Do exotic bumblebees and honeybees compete with native flower-visiting insects in Tasmania?

Honeybees, Apis mellifera, have been introduced by man throughout the globe. More recently, other bee species including various bumblebees (Bombus spp.) have been introduced to several new regions. Here we examine the impacts of honeybees and the bumblebee, Bombus terrestris, on native flower-visiting insects in Tasmania. To assess whether native insects have lower abundance or are excluded in areas that have been colonised by exotic bees, we quantified the abundance, diversity and floral preferences of flower-visiting insects at sites where bumblebees and honeybees were present, and compared them to sites where they were absent. This was achieved by hand searches at 67 sites, and by deploying sticky traps at 122 sites. Honeybees were by far the most abundant bee species overall, and dominated the bee fauna at most sites. There was considerable niche overlap between honeybees, bumblebees and native bees in terms of the flowers that they visited. Sites where bumblebees were established had similar species richness, diversity and abundance of native flower-visiting insects compared to sites where bumblebees were absent. In contrast, native bees were more than three times more abundant at the few sites where honeybees were absent, compared to those where they were present. Our results are suggestive of competition between honeybees and native bees, but exclusion experiments are needed to provide a definitive test.     

Competing drivers lead to non-linear native–exotic relationships in endangered temperate grassy woodlands

Relationships between the diversity and abundance of native versus exotic species underpin management of disturbance regimes for conservation. Theory predicts negative, positive or neutral relationships depending on respective drivers, with greatest potential benefit when natives and exotics show opposing responses to management. We examined drivers of exotic plant cover and relationships with native plant richness using 12-year burning, mowing and grazing experiments in two representative temperate grassy eucalypt woodlands with contrasting histories of frequent versus infrequent disturbance. We hypothesized that disturbance and high resources favour exotics, and assessed whether natives and exotics covary positively due to common external drivers or negatively due to contrasting external drivers and/or competition. Positive relationships with rainfall and disturbance explained >80 % of the variation in exotic cover at both sites, supporting our first hypothesis. Native–exotic relationships were non-linear, with native richness first increasing rapidly with increasing exotic cover, then levelling and beginning to decrease. Common external drivers, particularly inter-annual rainfall, explained initial positive relationships, highlighting a prevalence of positive relationships at long temporal (as well as large spatial) scales. At the historically frequently-burnt site, a concomitant increase in native richness and exotic cover after fire contributed to the positive relationship, indicating a management trade-off. At the long-unburnt site, exotics increased but natives decreased with fire, suggesting dual benefits of low fire frequency. We conclude that relationships between exotic cover and native richness emerge from interactions among external drivers and competitive responses, with responses to external drivers dominating at low resources and negative interactions gaining importance as resources increase.     

Do exotic generalist predators alter host plant preference of a native willow beetle?

1 Selection can favour herbivores that choose host plants benefitting their offspring either by enhancing growth rates or by increasing larval defences against native predators. For exotic predator species that feed on herbivores, their success with invading new habitats may depend upon overcoming defences used by native prey. Whether exotic predators can alter herbivore host choice has remained unexamined. Therefore, we compared the efficacy of larval defence by Chrysomela knabi (a native beetle species) that had fed on two native willow hosts: Salix sericea (a phenolic glycoside (PG)-rich species) and Salix eriocephala (a PG-poor species), when attacked by exotic generalist predators. In addition, the preference and performance of C. knabi on S. sericea and S. eriocephala was examined.
2  Chrysomela knabi preferred and performed better on S. sericea. In a common garden, adult C. knabi were nine-fold more common and oviposited five-fold more frequently on S. sericea than on S. eriocephala . In the laboratory, adult feeding preference on leaf discs and survival rates of larvae were both greater on S. sericea , and time to pupation was shorter.
3  Chrysomela knabi larvae produced significantly more salicylaldehyde when fed S. sericea leaves than when fed S. eriocephala leaves. Additionally, those larvae with greater salicylaldehyde had reduced predation by two exotic generalist predators, Harmonia axyridis larvae and juvenile Tenodera aridifolia sinensis .
4 The results obtained in the present study suggest that selection favoured the preference of C. knabi for PG-rich willow plants because larvae grew and survived better and that selection by common exotic generalist predators would reinforce this preference.     

Food web structure in exotic and native mangroves: a Hawaii–Puerto Rico comparison

Plant invasions can fundamentally alter detrital inputs and the structure of detritus-based food webs. We examined the detrital pathways in mangrove food webs in native (Puerto Rican) and introduced (Hawaiian) Rhizophora mangle forests using a dual isotope approach and a mixing model. Based on trophic-level fractionation of 0–1‰ for δ ¹³C and 2–3‰ for δ ¹⁵N, among the invertebrates, only nematodes, oligochaetes, and nereid polychaetes from native mangroves exhibited stable isotopes consistent with a mangrove-derived diet. Certain fauna, in particular tubificid oligochaetes, had δ ¹³C values consistent with the consumption of mangrove leaves, but they were depleted in ¹⁵N, suggesting their primary nitrogen source was low in ¹⁵N, and was possibly N₂-fixing bacteria. In introduced mangroves, all feeding groups appeared to rely heavily on non-mangrove sources, especially phytoplankton inputs. Mixing model results and discriminant analysis showed clear separation of introduced and native mangrove sites based on differential food source utilization within feeding groups, with stronger and more diverse use of benthic foods observed in native forests. Observed differences between native and invasive mangrove food webs may be due to Hawaiian detritivores being poorly adapted to utilizing the tannin-rich, nitrogen-poor mangrove detritus. In addition, differential utilization of mangrove detritus between native and introduced mangroves may be a consequence of forest age. We postulate that increasing mangrove forest age may promote diversification of bacterial food webs important in N and S cycling. Our results also suggest a potentially important role for sulfur bacteria in supporting the most abundant infaunal consumers, nematodes, in the most mature systems. Electronic supplementary material The online version of this article (doi:) contains supplementary material that is available to authorized users.     

Herbivore preference for native vs. exotic plants: generalist herbivores from multiple continents prefer exotic plants that are evolutionarily naïve

Enemy release and biotic resistance are competing, but not mutually exclusive, hypotheses addressing the success or failure of non-native plants entering a new region. Enemy release predicts that exotic plants become invasive by escaping their co-adapted herbivores and by being unrecognized or unpalatable to native herbivores that have not been selected to consume them. In contrast, biotic resistance predicts that native generalist herbivores will suppress exotic plants that will not have been selected to deter these herbivores. We tested these hypotheses using five generalist herbivores from North or South America and nine confamilial pairs of native and exotic aquatic plants. Four of five herbivores showed 2.4-17.3 fold preferences for exotic over native plants. Three species of South American apple snails (Pomacea sp.) preferred North American over South American macrophytes, while a North American crayfish Procambarus spiculifer preferred South American, Asian, and Australian macrophytes over North American relatives. Apple snails have their center of diversity in South America, but a single species (Pomacea paludosa) occurs in North America. This species, with a South American lineage but a North American distribution, did not differentiate between South American and North American plants. Its preferences correlated with preferences of its South American relatives rather than with preferences of the North American crayfish, consistent with evolutionary inertia due to its South American lineage. Tests of plant traits indicated that the crayfish responded primarily to plant structure, the apple snails primarily to plant chemistry, and that plant protein concentration played no detectable role. Generalist herbivores preferred non-native plants, suggesting that intact guilds of native, generalist herbivores may provide biotic resistance to plant invasions. Past invasions may have been facilitated by removal of native herbivores, introduction of non-native herbivores (which commonly prefer native plants), or both.     

A tritrophic approach to the preference–performance hypothesis involving an exotic and a native plant

Exotic plants often generate physical and chemical changes in native plant communities where they become established. A major challenge is to understand how novel plants may affect trophic interactions in their new habitats, and how native herbivores and their natural enemies might respond to them. We compared the oviposition preference and offspring performance of the crucifer specialist, Pieris brassicae, on an exotic plant, Bunias orientalis, and on a related native plant, Sinapis arvensis. Additionally, we studied the response of the parasitoid, Cotesia glomerata to herbivore-induced plant volatiles (HIPV) and determined the volatile blend composition to elucidate which compound(s) might be involved in parasitoid attraction. On both host plants we also compared the parasitism rate of P. brassicae by C. glomerata. Female butterflies preferred to oviposit on the native plant and their offspring survival and performance was higher on the native plant compared to the exotic. Although, headspace analysis revealed qualitative and quantitative differences in the volatile blends of both plant species, C. glomerata did not discriminate between the HIPV blends in flight-tent bioassays. Nevertheless, parasitism rate of P. brassicae larvae was higher on the native plant under semi-field conditions. Overall, P. brassicae oviposition preference may be more influenced by bottom-up effects of the host plant on larval performance than by top-down pressure exerted by its parasitoid. The potential for dietary breadth expansion of P. brassicae to include the exotic B. orientalis and the role of top-down processes played by parasitoids in shaping herbivore host shifts are further discussed.     

Enhancing gardens as habitats for plant-associated invertebrates: should we plant native or exotic species?

Domestic gardens, typically consisting of a mixture of native and non-native plants, support biodiversity. The relative value of these native and non-native plants for invertebrates is largely unknown. To address this a replicated field experiment with plots planted with one of three assemblages of non-invasive perennial and shrubby garden plants (treatments), based on plant origin [UK native, near-native (Northern Hemisphere) and Exotic (Southern Hemisphere)] was established. Over 4 years the invertebrates were recorded by Vortis suction sampler and amount of plant material measured. The abundance of above ground plant-inhabiting invertebrates increased with canopy cover and was higher on the native treatment. For several functional groups including herbivores and some predatory groups the near-native plants supported only marginally fewer individuals compared to native plots, with exotic plants being less favoured. The experiment demonstrated that gardens and other cultivated ornamental plantings support a wide range of plant-inhabiting invertebrates from primary functional groups regardless of the plants’ origin and the more plant matter (canopy cover) available the greater the abundance. Greater abundance of invertebrates will be supported by gardens and cultivated planting schemes with plantings biased towards native and near-native plants and that provide dense vegetation cover. However, exotic plants should not be dismissed as these are inhabited by some invertebrates.     

Forest fragment size and nutrient availability: complex responses of mycorrhizal fungi in native–exotic hosts

In the past few decades, it has been widely accepted that forest loss due to human actions alter the interactions between organisms. We studied the relationship between forest fragment size and arbuscular mycorrhizal fungi (AMF) and dark septate endophytes (DSE) colonization, and the AMF spore communities in the rhizosphere of two congeneric Euphorbia species (native and exotic/invasive). We hypothesized that these fungal variables will differ with fragment size and species status, and predicted that (a) AMF and DSE colonization together with AMF spore abundance and diversity would be positively related to forest fragment size; (b) these relationships will differ between the exotic and the native species; and (c) there will be a negative relationship between forest fragment size and the availability of soil nutrients (NH₄ ⁺, NO₃ ⁻, and phosphorus). This study was performed in the eight randomly selected forest fragments (0.86–1000 ha), immersed in an agricultural matrix from the Chaquean region in central Argentina. AMF root colonization in the native and exotic species was similar, and was positively related with forest fragment size. Likewise, AMF spore diversity and spore abundance were higher in the larger fragments. While DSE root colonization in the native host was positively related with forest fragment size, DSE colonization in the exotic host showed no relationship. Soil nutrients contents were negatively related with forest fragment size. In addition, NH₄ ⁺ and NO₃ ⁻ were negatively correlated with AMF spores abundance and root colonization and with DSE colonization in the native species. The results observed in this study show how habitat fragmentation might affect the interaction between key soil components, such as rhizospheric plant-fungal symbiosis and nutrient availability. These environmental changes may have important consequences on plant community composition and nutrient dynamics in this fragmented landscape.     

Intraspecific and interspecific pair-wise seedling competition between exotic annual grasses and native perennials: plant–soil relationships

Few studies have examined plant–soil relationships in competitive arenas between exotic and native plants in the western United States. A pair-wise competitive design was used to evaluate plant–soil relationships between seedlings of the exotic annual grasses Bromus tectorum and Taeniatherium caput-medusae and the native perennial grasses Elymus elymoides and Pseudoroegneria spicata. Two soils were tested: an arid soil (argid) occupied by E. elymoides and presently invaded by B. tectorum and a high elevation, high organic matter, soil (aquept) where none of the tested species would typically occur. Plant growth proceeded for 85 days at which time above-ground biomass and tissue nutrient concentrations were quantified. Soil also was collected from the rooting zone beneath each species and analyzed for various nutrient pools. The exotic species had significantly greater above-ground biomass than the natives and grew far better in the aquept soil than the argid soil. Growth of B. tectorum, and to some degree, T. caput-medusae was suppressed in intraspecific competition and enhanced, especially in the aquept soil, when competing with the natives. Although not significant, biomass of natives strongly trended downward when competing with the exotic grasses. Overall, concentrations of tissue nutrients were minimally affected by competition, but natives tended to be more negatively affected by competition with exotics. Except for phosphorus (P), all species had significantly greater nutrient concentrations when growing in the aquept soil compared to the argid soil. In both soils, exotics had significant greater tissue concentrations of manganese (Mn), magnesium (Mg), and iron (Fe), while natives had significantly greater nitrogen (N). Species affects on soil nutrient pools occurred mostly in the aquept soil with exotic species significantly decreasing pools of available N, potentially available N, and soil-solution pools of calcium (Ca²⁺), potassium (K⁺), and magnesium (Mg²⁺) relative to natives. Overall, the data suggest that, in the seedling state, B. tectorum is a superior competitor. Moreover, when the natives compete intra- or interspecifically, particularly in the aquept soil, availability of N and other nutrients in their rooting zone is consistently greater than when they compete interspecifically with the exotic grasses. These data suggest the exotics are able to co-opt nutrients in the rooting zone of the natives and perhaps gain a competitive advantage.     

Colilargo’s occupancy and the role of native and exotic plants in hantavirus expansion and transmission risk

Mammal Research - In this work, we examined the relationship of Oligoryzomys longicaudatus’ (the Andes virus [ANDV] host, commonly known as the colilargo) occupancy and the cover of dominant...     

Do heat and smoke increase emergence of exotic and native plants in the matorral of central Chile?

We studied the effect of heat shock and wood-fueled smoke on the emergence of native and exotic plant species in soil samples obtained in an evergreen shrubland of central Chile, located on the eastern foothills of the Coastal Range of Lampa. Immediately after collection samples were dried and stored under laboratory condition. For each two transect, 10 samples were randomly chosen, and one of the following treatments was applied: 1) Heat-shock treatment. 2) Plant-produced smoke treatment. 3) Combined heat-and-smoke treatment. 4) Control, corresponding to samples not subjected to treatment. Twenty-three species, representing 12 families, emerged from the soil samples. The best-represented families were Poaceae and Asteraceae. All of the emerged species were herbs, 21 were annuals, and 14 were exotic to Chile. Fire-related triggers used in this study did not increase the emergence and/or abundance of exotic species with respect to natives in soil samples. Interestingly, this study provides evidence that heat-shock can increase the emergence of native herbs.     

Plantations of exotic tree species in Britain: irrelevant for biodiversity or novel habitat for native species?

Novel or emergent ecosystems arising from human action present both threats and opportunities for biodiversity. It has been suggested that exotic species can “facilitate” or “inhibit” native biodiversity through habitat modification. In Britain, there is a discussion over the contribution to biodiversity of plantations of exotic conifer species as these are commonly thought to have little relevancy as a habitat for native biodiversity. To address this we compared the species richness of a range of different taxonomic groups (lichens, bryophytes, fungi, vascular plants, invertebrates and songbirds) in exotic and native forest stands of differing structural stages in northern and southern Britain. In terms of overall native species-richness there was no significant difference between the exotic and the native stands. In the north, six species groups showed higher values in the exotic Sitka spruce (Picea sitchensis) stands with the remaining six showing higher values in the native Scots pine (Pinus sylvestris) stands. Most notably, lichen species richness was much lower in the exotic stands compared to the native stands, whereas bryophyte and fungal species richness was proportionately higher in the exotic stands. In the south, five species groups (all invertebrate taxa) showed higher species richness in exotic Norway spruce (Picea abies) stands compared to native oak (Quercus robur) stands. Five species groups had higher species-richness in the oak stands, in particular lichens and fungi. It is concluded that emergent ecosystems of exotic conifer species are not irrelevant to biodiversity. Where already well-established they can provide habitat for native species particularly if native woodland is scarce and biodiversity restoration is an immediate priority.     

Enemy release or invasional meltdown? Deer preference for exotic and native trees on Isla Victoria,Argentina

Abstract How interactions between exotic species affect invasion impact is a fundamental issue on both theoretical and applied grounds. Exotics can facilitate establishment and invasion of other exotics (invasional meltdown) or they can restrict them by re‐establishing natural population control (as predicted by the enemy‐release hypothesis). We studied forest invasion on an Argentinean island where 43 species of Pinaceae, including 60% of the world's recorded invasive Pinaceae, were introduced c. 1920 but where few species are colonizing pristine areas. In this area two species of Palearctic deer, natural enemies of most Pinaceae, were introduced 80 years ago. Expecting deer to help to control the exotics, we conducted a cafeteria experiment to assess deer preferences among the two dominant native species (a conifer, Austrocedrus chilensis, and a broadleaf, Nothofagus dombeyi) and two widely introduced exotic tree species (Pseudotsuga menziesii and Pinus ponderosa). Deer browsed much more intensively on native species than on exotic conifers, in terms of number of individuals attacked and degree of browsing. Deer preference for natives could potentially facilitate invasion by exotic pines. However, we hypothesize that the low rates of invasion currently observed can result at least partly from high densities of exotic deer, which, despite their preference for natives, can prevent establishment of both native and exotic trees. Other factors, not mutually exclusive, could produce the observed pattern. Our results underscore the difficulty of predicting how one introduced species will effect impact of another one.     

Invading from the garden? A comparison of leaf herbivory for exotic and native plants in natural and ornamental settings

Abstract The enemies release hypothesis proposes that exotic species can become invasive by escaping from predators and parasites in their novel environment. Agrawal et al. (Enemy release? An experiment with congeneric plant pairs and diverse above‐ and below‐ground enemies. Ecology, 86, 2979–2989) proposed that areas or times in which damage to introduced species is low provide opportunities for the invasion of native habitat. We tested whether ornamental settings may provide areas with low levels of herbivory for trees and shrubs, potentially facilitating invasion success. First, we compared levels of leaf herbivory among native and exotic species in ornamental and natural settings in Cincinnati, Ohio, United States. In the second study, we compared levels of herbivory for invasive and noninvasive exotic species between natural and ornamental settings. We found lower levels of leaf damage for exotic species than for native species; however, we found no differences in the amount of leaf damage suffered in ornamental or natural settings. Our results do not provide any evidence that ornamental settings afford additional release from herbivory for exotic plant species.     

Patterns of native and exotic species richness in the urban flora of Brussels: rejecting the ‘rich get richer’ model

In this study we analyzed patterns of native and exotic species richness in the urban flora of Brussels (Belgium) using a coarse-scale systematic sampling grid of 1 km². The observed correlation between native and exotic richness within the grid cells sampled was then compared to the results of an adequate null model assuming no species interactions. In addition, ordinary least-squares and quantile regressions were used to analyze the relationship between the ratio of exotics to natives and the proportion of densely built up areas in each cell. Though the results obtained conform to the Eltonian expectation that exotic species preferably invade areas of low native species diversity, traditional niche-filling mechanisms seems inadequate to explain the observed pattern. Rather, aliens simply tend to have different environmental requirements than natives.     

Are differences in productivity between native and exotic trees in N.W. Patagonia related to differences in hydraulic conductance?

Afforestation with the exotic Pinus ponderosa is currently taking place within the natural distributional area of Austrocedrus chilensis, a native conifer of N.W. Patagonia. Annual productivity of the exotic species is double the productivity of the native one. In order to test the hypothesis that these differences in productivity are, at least in part, due to differences in hydraulic characteristics of both species, we measured or estimated several ecophysiological variables in A. chilensis and P. ponderosa trees growing in the same place. Water use (WU) and diameter growth were lower in A. chilensis than in P. ponderosa. Although predawn water potential was relatively constant during the whole growing season, A. chilensis trees showed lower values of this variable than P. ponderosa in a very dry period, suggesting different water sources. Under field conditions, canopy-stomatal (gs) and whole hydraulic conductances, specific hydraulic conductivity and photosynthetic rate (A) were lower in A. chilensis than P. ponderosa. In contrast, instantaneous WU efficiency was higher in A. chilensis than in P. ponderosa. However, gs and A in A. chilensis significantly increased in cut branches of this species suggesting hydraulic limitations on photosynthesis. We hypothesize that hydraulic characteristics of P. ponderosa permit high stomatal conductance for more hours a day than A. chilensis trees, without reaching threshold values of water potential. This can explain, at least in part, differences in C fixation and thus, in productivity between species. In addition, our results suggested a secondary limitation to C fixation in A. chilensis at the photosynthetic apparatus.     

Does salt stress increase the ability of the exotic legume Acacia longifolia to compete with native legumes in sand dune ecosystems?

Sand dune ecosystems are one of the areas most affected by the introduction of invasive species which represents a threat for biodiversity conservation. Their invasion patterns and spread may depend on their salinity tolerance, besides other factors. To test this hypothesis, we investigated the effects of salt stress on seed germination and on the activity of antioxidant enzymes (catalase, CAT; ascorbate peroxidase, APX; peroxidase, POX; and glutathione reductase, GR) in two legume species, an invasive, Acacia longifolia (Andrews.) Willd., and a native, Ulex europaeus (L.), very common in the sand dunes of the coast of Portugal. Salt stress was induced by adding NaCl at different concentrations, 0, 50, 100 and 200 mM, for 15 days. Results showed that the highest germination percentages were obtained in distilled water (control) and that, with increasing salt concentration, seed germination was delayed and decreased in both species. Inhibition of germination was higher in the native species, only 3% of seeds germinated at 100 mM and no seeds germinated at 200 mM NaCl. In the invasive species, the reduction was higher at 200 mM NaCl (16%). Considering the coefficient of germination velocity, a decrease in both species with increasing NaCl concentration was observed. The CAT and GR activities decreased in A. longifolia with increasing salinity. In turn, APX activity significantly increased as NaCl concentration increased while the POX activities declined at the highest NaCl concentration. On the other hand, at 50 mM NaCl lower activity of CAT and APX and higher GR and POX were found in U. europaeus. In both species, protein content increased as NaCl concentration increased. In addition, it seems that APX activities play an essential role in the scavenging reactive oxygen species (ROS). These results suggest that the seeds of the invasive legume A. longifolia are more tolerant to salinity than the native legume U. europaeus, and seem better equipped to handle the physiological stress of high salinity, which may contribute to its invasive ability in sand dunes.