Carvalheiro et al., 2014; Morales and Traveset, 2009; Pysek et al., 2011). Only if some or all of these conditions are met will the extent of pollinator sharing between the native and the invasive plant species rise to the point where there is an impact, positive or negative, on the native plant (e.g. Thijs et al., 2012). There have been no studies (to our knowledge at the time of writing) that have examined the impact of invasive alien wild plants on food crops, which represents a significant knowledge gap. The pollinator-mediated impacts of native co-flowering plant species on flowering invasive plants are not clear and have been less studied (Carvallo et al., 2013). 91 2. DRIVERS OF CHANGE OF POLLINATORS, POLLINATION NETWORKS AND POLLINATION
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POLLINATORS POLLINATION AND FOOD PRODUCTION

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