Native and Non-Native Supergeneralist Bee Species Have Different Effects on Plant-Bee Networks

Supergeneralists, defined as species that interact with multiple groups of species in ecological networks, can act as important connectors of otherwise disconnected species subsets. In Brazil, there are two supergeneralist bees: the honeybee Apis mellifera, a non-native species, and Trigona spinipes, a native stingless bee. We compared the role of both species and the effect of geographic and local factors on networks by addressing three questions: 1) Do both species have similar abundance and interaction patterns (degree and strength) in plant-bee networks? 2) Are both species equally influential to the network structure (nestedness, connectance, and plant and bee niche overlap)? 3) How are these species affected by geographic (altitude, temperature, precipitation) and local (natural vs. disturbed habitat) factors? We analyzed 21 plant-bee weighted interaction networks, encompassing most of the main biomes in Brazil. We found no significant difference between both species in abundance, in the number of plant species with which each bee species interacts (degree), and in the sum of their dependencies (strength). Structural equation models revealed the effect of A. mellifera and T. spinipes, respectively, on the interaction network pattern (nestedness) and in the similarity in bee’s interactive partners (bee niche overlap). It is most likely that the recent invasion of A. mellifera resulted in its rapid settlement inside the core of species that retain the largest number of interactions, resulting in a strong influence on nestedness. However, the long-term interaction between native T. spinipes and other bees most likely has a more direct effect on their interactive behavior. Moreover, temperature negatively affected A. mellifera bees, whereas disturbed habitats positively affected T. spinipes. Conversely, precipitation showed no effect. Being positively (T. spinipes) or indifferently (A. mellifera) affected by disturbed habitats makes these species prone to pollinate plant species in these areas, which are potentially poor in pollinators.     

Ecological engineering by a native leaf-cutting ant increases the performance of exotic plant species

Numerous mechanisms are proposed to explain why exotic plants successfully invade natural communities. However, the positive effects of native engineers on exotic plant species have received less consideration. We tested whether the nutrient-rich soil patches created by a native ecological engineer (refuse dumps from the leaf-cutting ant Acromyrmex lobicornis) increase the performance of exotic more than native plants. In a greenhouse experiment, individuals from several native and exotic species were planted in pots with refuse dumps (RDs) and non-nest soils (NNSs). Total plant biomass and foliar nutrient content were measured at the end of the experiment. We also estimated the cover of exotic and native plant species in external RDs from 54 field ant nests and adjacent areas. Greenhouse plants showed more biomass and foliar nutrient content in RDs than in NNS pots. Nevertheless, differences in the final mean biomass among RD and NNS plants were especially great in exotics. Accordingly, the cover of exotic plants was higher in field RDs than in adjacent, non-nest soils. Our results demonstrated that plants can benefit from the enhanced nutrient content of ant RDs, and that A. lobicornis acts as an ecosystem engineer, creating a substrate that especially increases the performance of exotics. This supports the fluctuating resource hypothesis as a mechanism to promote biological invasions, and illustrates how this hypothesis may operate in nature. Since ant nests and exotic plants are more common in disturbed than in pristine environments, the role of ant nests in promoting biological invasions might be of particular interest. Proposals including the use of engineer species to restore disturbed habitats should be planned with caution because of their potential role in promoting invasions.     

Influence of Apis mellifera L. (Hymenoptera: Apidae) on the Use of the Most Abundant and Attractive Floral Resources in a Plant Community

Some factors influence the distribution of abundance of floral visitors, especially the amount and quality of the floral resources available, the size of the area occupied by the visitor, habitat heterogeneity, and the impact caused by natural enemies and introduced species. The objective of this research was to evaluate the distribution of abundance of the foraging activity of native floral visitors and Apis mellifera L. in the most abundant and attractive food sources in a secondary forest fragment with features of Cerrado-Atlantic Forest. Some plant species were selected and the frequency of foraging made by floral visitors was recorded. A high abundance of visits in flowers was performed by A. mellifera. Two factors may have influenced this result: (1) the occupation of the forest fragment predominantly by vines and shrubs at the expenses of vegetation with arboreal characteristics that favored the encounter of the flowering plants by A. mellifera; (2) rational beekeeping of A. mellifera, causing the number of natural swarms which originate annually from colonies of commercial apiaries and colonies previously established in the environment to be very high, thus leading to an increase in the population size of this bee species in the study site. The frequent occurrence of human-induced fire and deforestation within the forest fragment may have reduced the population size of the bee species, including A. mellifera. As the populations of A. mellifera have the capacity to quickly occupy the environment, this species possibly became dominant after successive disturbances made in the forest fragment.     

Minimal Effects of an Invasive Flowering Shrub on the Pollinator Community of Native Forbs

Biological invasions can strongly influence species interactions such as pollination. Most of the documented effects of exotic plant species on plant-pollinator interactions have been observational studies using single pairs of native and exotic plants, and have focused on dominant exotic plant species. We know little about how exotic plants alter interactions in entire communities of plants and pollinators, especially at low to medium invader densities. In this study, we began to address these gaps by experimentally removing the flowers of a showy invasive shrub, Rosa multiflora, and evaluating its effects on the frequency, richness, and composition of bee visitors to co-flowering native plants. We found that while R. multiflora increased plot-level richness of bee visitors to co-flowering native plant species at some sites, its presence had no significant effects on bee visitation rate, visitor richness, bee community composition, or abundance overall. In addition, we found that compared to co-flowering natives, R. multiflora was a generalist plant that primarily received visits from generalist bee species shared with native plant species. Our results suggest that exotic plants such as R. multiflora may facilitate native plant pollination in a community context by attracting a more diverse assemblage of pollinators, but have limited and idiosyncratic effects on the resident plant-pollinator network in general.     

No impact of a native beetle on exotic plant performance and competitive ability due to plant compensation

The novel associations between invasive plants and their natural enemies in the introduced range have recently received increasing attention; however, the effects of novel enemies on exotic plant performance and competition with native species remain poorly explored. Here, we tested the impact of herbivory by a native beetle, Cassida piperata, on the performance of the exotic species Alternanthera philoxeroides and competition with a native congener, Alternanthera sessilis, using common garden experiments in central China. We found A. philoxeroides was able to fully compensate for intense herbivory by C. piperata. Herbivory by C. piperata that released at the average density in this region had no impact on competition between the native and exotic plant species. Our results indicate that herbivory by novel enemies may not reduce exotic plant performance due to plant compensation. However, high tolerance to herbivory may not confer a competitive advantage for exotic species compared to less tolerant native competitors if the herbivore damage is below a certain threshold. Thus, it is necessary to assess the impact of novel enemies on exotic plant performance and competition with native plants along gradients of insect densities. This may lead to a better understanding of how best to exploit the role of native herbivores in facilitating or slowing plant invasions.     

Additive effects of exotic plant abundance and land-use intensity on plant–pollinator interactions

The continuing spread of exotic plants and increasing human land-use are two major drivers of global change threatening ecosystems, species and their interactions. Separate effects of these two drivers on plant–pollinator interactions have been thoroughly studied, but we still lack an understanding of combined and potential interactive effects. In a subtropical South African landscape, we studied 17 plant–pollinator networks along two gradients of relative abundance of exotics and land-use intensity. In general, pollinator visitation rates were lower on exotic plants than on native ones. Surprisingly, while visitation rates on native plants increased with relative abundance of exotics and land-use intensity, pollinator visitation on exotic plants decreased along the same gradients. There was a decrease in the specialization of plants on pollinators and vice versa with both drivers, regardless of plant origin. Decreases in pollinator specialization thereby seemed to be mediated by a species turnover towards habitat generalists. However, contrary to expectations, we detected no interactive effects between the two drivers. Our results suggest that exotic plants and land-use promote generalist plants and pollinators, while negatively affecting specialized plant–pollinator interactions. Weak integration and high specialization of exotic plants may have prevented interactive effects between exotic plants and land-use. Still, the additive effects of exotic plants and land-use on specialized plant–pollinator interactions would have been overlooked in a single-factor study. We therefore highlight the need to consider multiple drivers of global change in ecological research and conservation management.     

Phenotypic plasticity of reproductive traits in response to food availability in invasive and native species of nematode

Invasive species cause severe ecological and economic damage; however, the mechanisms underlying their successful invasion often remain elusive. In the case of Bursaphelenchus xylophilus, a global quarantine pest which invaded Asia and Europe, it has been suggested that this species possesses highly competitive abilities, which promotes its establishment and rapid spread. To explore biological traits that may explain its highly competitive abilities, we focused on expression of phenotypic plasticity in response to the food conditions experienced by the females during their development as juveniles in the invasive species B. xylophilus and native species Bursaphelenchus mucronatus. We report an unexpected significant difference of phenotypic trade-off between egg number and egg size in the invasive species B. xylophilus and native species B. mucronatus. This leads to superior propagation ability of invasive species, under high and low food conditions in culture. These effects reflect adaptive optimal resource allocation where more eggs are produced in favorable environments to enhance population viability. Furthermore, we show that B. xylophilus eggs hatched earlier than B. mucronatus when their parents experienced high food availability. Thus, this study revealed, for the first time, phenotypic plasticity of reproductive traits in B. xylophilus which empowers the species a competitive advantage relative to their native counterpart B. mucronatus when they are under different range of food availability. These results are a step towards answering the vital question of how an exotic invasive species exclude a native species from its original niche.     

A global meta‐analysis of exotic versus native leaf decay in stream ecosystems

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Native flower strips increase visitation by non-bee insects to avocado flowers and promote yield

Pollination is an essential ecosystem service for pollinator-dependent crops and plant communities. Apis mellifera L. is by far the most commonly used species to obtain this service in agriculture. However, there is growing evidence of the importance for crop yields of the service provided by wild bees and non-bee insect pollinators. Establishing flower resources in agricultural landscapes is a management practice that can increase insect pollinator populations and improve crop yields. We established perennial native flower strips (NFS) in four avocado orchards in central Chile during autumn 2017. We monitored flower visitors and counted newly formed fruits in avocados near and far from NFS in spring 2019, to assess flower visitor groups, flower visitation rates and fruit numbers. Only A. mellifera visited avocado flowers within bees, whereas both the managed A. mellifera and wild bees were the main visitors to the NFS. NFS increased visitation rates to adjacent avocado of flies, and with a trend for the sum of all non-managed flower visitors (i.e. excluding A. mellifera). However, there were no differences in the rates of A. mellifera visitation to avocados near and far from NFS. Avocado fruit numbers were higher among avocados near NFS than among those farther away. This difference could be due to better pollination by the increased visits to avocado flowers by flies or other wild insects. Therefore, NFS could contribute to crop fruit number, as well as conservation of native flora, wild bees and non-bee pollinators on fruit farms in the “Central Chile” biodiversity hotspot.     

An invasive plant provides refuge to native plant species in an intensely grazed ecosystem

Invasion by exotic plant species and herbivory can individually alter native plant species diversity, but their interactive effects in structuring native plant communities remain little studied. Many exotic plant species escape from their co-evolved specialized herbivores in their native range (in accordance with the enemy release hypothesis). When these invasive plants are relatively unpalatable, they may act as nurse plants by reducing herbivore damage on co-occurring native plants, thereby structuring native plant communities. However, the potential for unpalatable invasive plants to structure native plant communities has been little investigated. Here, we tested whether presence of an unpalatable exotic invader Opuntia ficus-indica was associated with the structure of native plant communities in an ecosystem with a long history of grazing by ungulate herbivores. Along 17 transects (each 1000 m long), we conducted a native vegetation survey in paired invaded and uninvaded plots. Plots that harboured O. ficus-indica had higher native plant species richness and Shannon–Wiener diversity H′ than uninvaded plots. However, mean species evenness J was similar between invaded and uninvaded plots. There was no significant correlation between native plant diversity and percentage plot cover by O. ficus-indica. Presence of O. ficus-indica was associated with a compositional change in native community assemblages between paired invaded and uninvaded plots. Although these results are only correlative, they suggest that unpalatable exotic plants may play an important ecological role as refugia for maintenance of native plant diversity in intensely grazed ecosystems.     

Pollination Services Provided by Bees in Pumpkin Fields Supplemented with Either Apis mellifera or Bombus impatiens or Not Supplemented

Pollinators provide an important service in many crops. Managed honey bees (Apis mellifera L.) are used to supplement pollination services provided by wild bees with the assumption that they will enhance pollination, fruit set and crop yield beyond the levels provided by the wild bees. Recent declines in managed honey bee populations have stimulated interest in finding alternative managed pollinators to service crops. In the eastern U.S., managed hives of the native common eastern bumble bee (Bombus impatiens Cresson) may be an excellent choice. To examine this issue, a comprehensive 2-yr study was conducted to compare fruit yield and bee visits to flowers in pumpkin (Cucurbita pepo L.) fields that were either supplemented with A. mellifera hives, B. impatiens hives or were not supplemented. We compared pumpkin yield, A. mellifera flower visitation frequency and B. impatiens flower visitation frequency between treatments. Results indicated that supplementing pumpkin fields with either A. mellifera or B. impatiens hives did not increase their visitation to pumpkin flowers or fruit yield compared with those that were not supplemented. Next, the relationship between frequency of pumpkin flower visitation by the most prominent bee species (Peponapis pruinosa (Say), B. impatiens and A. mellifera) and fruit yield was determined across all pumpkin fields sampled. Fruit yield increased as the frequency of flower visits by A. mellifera and B. impatiens increased in 2011 and 2012, respectively. These results suggest that supplementation with managed bees may not improve pumpkin production and that A. mellifera and B. impatiens are important pollinators of pumpkin in our system.     

Pollination and breeding system of Melochia tomentosa L. (Malvaceae), a keystone floral resource in the Brazilian Caatinga

The main goals of the present paper were to investigate the floral biology and the breeding system of Melochia tomentosa in a semi-arid region in Brazil, comparing the role of Apis mellifera with other native pollinators, and to discuss the importance of this plant species as a floral resource for the local fauna in maintaining different guilds of specialized pollinators in the Caatinga. M. tomentosa is very common in Caatinga areas and blooms year-round with two flowering peaks, one in the wet and another in the dry period. The pink, bright-colored flowers are distylous and both morphs are homogamous. The trichomatic nectary is located on the inner surface of the connate sepals, and the nectar (ca. 7 μl) is accumulated in the space between the corolla and the calyx. Nectar sugar concentration reaches an average of 28%. The results of controlled pollination experiments show that M. tomentosa is self-incompatible. Pollen viability varies from 94% to 98%. In spite of being visited by several pollen vectors, flower attributes of M. tomentosa point to melittophily, and A. mellifera was the most frequent visitor and the principal pollinator. Although honeybees are exotic, severely competing with native pollinators, they are important together with other native bees, like Centris and Xylocopa species, for the fruit set of M. tomentosa. This species represents a keystone floral resource favoring the maintenance of many species of bees, butterflies and hummingbirds through the year in this deciduous tropical dry forest.     

生物入侵对鸟类的生态影响

生物入侵是全球生物多样性面临的最主要威胁之一, 入侵种在改变入侵地环境的同时也使当地的生物受到极大影响。鸟类在生态系统中处于较高的营养级, 生态系统中任何一个环节的变化都可能对鸟类造成一定的影响。本文回顾了哺乳动物、鸟类、无脊椎动物和植物等不同生物类群的入侵对本地鸟类生态影响方面的研究进展。外来生物对鸟类的影响主要表现在以下几方面: (1)外来哺乳动物对成鸟、幼鸟或鸟卵的捕食作用; (2)外来鸟类与本地鸟类竞争栖息地和食物资源, 与当地的近缘种杂交而造成基因流失; (3)外来无脊椎动物改变本地鸟类的栖息环境和食物状况, 甚至直接捕食本地鸟类; (4)外来植物入侵改变入侵地的植物群落组成和结构, 造成本地鸟类的栖息地丧失或破碎化, 并通过改变入侵地生态系统的食物链结构而对高营养级的鸟类产生影响。最后, 作者还提出了该领域有待解决的问题和今后可能的研究方向。     

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.     

Drosophilid Assemblages at Different Urbanization Levels in the City of Porto Alegre, State of Rio Grande do Sul, Southern Brazil

The present study analyzed the drosophilid assemblages in different levels of urbanization in the city of Porto Alegre, Rio Grande do Sul, Brazil. Collections were carried out in 2008 in three different environments: a highly urbanized area????Jardim Botanico,?? a forested area with intermediary urbanization????Parque Gabriel Knijnik,?? and in a relatively well-preserved forested area, although threatened by the urban growth????Morro Santana.?? In Jardim Botanico, 36 species belonging to four genera were found, with high abundance of exotic species as Drosophila simulans Sturtevant and Zaprionus indianus (Gupta). In Parque Gabriel Knijnik, 33 species that belonged to four genera were found, with higher abundances of native species belonging to the Drosophila tripunctata species group and Drosophila willistoni species subgroup, and lower abundance of exotic species. As for Morro Santana, 32 species and three genera were found, with higher abundances of native groups, low representativeness of exotic species, and absence of Zaprionus indianus. The analysis of the Jaccard index showed higher similarity in the species composition between samples collected in summer and autumn, and between samples collected in winter and spring. On the other hand, the Morisita index differentiated Jardim Botanico from the other two studied sites. Our results show that Morro Santana is an important area of native biodiversity, reinforcing, therefore, the inclusion of this area in the project for the creation of an ecological corridor as proposed by the Ministry of the Environment of Brazil.     

外来植物紫茎泽兰入侵对土壤理化性质及丛枝菌根真菌(AMF)群落的影响

为了揭示外来植物紫茎泽兰入侵对入侵地土壤丛枝菌根真菌(AMF)群落及相关肥力的影响,比较测定了紫茎泽兰不同入侵程度土壤理化性质、AMF侵染率及AMF群落的差异。结果表明,紫茎泽兰入侵降低了土壤pH,使土壤中有机碳、全氮和速效钾含量分别增加83.0%,106.9%和111.0%;尽管对全磷含量没有显著影响,但有机磷含量呈升高的趋势,而速效磷呈降低的趋势。紫茎泽兰入侵降低了本地植物的AMF侵染率;随着入侵程度的加深,土壤中以膨胀无梗囊霉(Acauospora dilatata)为优势种的AMF群落结构逐渐转变为以近明球囊霉(Glomus claroideum )为优势种的结构,紫茎泽兰可在其根周选择培育近明球囊霉,而对其它AMF种,特别是对膨胀无梗囊霉则存在抑制作用;基于各AMF种多度的聚类分析表明,形成紫茎泽兰单优群落土壤中各AMF种多度与未入侵的本地植物群落及入侵程度较轻的紫茎泽兰与本地植物群落之间存在明显分歧。综合分析推断认为,紫茎法兰入侵改变了入侵地土壤理化性状,抑制AMF对土著植物的侵染,改变AMF群落,并在其根周选择培育近明球囊霉,这可能是紫茎泽兰入侵及扩张的重要途径之一。     

Physiological responses to short-term water and light stress in native and invasive plant species in southern California

As climate variability increases in low-resource environments, the ability of native and invasive species to tolerate stress and respond to large, ephemeral resource pulses will strongly influence plant fitness and, consequently, competitive outcomes. We examined how native and invasive species occurring in arid coastal sage scrub communities in southern California responded to water and high-light stress. We also examined how plants responded to irrigation following short-term water stress. While species responded differently to water and light treatments, no general pattern emerged between native and invasive species. Photosynthetic function of Ricinus communis (invasive) and Salvia mellifera (native) was most robust to water stress and most responsive to irrigation following water stress. Leaf transpiration data suggested that Ricinus and Salvia maintained photosynthetic function by high water use efficiency rather than higher water status via large root biomass. Brassica nigra (invasive) and Encelia californica (native) were more resistant to photoinhibition in response to high-light stress than Ricinus, Salvia, Artemesia californica (native) or Nicotiana glauca (invasive). Our data suggest that native and invasive species in these arid systems display a range of physiological responses to stress and that strategies for invasive species control or native ecosystem restoration based on plant responses to stress may require species-specific approaches.     

Use of an exotic host plant shifts immunity,chemical defense,and viral burden in wild populations of a specialist insect herbivore

Defense against natural enemies constitutes an important driver of herbivore host range evolution in the wild. Populations of the Baltimore checkerspot butterfly, Euphydryas phaeton (Nymphalidae), have recently incorporated an exotic plant, Plantago lanceolata (Plantaginaceae), into their dietary range. To understand the tritrophic consequences of utilizing this exotic host plant, we examined immune performance, chemical defense, and interactions with a natural entomopathogen (Junonia coenia densovirus, Parvoviridae) across wild populations of this specialist herbivore. We measured three immune parameters, sequestration of defensive iridoid glycosides (IGs), and viral infection load in field‐collected caterpillars using either P. lanceolata or a native plant, Chelone glabra (Plantaginaceae). We found that larvae using the exotic plant exhibited reduced immunocompetence, compositional differences in IG sequestration, and higher in situ viral burdens compared to those using the native plant. On both host plants, high IG sequestration was associated with reduced hemocyte concentration in the larval hemolymph, providing the first evidence of incompatibility between sequestered chemical defenses and the immune response (i.e., the “vulnerable host” hypothesis) from a field‐based study. However, despite this negative relationship between IG sequestration and cellular immunity, caterpillars with greater sequestration harbored lower viral loads. While survival of virus‐infected individuals decreased with increasing viral burden, it ultimately did not differ between the exotic and native plants. These results provide evidence that: (1) phytochemical sequestration may contribute to defense against pathogens even when immunity is compromised and (2) herbivore persistence on exotic plant species may be facilitated by sequestration and its role in defense against natural enemies.     

Biodiversity Loss following the Introduction of Exotic Competitors: Does Intraguild Predation Explain the Decline of Native Lady Beetles?

Exotic species are widely accepted as a leading cause of biodiversity decline. Lady beetles (Coccinellidae) provide an important model to study how competitor introductions impact native communities since several native coccinellids have experienced declines that coincide with the establishment and spread of exotic coccinellids. This study tested the central hypothesis that intraguild predation by exotic species has caused these declines. Using sentinel egg experiments, we quantified the extent of predation on previously-common (Hippodamia convergens) and common (Coleomegilla maculata) native coccinellid eggs versus exotic coccinellid (Harmonia axyridis) eggs in three habitats: semi-natural grassland, alfalfa, and soybean. Following the experiments quantifying egg predation, we used video surveillance to determine the composition of the predator community attacking the eggs. The extent of predation varied across habitats, and egg species. Native coccinellids often sustained greater egg predation than H. axyridis. We found no evidence that exotic coccinellids consumed coccinellid eggs in the field. Harvestmen and slugs were responsible for the greatest proportion of attacks. This research challenges the widely-accepted hypothesis that intraguild predation by exotic competitors explains the loss of native coccinellids. Although exotic coccinellids may not be a direct competitor, reduced egg predation could indirectly confer a competitive advantage to these species. A lower proportion of H. axyridis eggs removed by predators may have aided its expansion and population increase and could indirectly affect native species via exploitative or apparent competition. These results do not support the intraguild predation hypothesis for native coccinellid decline, but do bring to light the existence of complex interactions between coccinellids and the guild of generalist predators in coccinellid foraging habitats.