Biotic and abiotic controls of Argentine ant invasion success at local and landscape scales

Although the ecological success of introduced species hinges on biotic interactions and physical conditions, few experimental studies--especially on animals--have simultaneously investigated the relative importance of both types of factors. The lack of such research may stem from the common assumption that native and introduced species exhibit similar environmental tolerances. Here we combine experimental and spatial modeling approaches (1) to determine the relative importance of biotic and abiotic controls of Argentine ant (Linepithema humile) invasion success, (2) to examine how the importance of these factors changes with spatial scale in southern California (USA), and (3) to assess how Argentine ants differ from native ants in their environmental tolerances. A factorial field experiment that combined native ant removal with irrigation revealed that Argentine ants failed to invade any dry plots (even those lacking native ants) but readily invaded all moist plots. Native ants slowed the spread of Argentine ants into irrigated plots but did not prevent invasion. In areas without Argentine ants, native ant species showed variable responses to irrigation. At the landscape scale, Argentine ant occurrence was positively correlated with minimum winter temperature (but not precipitation), whereas native ant diversity increased with precipitation and was negatively correlated with minimum winter temperature. These results are of interest for several reasons. First, they demonstrate that fine-scale differences in the physical environment can eclipse biotic resistance from native competitors in determining community susceptibility to invasion. Second, our results illustrate surprising complexities with respect to how the abiotic factors limiting invasion can change with spatial scale, and third, how native and invasive species can differ in their responses to the physical environment. Idiosyncratic and scale-dependent processes complicate attempts to forecast where introduced species will occur and how their range limits may shift as a result of climate change.     

Advertising to the enemy: enhanced floral fragrance increases beetle attraction and reduces plant reproduction

Many organisms face challenges in avoiding predation while searching for mates. For plants, emitting floral fragrances to advertise reproductive structures could increase the attraction of detrimental insects along with pollinators. Very few studies have experimentally evaluated the costs and benefits of fragrance emission with explicit consideration of how plant fitness is affected by both pollinators and florivores. To determine the reproductive consequences of increasing the apparency of reproductive parts, we manipulated fragrance, pollination, and florivores in the wild Texas gourd, Cucurbita pepo var. texana. With enhanced fragrance we found an increase in the attraction of florivores, rather than pollinators, and a decrease in seed production. This study is the first to demonstrate that enhanced floral fragrance can increase the attraction of detrimental florivores and decrease plant reproduction, suggesting that florivory as well as pollination has shaped the evolution of floral scent.     

Effects of vegetation cover, presence of a native ant species, and human disturbance on colonization by Argentine ants

The spread of non-native invasive species is affected by human activity, vegetation cover, weather, and interaction with native species. We analyzed data from a 17-year study of the distribution of the non-native Argentine ant (Linepithema humile) and the native winter ant (Prenolepis imparis) in a preserve in northern California (U.S.A.). We conducted logistic regressions and used model selection to determine whether the following variables were associated with changes in the distribution of each species: presence of conspecifics at neighboring sites, distance to development (e.g., roads, buildings, and landscaped areas), proportion of vegetation cover taller than 0.75 m, elevation, distance to water, presence of both species at a site, temperature, and rainfall. Argentine ants colonized unoccupied sites from neighboring sites, but the probability of appearance and persistence decreased as distance to development, vegetation cover, and elevation increased. Winter ants appeared and persisted in sites with relatively high vegetation cover (i.e., highly shaded sites). Presence of the 2 species was negatively associated in sites with high vegetation cover (more winter ants) and sites near development (more Argentine ants). Probability of colonization of Argentine ants decreased where winter ants were most persistent. At sites near development within the preserve, abundant Argentine ant populations may be excluding winter ants. The high abundance of Argentine ants at these sites may be due to immigration from suburban areas outside the preserve, which are high-quality habitat for Argentine ants. In the interior of the preserve, distance from development, low-quality habitat, and interaction with winter ants may in combination exclude Argentine ants. Interactions among the variables we examined were associated with low probabilities of Argentine ant colonization in the preserve.     

Herbivory-mediated pollinator limitation: negative impacts of induced volatiles on plant-pollinator interactions

Although induced plant responses to herbivory are well studied as mechanisms of resistance, how induction shapes community interactions and ultimately plant fitness is still relatively unknown. Using a wild tomato, Solanum peruvianum, native to the Peruvian Andes, we evaluated the disruption of pollination as a potential ecological cost of induced responses. More specifically, we tested the hypothesis that metabolic changes in herbivore-attacked plants, such as the herbivore-induced emission of volatile organic compounds (VOCs), alter pollinator behavior and consequentially affect plant fitness. We conducted a series of manipulative field experiments to evaluate the role of herbivore-induced vegetative and floral VOC emissions as mechanisms by which herbivory affects pollinator behavior. In field surveys and bioassays in the plants' native habitat, we found that real and simulated herbivory (methyl jasmonate application) reduced attractiveness of S. peruvianum flowers to their native pollinators. We show that reduced pollinator preference, not resource limitation due to leaf tissue removal, resulted in reduced seed set. Solitary bee pollinators use floral plant volatiles, emitted in response to herbivory or methyl jasmonate treatment, as cues to avoid inflorescences on damaged plants. This herbivory-induced pollinator limitation can be viewed as a general cost of induced plant responses as well as a specific cost of herbivory-induced volatile emission.     

Separating selection by diurnal and nocturnal pollinators on floral display and spur length in Gymnadenia conopsea

Most plants attract multiple flower visitors that may vary widely in their effectiveness as pollinators. Floral evolution is expected to reflect interactions with the most important pollinators, but few studies have quantified the contribution of different pollinators to current selection on floral traits. To compare selection mediated by diurnal and nocturnal pollinators on floral display and spur length in the rewarding orchid Gymnadenia conopsea, we manipulated the environment by conducting supplemental hand-pollinations and selective pollinator exclusions in two populations in central Norway. In both populations, the exclusion of diurnal pollinators significantly reduced seed production compared to open pollination, whereas the exclusion of nocturnal pollinators did not. There was significant selection on traits expected to influence pollinator attraction and pollination efficiency in both the diurnal and nocturnal pollination treatment. The relative strength of selection among plants exposed to diurnal and nocturnal visitors varied among traits and populations, but the direction of selection was consistent. The results suggest that diurnal pollinators are more important than nocturnal pollinators for seed production in the study populations, but that both categories contribute to selection on floral morphology. The study illustrates how experimental manipulations can link specific categories of pollinators to observed selection on floral traits, and thus improve our understanding of how species interactions shape patterns of selection.     

Density-mediated, context-dependent consumer-resource interactions between ants and extrafloral nectar plants

Interspecific interactions are often mediated by the interplay between resource supply and consumer density. The supply of a resource and a consumer's density response to it may in turn yield context-dependent use of other resources. Such consumer-resource interactions occur not only for predator-prey and competitive interactions, but for mutualistic ones as well. For example, consumer-resource interactions between ants and extrafloral nectar (EFN) plants are often mutualistic, as EFN resources attract and reward ants which protect plants from herbivory. Yet, ants also commonly exploit floral resources, leading to antagonistic consumer-resource interactions by disrupting pollination and plant reproduction. EFN resources associated with mutualistic ant-plant interactions may also mediate antagonistic ant-flower interactions through the aggregative density response of ants on plants, which could either exacerbate ant-flower interactions or alternatively satiate and distract ants from floral resources. In this study, we examined how EFN resources mediate the density response of ants on senita cacti in the Sonoran Desert and their context-dependent use of floral resources. Removal of EFN resources reduced the aggregative density of ants on plants, both on hourly and daily time scales. Yet, the increased aggregative ant density on plants with EFN resources decreased rather than increased ant use of floral resources, including contacts with and time spent in flowers. Behavioral assays showed no confounding effect of floral deterrents on ant-flower interactions. Thus, ant use of floral resources depends on the supply of EFN resources, which mediates the potential for both mutualistic and antagonistic interactions by increasing the aggregative density of ants protecting plants, while concurrently distracting ants from floral resources. Nevertheless, only certain years and populations of study showed an increase in plant reproduction through herbivore protection or ant distraction from floral resources. Despite pronounced effects of EFN resources mediating the aggregative density of ants on plants and their context-dependent use of floral resources, consumer-resource interactions remained largely commensalistic.     

Predispersal seed herbivores, not pollinators, exert selection on floral traits via female fitness

Herbivores that oviposit in flowers of animal-pollinated plants depend on pollinators for seed production and are therefore expected to choose flowers that attract pollinators. This provides a mechanism by which seed herbivores and pollinators could impose conflicting selection on floral traits. We measured phenotypic selection on floral traits of Lobelia siphilitica (Lobeliaceae) via female fitness to determine the relative strength of selection by pollinators and a specialist predispersal seed herbivore. We were able to attribute selection on flowering phenology to the herbivores. However, no selection could be attributed to pollinators, resulting in no conflicting selection on floral traits. Unlike pollinators, whose preference for certain floral traits does not always translate into higher fitness, any discrimination by seed herbivores is likely to decrease fitness of the preferred floral phenotype. Thus predispersal seed herbivores may be a significant agent of selection on floral traits.     

Mutualism as reciprocal exploitation: African plant-ants defend foliar but not reproductive structures

The foundation of many plant-ant mutualisms is ant protection of plants from herbivores in exchange for food and/or shelter. While the role of symbiotic ants in protecting plants from stem- and leaf-feeding herbivores has been intensively studied, the relationship between ant defense and measures of plant fitness has seldom been quantified. We studied ant aggression, damage by herbivores and seed predators, and fruit production among Acacia drepanolobium trees occupied by four different acacia-ant species in an East African savanna. Levels of ant aggression in response to experimental disturbance differed strongly among the four species. All four ant species recruited more strongly to new leaf growth on host plants following disturbance, while recruitment to developing fruits was on average an order of magnitude lower. Host plants occupied by more aggressive ant species suffered significantly less vegetative damage from leaf-feeding insects, stem-boring beetles, and vertebrate browsers than host plants occupied by less aggressive ant species. However, there were no differences among fruiting host plants occupied by different ant species in levels of seed predation by bruchid seed predators. Fruit production on host trees was significantly correlated with tree stem diameter but not with the identity of resident ants. Our results demonstrate that defense of host plants may differ substantially among ant species and between vegetative and reproductive structures and that fruit production is not necessarily correlated with high levels of aggression by resident ants.     

Mimics and magnets: the importance of color and ecological facilitation in floral deception

Plants that lack floral rewards can attract pollinators if they share attractive floral signals with rewarding plants. These deceptive plants should benefit from flowering in close proximity to such rewarding plants, because pollinators are locally conditioned on floral signals of the rewarding plants (mimic effect) and because pollinators are more abundant close to rewarding plants (magnet effect). We tested these ideas using the non-rewarding South African plant Eulophia zeyheriana (Orchidaceae) as a study system. Field observations revealed that E. zeyheriana is pollinated solely by solitary bees belonging to a single species of Lipotriches (Halictidae) that appears to be closely associated with the flowers of Wahlenbergia cuspidata (Campanulaceae), a rewarding plant with which the orchid is often sympatric. The pale blue color of the flowers of E. zeyheriana differs strongly from flowers of its congeners, but is very similar to that of flowers of W. cuspidata. Analysis of spectral reflectance patterns using a bee vision model showed that bees are unlikely to be able to distinguish the two species in terms of flower color. A UV-absorbing sunscreen was applied to the flowers of the orchid in order to alter their color, and this resulted in a significant decline in pollinator visits, thus indicating the importance of flower color for attraction of Lipotriches bees. Pollination success in the orchid was strongly affected by proximity to patches of W. cuspidata. This was evident from one of two surveys of natural populations of the orchid, as well as experiments in which we translocated inflorescences of the orchid either into patches of W. cuspidata or 40 m outside such patches. Flower color and location of E. zeyheriana plants relative to rewarding magnet patches are therefore key components of the exploitation by this orchid of the relationship between W. cuspidata and Lipotriches bee pollinators.     

Homopteran chemical signatures reduce aggression of tending ants

Summary. Because generalist ants are aggressive towards foreign insects, the recognition of homopterans by tending ants is critical in ant/homopteran trophobiosis. Herein we report experimental evidence indicating that Argentine ant, Linepithema humile (Mayr) (Hymenoptera: Formicidae) learn to associate the production of honeydew with the chemical characteristics of homopteran cuticle, suppressing ant aggression and allowing the ants to tend homopterans. Although chemically-mediated associative learning is well understood in honeybee foraging, to our knowledge, it has not been reported before in ant/homopteran trophobiosis.     

Facilitation in an insect-pollinated herb with a floral display dimorphism

Population context should influence pollination success and selection on floral display in animal-pollinated plants because attraction of pollinators depends not only on the characteristics of individual plants, but also on the attractiveness of co-occurring conspecifics. The insect-pollinated herb Primula farinosa is polymorphic for inflorescence height. Natural populations may include both long-scaped plants, which present their flowers well above the soil surface, and short-scaped plants, with their flowers positioned close to the ground. We experimentally tested whether seed production in short-scaped P. farinosa varied with local morph frequency and surrounding vegetation height. In tall vegetation, short-scaped plants in polymorphic populations produced more fruit and tended to produce more seeds than short-scaped plants did in monomorphic populations. In low vegetation, population composition did not significantly affect fruit and seed output of short-scaped plants. The results suggest that long-scaped plants facilitate short-scaped plants in terms of pollinator attraction and that the facilitation effect is contingent on the height of the surrounding vegetation. The documented facilitation should contribute to the maintenance of the scape length polymorphism in ungrazed areas where litter accumulates and vegetation grows tall.     

Density-dependent effects of ants on selection for bumble bee pollination in Polemonium viscosum

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% +/- 6% [mean +/- SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20-80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum.     

Trait-mediated effects on flowers: artificial spiders deceive pollinators and decrease plant fitness

Although predators can affect foraging behaviors of floral visitors, rarely is it known if these top-down effects of predators may cascade to plant fitness through trait-mediated interactions. In this study we manipulated artificial crab spiders on flowers of Rubus rosifolius to test the effects of predation risk on flower-visiting insects and strength of trait-mediated indirect effects to plant fitness. In addition, we tested which predator traits (e.g., forelimbs, abdomen) are recognized and avoided by pollinators. Total visitation rate was higher for control flowers than for flowers with an artificial crab spider. In addition, flowers with a sphere (simulating a spider abdomen) were more frequently visited than those with forelimbs or the entire spider model. Furthermore, the presence of artificial spiders decreased individual seed set by 42% and fruit biomass by 50%. Our findings indicate that pollinators, mostly bees, recognize and avoid flowers with predation risk; forelimbs seem to be the predator trait recognized and avoided by hymenopterans. Additionally, predator avoidance by pollinators resulted in pollen limitation, thereby affecting some components of plant fitness (fruit biomass and seed number). Because most pollinator species that recognized predation risk visited many other plant species, trait-mediated indirect effects of spiders cascading down to plant fitness may be a common phenomenon in the Atlantic rainforest ecosystem.     

Ecological and evolutionary mechanisms for low seed: ovule ratios: need for a pluralistic approach?

Central to the ecology and evolution of a broad range of plants is understanding why they routinely have submaximal reproduction manifested as low seed : ovule and fruit : flower ratios. We know much less about the processes responsible for low seed : ovule ratios than we do for fruit : flower ratios. Current hypotheses for low seed : ovule ratios are largely drawn from those for fruit : flower ratios, including proximate (ecological) causes of pollen limitation, resource limitation, and pollen quality, as well as the ultimate (evolutionary) hypothesis of "bet hedging" on stochastic pollination. Yet, such mechanisms operating on fruit : flower ratios at the whole-plant level may not best explain low seed : ovule ratios at the individual-flower level. We tested each of these proximate and ultimate causes for low seed : ovule ratios using the specialized pollination mutualism between senita cacti (Pachycereus schottii) and senita moths (Upiga virescens). Seed : ovule ratios were consistently low (approximately 0.61). Such excess ovule production by senita likely has a strong genetic component given the significant differences among plants in ovule number and the consistency in ovule production by plants within and among flowering seasons. Excess ovule production and low seed : ovule ratios could not be explained by pollen limitation, resource limitation, pollen quality, or bet hedging. Nevertheless, phenotypic selection analyses did show significant selection gradients for increased ovule number, suggesting that other evolutionary processes may be responsible for excess ovule production and low seed : ovule ratios. In contrast, low fruit : flower ratios at the whole-plant level were explained by an apparent equilibrium between pollen and resource limitation. Thus, mechanisms responsible for low fruit : flower ratios at the whole-plant level are not necessarily in accord with those of low seed : ovule ratios at the individual-flower level. This suggests that we may need to adopt a more pluralistic approach to seed : ovule ratios and consider alternative hypotheses, including a greater array of proximate and ultimate causes. Initial results of this study suggest that floral allometry, selection on correlated floral traits, stigma clogging with pollen grains, and style clogging with pollen tubes may provide promising avenues for understanding low seed : ovule ratios.     

Ants as a Measure of Effectiveness of Habitat Conservation Planning in Southern California

Abstract: In the United States multispecies habitat conservation plans were meant to be the solution to conflicts between economic development and protection of biological diversity. Although now widely applied, questions exist concerning the scientific credibility of the conservation planning process and effectiveness of the plans. We used ants to assess performance of one of the first regional conservation plans developed in the United States, the Orange County Central‐Coastal Natural Community Conservation Plan (NCCP), in meeting its broader conservation objectives of biodiversity and ecosystem‐level protection. We collected pitfall data on ants for over 3 years on 172 sites established across a network of conservation lands in coastal southern California. Although recovered native ant diversity for the study area was high, site‐occupancy models indicated the invasive and ecologically disruptive Argentine ant (Linepithema humile) was present at 29% of sites, and sites located within 200 m of urban and agricultural areas were more likely to have been invaded. Within invaded sites, native ants were largely displaced, and their median species richness declined by more than 60% compared with uninvaded sites. At the time of planning, 24% of the 15,133‐ha reserve system established by Orange County NCCP fell within 200 m of an urban or agricultural edge. With complete build out of lands surrounding the reserve, the proportion of the reserve system vulnerable to invasion will grow to 44%. Our data indicate that simply protecting designated areas from development is not enough. If habitat conservation plans are to fulfill their conservation promise of ecosystem‐level protection, a more‐integrated and systematic approach to the process of habitat conservation planning is needed.     

Balancing between mutualism and exploitation: the symbiotic interaction between Lasius ants and aphids

Nests of Lasius niger (L.) ants were given varied food regimens to test whether their behaviour towards an aphid partner, Aphis fabae (Scop.), changed with alternative food supplies. Honeydew collection and predation on aphids were measured by video monitoring the movement of ants between their nest and an aphid aggregation. Data collected from the aphid aggregations enabled comparisons between remaining aphid biomass and between the tending intensities of the ants. I tested how ant behaviour was influenced by their access to alternative prey and sugar. The results showed that ants accepted a honey solution as a substitute for the honeydew produced by aphids. Ants not only attended their aphid partners, but also preyed on them. The average predation rate increased eightfold when ants were offered the alternative of sugar, whereas alternative prey had no significant effect. In contrast, ant-tending intensity decreased with alternative sugar whereas alternative prey elicited no effect.     

Evolution of polyploidy and the diversification of plant-pollinator interactions

One of the major mechanisms of plant diversification has been the evolution of polyploid populations that differ from their diploid progenitors in morphology, physiology, and environmental tolerances. Recent studies have indicated that polyploidy may also have major effects on ecological interactions with herbivores and pollinators. We evaluated pollination of sympatric diploid and tetraploid plants of the rhizomatous herb Heuchera grossulariifolia (Saxifragaceae) along the Selway and Salmon Rivers of northern Idaho, USA, during four consecutive years. Previous molecular and ecological analyses had indicated that the tetraploid populations along these two river systems are independently derived and differ from each other in multiple traits. In each region, we evaluated floral visitation rate by all insect visitors, pollination efficacy of all major visitors, and relative contribution of all major pollinators to seed set. In both regions, diploid and tetraploid plants attracted different suites of floral visitors. Most pollination was attributable to several bee species and the moth Greya politella. Lasioglossum bees preferentially visited diploid plants at Lower Salmon but not at Upper Selway, queen Bombus centralis preferentially visited tetraploids at both sites, and worker B. centralis differed between sites in their cytotype preference. Hence, diploid and autotetraploid H. grossulariifolia plants act essentially as separate ecological species and may experience partial reproductive isolation through differential visitation and pollination by their major floral visitors. Overall the results, together with recent results from other studies, suggest that the repeated evolution of polyploidy in plants may contribute importantly to the structure and diversification of ecological interactions in terrestrial communities.     

Defensive behavior of ants in a mutualistic relationship with aphids

Mutualistic relationships between ants and aphids are well studied but it is unknown if aphid-attending ants place a greater relative importance on defending aphids from aphid-predators or from competing ant colonies. We tested the hypothesis that aphid-attending ants defend their aphids against aphid-predators more aggressively than against ants from neighboring colonies. We conducted introduction trials by placing an individual non-predatory insect, an aphid-predator, or a foreign conspecific ant on the leaf of a resident ant. We found that ants did not attack non-predatory insects, but did attack competing ants and aphid-predators. When we presented resident ants with both the threats (i.e., predator and competitor) at the same time, residents always attacked potential competitors as opposed to aphid-predators. We suggest this behavior may reduce the likelihood of raids by neighboring colonies. Ants appear to balance both the energetic costs of making an attack and the costs associated with losing aphids to a predator, against the benefits of signaling their defensive ability to rivals and/or preventing rivals from gaining knowledge of a potential food resource.     

Relative Effects of Disturbance on Red Imported Fire Ants and Native Ant Species in a Longleaf Pine Ecosystem

Abstract: The degree to which changes in community composition mediate the probability of colonization and spread of non‐native species is not well understood, especially in animal communities. High species richness may hinder the establishment of non‐native species. Distinguishing between this scenario and cases in which non‐native species become established in intact (lacking extensive anthropogenic soil disturbance) communities and subsequently diminish the abundance and richness of native species is challenging on the basis of observation alone. The red imported fire ant (Solenopsis invicta), an invasive species that occurs throughout much of the southeastern United States, is such an example. Rather than competitively displacing native species, fire ants may become established only in disturbed areas in which native species richness and abundance are already reduced. We used insecticide to reduce the abundance of native ants and fire ants in four experimental plots. We then observed the reassembly and reestablishment of the ants in these plots for 1 year after treatment. The abundance of fire ants in treated plots did not differ from abundance in control plots 1 year after treatment. Likewise, the abundance of native ants increased to levels comparable to those in control plots after 1 year. Our findings suggest that factors other than large reductions in ant abundance and species density (number of species per unit area) may affect the establishment of fire ants and that the response of native ants and fire ants to disturbance can be comparable.     

Effects of Native and Non‐Native Vertebrate Mutualists on Plants

Abstract: Extinctions can leave species without mutualist partners and thus potentially reduce their fitness. In cases where non‐native species function as mutualists, mutualism disruption associated with species’ extinction may be mitigated. To assess the effectiveness of mutualist species with different origins, we conducted a meta‐analysis in which we compared the effectiveness of pollination and seed‐dispersal functions of native and non‐native vertebrates. We used data from 40 studies in which a total of 34 non‐native vertebrate mutualists in 20 geographic locations were examined. For each plant species, opportunistic non‐native vertebrate pollinators were generally less effective mutualists than native pollinators. When native mutualists had been extirpated, however, plant seed set and seedling performance appeared elevated in the presence of non‐native mutualists, although non‐native mutualists had a negative overall effect on seed germination. These results suggest native mutualists may not be easily replaced. In some systems researchers propose taxon substitution or the deliberate introduction of non‐native vertebrate mutualists to reestablish mutualist functions such as pollination and seed dispersal and to rescue native species from extinction. Our results also suggest that in places where all native mutualists are extinct, careful taxon substitution may benefit native plants at some life stages.