Developmental database for phenology models: related insect and mite species have similar thermal requirements

Two values of thermal requirements, the lower developmental threshold (LDT), that is, the temperature at which development ceases, and the sum of effective temperatures, that is, day degrees above the LDT control the development of ectotherms and are used in phenology models to predict time at which the development of individual stages of a species will be completed. To assist in the rapid development of phenology models, we merged a previously published database of thermal requirements for insects, gathered by online search in CAB Abstracts, with independently collected data for insects and mites from original studies. The merged database comprises developmental times at various constant temperatures on 1,054 insect and mite species, many of them in several populations, mostly pests and their natural enemies, from all over the world. We show that closely related species share similar thermal requirements and therefore, for a species with unknown thermal requirements, the value of LDT and sum of effective temperatures of its most related species from the database can be used.     

Do higher resource capture ability and utilization efficiency facilitate the successful invasion of native plants?

The great damage caused by native invasive species on natural ecosystems is prompting increasing concern worldwide. Many studies have focused on exotic invasive species. In general, exotic invasive plants have higher resource capture ability and utilization capacity, and lower leaf construction cost (CC) compared to noninvasive plants. However, the physiological mechanisms that determine the invasiveness of native plants are poorly understood. We hypothesized that native invaders, like exotic invaders, may have higher resource capture ability and utilization efficiency compared to native noninvaders. To test this hypothesis, ecophysiological traits including light-saturated photosynthetic rate (A_max), specific leaf area (SLA), photosynthetic nitrogen use-efficiency (PNUE), photosynthetic energy-use efficiency (PEUE), and mass-based and area-based leaf construction cost (CC_mass and CC_area) were measured. We compared the above traits between three pairs of native invasive and noninvasive native species, and between three pairs of exotic invasive and noninvasive species in Guangzhou, southern China. Our results showed that the native invaders had higher A_max, SLA, PNUE, PEUE and lower CC_mass, CC_area, compared to native noninvaders and that these traits were also found in the exotic invaders. PNUE and PEUE in the native invaders were 150.3 and 129.0% higher, respectively, than in noninvasive native species, while these same measures in exotic invaders were 43.0 and 94.2% higher, respectively, than in exotic noninvasive species. The results indicated that native invaders have higher resource capture ability and resource utilization efficiency, suggesting that these traits may be a common biological foundation underlying successful invasion by both native and exotic invasives.     

Evolutionary ecology of microsporidia associated with the invasive ladybird Harmonia axyridis

Invasive species are characterized by the rapid growth and spread of their populations after establishing a foothold in new habitats, and there are now many examples of such species negatively affecting biodiversity and the economy. It is unclear why some species can become successful invaders, whereas most (even if closely related) remain noninvasive. We previously proposed a hypothesis that parasites associated with invading species can promote their invasive success if they are harmless toward the invaders but harmful to their competitors and/or predators in the newly colonized habitat. Here we discuss whether microsporidia that have recently been discovered in the invasive ladybird Harmonia axyridis contribute to its invasive success. We show that all H. axyridis beetles sourced from diverse collection sites all over the world carry abundant microsporidia. This suggests that both native and invasive H. axyridis populations are associated with these tolerated parasites, which were likely to have existed in native populations before expansion rather than being acquired in newly colonized areas. We describe the pathogenesis of the microsporidia during different developmental stages of H. axyridis and we address the possibility that the predation of its infected eggs and larvae by competing native ladybird species may lead to their infection and ultimately to their decline. Finally, we discuss our initial hypothesis: microsporidia that are tolerated by an invasive vector insect can be active against susceptible native competitors and/or predator species.     

Specific leaf area relates to the differences in leaf construction cost,photosynthesis, nitrogen allocation,and use efficiencies between invasive and noninvasive alien congeners

Comparisons between invasive and native species may not characterize the traits of invasive species, as native species might be invasive elsewhere if they were introduced. In this study, invasive Oxalis corymbosa and Peperomia pellucida were compared with their respective noninvasive alien congeners. We hypothesized that the invasive species have higher specific leaf (SLA) than their respective noninvasive alien congeners, and analyzed the physiological and ecological consequences of the higher SLA. Higher SLA was indeed the most important trait for the two invaders, which was associated with their lower leaf construction cost, higher nitrogen (N) allocation to photosynthesis and photosynthetic N use efficiency (PNUE). The higher N allocation to photosynthesis of the invaders in turn increased their PNUE, N content in photosynthesis, biochemical capacity for photosynthesis, and therefore light-saturated photosynthetic rate. The above resource capture-, use- and growth-related traits may facilitate the two invaders' invasion, while further comparative studies on a wider range of invasive and noninvasive congeners are needed to understand the generality of this pattern and to fully assess the competitive advantages afforded by these traits.     

Low resource availability limits weed invasion of tropical savannas

The savanna biome is one of the least invaded among global biomes, although the mechanisms underpinning its resistance to alien species relative to other biomes is not well understood. Invaders generally are at the resource acquisitive end of functional global plant trait variation and in low-resource savanna environments we might expect that successful invaders will only outperform native species under resource rich or highly disturbed conditions. However, invaders may also directly exploit resource stressed environments using resource conservative traits in some situations. It’s also possible that successful invaders and native species largely overlap in their trait profiles indicating site specific environmental factors are responsible for invader success in particular contexts rather than a general trait and functional divergence between invaders and native species. To address these various hypotheses, we compared a suite of morphological and physiological traits in graminoid and herbaceous native and co-occurring invasive plant species across a range of habitats in savannas of the Kimberley region of northern Australia. Invader grass species had traits associated with resource acquisition and fast growth rates, such as high SLA and leaf nutrient contents. In contrast, dominant native perennial grasses had traits characteristic of resource conservation and slow growth in resource stressed conditions. Trait profiles among invasive forbs and legumes exhibited stress tolerant traits relative to their native counterparts. Invaders also displayed strong divergence in reproductive traits, suggesting diverse responses to disturbance not indicated by leaf economic traits alone. These results suggest that savannas may be resistant to invaders with resource acquisitive traits due to their strong resource limitation.     

Protected areas offer refuge from invasive species spreading under climate change

Protected areas (PAs) are intended to provide native biodiversity and habitats with a refuge against the impacts of global change, particularly acting as natural filters against biological invasions. In practice, however, it is unknown how effective PAs will be in shielding native species from invasions under projected climate change. Here, we investigate the current and future potential distributions of 100 of the most invasive terrestrial, freshwater, and marine species in Europe. We use this information to evaluate the combined threat posed by climate change and invasions to existing PAs and the most susceptible species they shelter. We found that only a quarter of Europe's marine and terrestrial areas protected over the last 100 years have been colonized by any of the invaders investigated, despite offering climatically suitable conditions for invasion. In addition, hotspots of invasive species and the most susceptible native species to their establishment do not match at large continental scales. Furthermore, the predicted richness of invaders is 11%–18% significantly lower inside PAs than outside them. Invasive species are rare in long‐established national parks and nature reserves, which are actively protected and often located in remote and pristine regions with very low human density. In contrast, the richness of invasive species is high in the more recently designated Natura 2000 sites, which are subject to high human accessibility. This situation may change in the future, since our models anticipate important shifts in species ranges toward the north and east of Europe at unprecedented rates of 14–55 km/decade, depending on taxonomic group and scenario. This may seriously compromise the conservation of biodiversity and ecosystem services. This study is the first comprehensive assessment of the resistance that PAs provide against biological invasions and climate change on a continental scale and illustrates their strategic value in safeguarding native biodiversity.     

Superior performance and nutrient-use efficiency of invasive plants over non-invasive congeners in a resource-limited environment

Invasive plant species are often found to have advantages over native species in growth-related traits, such as photosynthetic rate, in disturbed or resource-rich environments. However, resource-use efficiency, rather than opportunistic resource capture, may confer more advantages when resources are scarce. In this study, performance and functional traits of invasive and non-invasive members of the genus Pinus were contrasted under the condition of nutrient limitations. Invasive species outperformed non-invasive congeners by growing 28% faster, on average. Invasives and non-invasives did not differ in biomass allocation traits (root-weight ratio, stem-weight ratio, leaf-weight ratio, leaf area ratio, root: shoot coefficient), but invaders had thinner and/or less dense leaves, as shown by a significantly lower leaf mass per area and leaf dry mass fraction. No differences between invasives and non-invasives were apparent in area-based leaf content of nitrogen, chlorophyll, or total protein, nor did the two groups differ in how efficiently they took up nutrients (specific absorption rate per unit root mass). The trait most strongly associated with invasives’ superior performance was photosynthetic nitrogen-use efficiency. Non-invaders were more water-use efficient. The results suggests that the relative performance of invasive and non-invasive species is context-dependent. Invaders may allocate leaf nitrogen more efficiently to maximize photosynthesis and growth in nitrogen-poor soils, while non-invaders with more heavily defended leaves may have an advantage in drier areas. Rather than searching for a suite of traits that constitutes “invasiveness”, it may be necessary to identify potential invaders by traits that are most adaptive to the local resource context.     

Determinants of the geographical extent of invasive plants in China: effects of biogeographical origin,life cycle and time since introduction

Plant invasions cause serious impacts to environment and human economy, and lots of researches have been paid to examine factors that determine the geographical extent of plant invaders. Many intrinsic species attributes, external environmental factors, and time since introduction have all been proposed to explain invasive success. Data on invasive plants distribution in province regions in China were collected, and species attributes and historical factors such as biogeographical origin, life cycle, and time since introduction were determined. While some annuals may eventually occupy the whole country area, perennials are unlikely to do so, particularly those from Central and South America. Time since introduction is significantly related to the number of provinces occupied by an invader; nevertheless after control of certain residence time, annuals are not likely to more provinces than the perennial herbs. High-impact invaders tend to spread faster than other invaders, but only when their life cycle is taken into account. This suggests that other plant traits, instead of life cycle, along with extrinsic factors, greatly contribute to invasive success and distribution area. Invasive plants in China are mainly distributed in human-made habitats, and their long-distance dispersal is primarily associated with human activities. Future studies need to consider interactions between species traits, environmental conditions, and human activities for further insights in researches alike.     

Invasive forbs differ functionally from native graminoids, but are similar to native forbs

? Exotic plant invasions can alter ecosystem processes, particularly if the invasive species are functionally different from native species. We investigated whether such alterations can be explained by differences in functional traits between native and invasive plants of the same functional group or by differences in functional group affiliation. ? We compared six invasive forbs in Europe with six native forbs and six native graminoids in leaf and whole-plant traits, plasticity in response to nutrient supply and interspecific competition, litter decomposition rate, effects on soil nutrient availability, and allelopathy. All traits were measured in a series of pot experiments, and leaf traits additionally in the field. ? Invasive forbs differed from native forbs for only a few traits; they had less leaf chlorophyll and lower phosphorus (P) uptake from soil, but they tended to have a stronger allelopathic effect. The invasive forbs differed in many traits from the native graminoids, their leaves had lower tissue densities and a shorter life span, their litter decomposed faster and they had a lower nitrogen-use efficiency. ? Our results suggest that invasive forbs have the potential to alter ecosystem properties when invading graminoid-dominated and displacing native graminoids but not when displacing native forbs.     

Introduced Brassica nigra populations exhibit greater growth and herbivore resistance but less tolerance than native populations in the native range

Rapid post-introduction evolution has been found in many invasive plant species, and includes changes in defence (resistance and tolerance) and competitive ability traits. Here, we explored the post-introduction evolution of a trade-off between resistance to and tolerance of herbivory, which has received little attention. In a common garden experiment in a native range, nine invasive and 16 native populations of Brassica nigra were compared for growth and defence traits. Invasive populations had higher resistance to, but lower tolerance of, herbivore damage than native populations. Invasive populations survived better and produced more seeds than native ones when released from herbivores; but fitness was equivalent between the regions under ambient herbivory. The invasive populations grew taller, and produced more biomass and lighter seeds than natives, irrespective of insecticide treatment. In addition to supporting the idea of post-introduction rapid evolution of plant traits, our results also contribute to an emerging pattern of both increasing resistance and growth in invasive populations, contrary to the predictions of earlier theories of resistance-growth trade-offs.     

Eating their way to the top? Mechanisms underlying the success of invasive insect generalist predators

Insect generalist predators have been introduced outside of their native range intentionally to improve biological control, or accidentally during commerce, and can subsequently become invasive. Invasive insect generalist predators (IIGP) have widespread impacts on invaded communities because they consume both herbivores and other predators. Also, they often reach higher densities than and displace similar native species. Reflecting the complexity of their ecological roles, a wide variety of mechanisms might contribute to invasive success by IIGP. These species often drive resources to lower levels than do natives, leading to intense resource competition and sometimes competitive exclusion of other predators. The broader range of resources used by many IIGP can heighten their competitive advantage, particularly when IIGP exploit modified habitats. In either case, IIGP improve herbivore suppression by depressing prey densities below pre-invasion levels. Coexistence among native and invasive generalists is fostered when species differ substantially in their niche requirements. In this case, a larger proportion of the total prey population is subject to attack post- than pre-invasion, which strengthens prey suppression. On the other hand, some IIGP feed heavily on other predators, at times leading to a weakening of prey suppression. Future research should continue to explore the roles of competition and niche partitioning on larger spatial scales, and in both the native and invasive ranges of IIGP. Additionally, combining data from empirical studies with theory might be an effective way to predict the spread and community impacts of IIGP invasions.     

Global compositional variation among native and non-native regional insect assemblages emphasizes the importance of pathways

Insects are among the world’s most ecologically and economically important invasive species. Here we assemble inventories of native and non-native species from 20 world regions and contrast relative numbers among these species assemblages. Multivariate ordination indicates that the distribution of species among insect orders is completely different between native and non-native assemblages. Some orders, such as the Psocoptera, Dictyoptera, Siphonaptera, Thysanoptera, and Hemiptera, are always over-represented in the non-native compared to native assemblages. Other orders, such as the Plecoptera, Trichoptera, Ephemeroptera, Odonata, Mecoptera and Microcoryphila, are consistently under-represented in non-native assemblages. These patterns most likely arise both as a result of variation among taxa in their association with invasion pathways responsible for transporting species among world regions, as well as variation in life-history traits that affect establishment potential. However, our results indicate that species compositions associated with invasiveness are fundamentally different from compositions related to insularity, indicating that colonization of islands selects for a different group of insect taxa than does selection for successful invaders. Native and non-native assemblage compositions were also related, to a lesser extent, to latitude of the region sampled. Together, these results illustrate the dominant role of invasion pathways in shaping the composition of non-native insect assemblages. They also emphasize the difference between natural background colonization of islands and anthropogenic colonization events, and imply that biological invasions are not a simple subset of a long-standing ecological process.     

The cane toad's (Chaunus [Bufo] marinus) increasing ability to invade Australia is revealed by a dynamically updated range model

Invasive species threaten biological diversity throughout the world. Understanding the dynamics of their spread is critical to mitigating this threat. In Australia, efforts are underway to control the invasive cane toad (Chaunus [Bufo] marinus). Range models based on their native bioclimatic envelope suggest that the cane toad is nearing the end of its invasion phase. However, such models assume a conserved niche between native and invaded regions and the absence of evolution to novel habitats. Here, we develop a dynamically updated statistical model to predict the growing extent of cane toad range based on their current distribution in Australia. Results demonstrate that Australian cane toads may already have the ability to spread across an area that almost doubles their current range and that triples projections based on their native distribution. Most of the expansion in suitable habitat area has occurred in the last decade and in regions characterized by high temperatures. Increasing use of extreme habitats may indicate that novel ecological conditions have facilitated a broader realized niche or that toad populations at the invasion front have evolved greater tolerance to extreme abiotic conditions. Rapid evolution to novel habitats combined with ecological release from native enemies may explain why some species become highly successful global invaders. Predicting species ranges following invasion or climate change may often require dynamically updated range models that incorporate a broader realization of niches in the absence of natural enemies and evolution in response to novel habitats.     

Photoperiod-temperature interaction-a new form of seasonal control of growth and development in insects and in particular a Carabid Beetle, <Emphasis Type="Italic">Amara communis</Emphasis> (Coleoptera: Carabidae)

Amara communis larvae were found to develop significantly faster and to have higher growth rate at short-day (12 h) as compared to long-day (22 h) photoperiods at all used temperatures (16, 18, 20, and 22°C). The coefficient of linear regression of larval development rate on temperature was significantly higher at the short day than at the long day. The thermal developmental thresholds appeared similar at both photoperiods. Body weight of young beetles reared under different photoperiods was almost the same. Thus, photoperiod does not simply accelerate or decelerate insect development, but modifies the thermal reaction norm. At short days, larval development becomes faster and more temperature-dependent, which provides a timely completion of development at the end of summer. The analysis of literature data has allowed us to find the photoperiodic modification of thermal requirements for development in 5 insect orders: Orthoptera, Heteroptera, Coleoptera, Lepidoptera, and Diptera. Modification may result in significant changes in the slope of the regression line, and hence the sum of degree-days, and in the thermal developmental threshold. Consequently, the thermal requirements for development in many insects gradually vary during summer under the effect of changing day-length, which may have adaptive significance. Thus, the photoperiodic modification of thermal reaction norms acts as a specific form of seasonal control of insect development.     

Flowering phenology of invasive alien plant species compared with native species in three Mediterranean-type ecosystems

Background and Aims

Flowering phenology is a potentially important component of success of alien species, since elevated fecundity may enhance invasiveness. The flowering patterns of invasive alien plant species and related natives were studied in three regions with Mediterranean-type climate: California, Spain and South Africa''s Cape region.

Methods

A total of 227 invasive–native pairs were compared for seven character types across the regions, with each pair selected on the basis that they shared the same habitat type within a region, had a common growth form and pollination type, and belonged to the same family or genus.

Key Results

Invasive alien plant species have different patterns of flowering phenology from native species in the three regions. Whether the alien species flower earlier, later or at the same time as natives depends on the climatic regime in the native range of the aliens and the proportion of species in the invasive floras originating from different regions. Species invading at least two of the regions displayed the same flowering pattern, showing that flowering phenology is a conservative trait. Invasive species with native ranges in temperate climates flower earlier than natives, those from Mediterranean-type climates at the same time, and species from tropical climates flower later. In California, where the proportion of invaders from the Mediterranean Basin is high, the flowering pattern did not differ between invasive and native species, whereas in Spain the high proportion of tropical species results in a later flowering than natives, and in the Cape region early flowering than natives was the result of a high proportion of temperate invaders.

Conclusions

Observed patterns are due to the human-induced sympatry of species with different evolutionary histories whose flowering phenology evolved under different climatic regimes. The severity of the main abiotic filters imposed by the invaded regions (e.g. summer drought) has not been strong enough (yet) to shift the flowering pattern of invasive species to correspond with that of native relatives. It does, however, determine the length of the flowering season and the type of habitat invaded by summer-flowering aliens. Results suggest different implications for impacts at evolutionary time scales among the three regions.Key words: Biological invasions, flowering phenology, genetic inertia, Cape Floristic Region, California, Spain, Mediterranean-type ecosystems, water availability, climatic origin     

Variation in phenotypic plasticity for native and invasive populations of Bromus tectorum

Phenotypic plasticity is often considered important for invasive plant success, yet relatively few studies have assessed plasticity in both native and invasive populations of the same species. We examined the plastic response to temperature for Bromus tectorum populations collected from similar shrub-steppe environments in the Republics of Armenia and Georgia, where it is native, and along an invasive front in California and Nevada. Plants were grown in growth chambers in either ‘warm’ (30/20 °C, day/night) or ‘cold’ (10/5 °C) conditions. Invasive populations exhibited greater adaptive plasticity than natives for freezing tolerance (as measured by chlorophyll a fluorescence), such that invasive populations grown in the cold treatment exhibited the highest tolerance. Invasive populations also exhibited more rapid seedling emergence in response to warm temperatures compared to native populations. The climatic conditions of population source locations were related to emergence timing for invasive populations and to freezing tolerance across all populations combined. Plasticity in growth-related traits such as biomass, allocation, leaf length, and photosynthesis did not differ between native and invasive populations. Rather, some growth-related traits were very plastic across all populations, which may help to dampen differences in biomass in contrasting environments. Thus, invasive populations were found to be particularly plastic for some important traits such as seedling emergence and freezing tolerance, but plasticity at the species level may also be an important factor in the invasive ability of B. tectorum.     

Existing and emerging high impact invasive species are characterized by higher functional responses than natives

Predicting ecological impacts of invasive species and identifying potentially damaging future invaders are research priorities. Since damage by invaders is characterized by their depletion of resources, comparisons of the ‘functional response’ (FR; resource uptake rate as a function of resource density) of invaders and natives might predict invader impact. We tested this by comparing FRs of the ecologically damaging ‘world''s worst’ invasive fish, the largemouth bass (Micropterus salmoides), with a native equivalent, the Cape kurper (Sandelia capensis), and an emerging invader, the sharptooth catfish (Clarias gariepinus), with the native river goby (Glossogobius callidus), in South Africa, a global invasion hotspot. Using tadpoles (Hyperolius marmoratus) as prey, we found that the invaders consumed significantly more than natives. Attack rates at low prey densities within invader/native comparisons reflected similarities in predatory strategies; however, both invasive species displayed significantly higher Type II FRs than the native comparators. This was driven by significantly lower prey handling times by invaders, resulting in significantly higher maximum feeding rates. The higher FRs of these invaders are thus congruent with, and can predict, their impacts on native communities. Comparative FRs may be a rapid and reliable method for predicting ecological impacts of emerging and future invasive species.     

Are invasive plant species better competitors than native plant species? – evidence from pair‐wise experiments

Invasive plants often appear to be more competitive than native species, but there have been few tests of this hypothesis. We reviewed published pair-wise experiments between invading and native plant species. Although the designs that have been used allow only limited inferences, the available data suggest that the effect of invasive species on native species is usually stronger than vice versa. Furthermore, mixtures of invasive and native species are generally less productive than monocultures of the native species, but not less than monocultures of the invasive species. However, the selection of invaders and natives for study has not been random, and the data could be biased towards highly competitive invaders and natives that are weaker than average competitors. We attempt to clarify confusion surrounding the concept of competitive superiority in the context of plant invasions, and we discuss the limitations of the methods that have been used to investigate competition between invasive and native species. To rigorously test the generality of the hypothesis that invaders are better competitors than natives we need to compare the effects of closely related native and invasive species on each other. We suggest that the influence of an invading species on total plant community biomass is an important clue in understanding the role of competition in a plant invasion. The role of competition in the establishment and naturalization stages of the invasion process may be very different from its role in the "outbreak" stage.     

Global resource acquisition patterns of invasive and native plant species do not hold at the regional scale in Mediterranean type ecosystems

Invasive species may outperform native species by acquiring more resources or by efficiently using limited resources. Studies comparing leaf traits as a metric of carbon capture strategies in native and invasive species have come to different conclusions. Some studies suggest that invasive species are better at acquiring resources, but that native and invasive species use resources similarly. Other studies have found that native and invasive species differ in resource use efficiency, which implies different biochemical or physiological mechanisms of carbon capture. To resolve this debate, we examined relationships among four leaf traits (photosynthetic rate, specific leaf area, foliar nitrogen, foliar phosphorus) in co-occurring native and invasive species from eight plant communities across five Mediterranean-climate ecosystems. We performed standardized major axis regression for all trait combinations within and across sites, testing for slope homogeneity and shifts in elevation (y-intercept) or along a common slope between species groups. Across the global dataset, native and invasive species had similar carbon capture strategies (i.e., similar slopes), with invasive species occupying a position of greater resource acquisition. However, these patterns did not hold when regions were analyzed individually. Regional differences may be driven by differences in life form between native and invasive species, and variation in soil resource availability among regions. Our context-dependent results reveal not only that management of invasive species will differ across regions but also that global comparisons of invasive and native species can be misleading.     

The invasive <Emphasis Type="Italic">Lespedeza cuneata</Emphasis> attracts more insect pollinators than native congeners in tallgrass prairie with variable impacts

Invasive plant species can potentially exert competitive or facilitative effects on insect pollination services of native species. Factors that influence these effects include the degree of shared pollinator species, synchronous flowering phenology, similar flower morphology and color, relatedness of invasive and natives, and showiness and densities of flowers. We investigated such plant-pollinator dynamics by comparing the invasive Lespedeza cuneata and three native congeners, all sympatric with synchronous flowering, using in situ populations over 2 years during peak floral displays. Insect visitation rates of the invasive were significantly higher per plant in both years than on the native species. The invasive exerted a competitive effect on visitation of the two native species with fewer shared pollinators, and a facilitative effect on visitation of the native species with the highest degree of shared insect visitors. Positive correlations were found between floral density and visitation rate per plant in all the native species. Although no such correlation was found for the invasive, floral density in L. cuneata was at least twenty times higher than in the native species and likely saturated the response of the pollinator community. Analyses of insect visitor taxonomic data indicated the insect communities visiting each of the Lespedeza species were generally similar though with species-specific differences. The main exception was that the common honeybee, Apis mellifera, was a primary visitor to the invasive plant species, yet was never observed on the native Lespedeza species.