Non-native Range Research Articles

Biogeographic differences in the effects of Centaurea stoebe on the soil nitrogen cycle: novel weapons and soil microbes

The success of some invasive plants may be due in part to native organisms lacking adaptation to species-specific biochemical traits of invaders—the Novel Weapons Hypothesis. We tested this hypothesis in the context of soil microbial communities by comparing the effects of Centaurea stoebe and the root exudate (±)-catechin, on ammonification and nitrification in both the non-native and native ranges of this species. In a non-native range (Montana), soil nitrate (NO3−) concentrations were lower in invaded than uninvaded grasslands. This did not appear to be due only to higher uptake rates as both C. stoebe plants and catechin significantly reduced resin extractable NO3−, the maximum rate of nitrification, and gross nitrification in Montana soils. Thus, reduced NO3− in invaded communities may be due in part to the inhibition of nitrifying bacteria by secondary metabolites produced by C. stoebe. The effects of C. stoebe on N-related processes were different in Romanian grasslands, where C. stoebe is native. In Romanian soil, C. stoebe had no effect on resin extractable NH4+ or NO3− (compared to other plant species), the maximum rate of nitrification, nor gross nitrification. A relatively high concentration of catechin reduced the maximum rate of nitrification in situ, but substantially less than in Montana. In vivo, gross ammonification was lowest when treated with catechin. Our results suggest biogeographic differences in the way a plant species alters nitrogen cycling through the direct effects of root exudates and adds to a growing body of literature demonstrating the important belowground effects of invasive plants.

Scent chemistry and pollinator attraction in the deceptive trap flowers of Ceropegia dolichophylla

Ceropegia species (Apocynaceae, Asclepiadoideae) have pitfall flowers and are pollinated by small flies through deception. It has been suggested that these flies are attracted by floral scent. However, the scent that is emitted from Ceropegia flowers has not been studied using headspace and gas chromatography mass spectrometry methods. It has also been unclear whether or not the flowers are mimics of particular models that attract flies. In the present study, we determined the composition as well as the spatial and temporal patterns of floral scent emitted by C. dolichophylla. Furthermore, we determined the pollinators in the native (China) and non-native (Germany) range of this species, and tested the capability of the floral scent to attract flies in the non-native range. Our data demonstrate that the floral scent, which is emitted from morning until evening, primarily from the tips of the corolla lobes, consists mainly of spiroacetals and aliphatic compounds. Milichiid flies were common visitors/pollinators in the native as well as non-native range, and were attracted by floral scent in bioassays performed in the non-native range. The compounds emitted by C. dolichophylla are unusual for flowers, but are well known from insect pheromones and occur in the glandular secretions of insects. The milichiid flies that visit and pollinate the flowers are kleptoparasites that feed on the prey (haemolymph or other secretions) of predatory arthropods, e.g. spiders, to which they are attracted by scent. Our data thus suggest that the floral scent of C. dolichophylla mimics the feeding sites of kleptoparasitic flies.

Acroptilon repens, an Asian invader, has stronger competitive effects on species from America than species from its native range

The ability to competitively suppress native species is key to successful invasion. Since invasions involve an increase in abundance or dominance of a species in its non-native range, competitive effects might be expected to be stronger in the non-native range of an invader; however, there have been few comparisons of the competitive effects of invasive plants on species from invaded ranges versus species from native ranges. We compared the competitive and allelopathic effects of Acroptilon repens on native North American species to effects on related species from the native range of Acroptilon in Uzbekistan. We also compared the competitive interactions among these North American and Eurasian species, in the absence of Acroptilon, examining the hypothesis that particular regional species pools may show differences in competitive ability. The results showed that Acroptilon had stronger competitive effects against native North American species than against species native to Uzbekistan. There was no difference in the competitive effects among Eurasians and North Americans. The effects of leachates collected from Acroptilon roots were weak but more negative on species from North America than on species from Uzbekistan. Our results suggest that inherently stronger competitive and allelopathic effects of Acroptilon on North American plants than on plants from its native range may contribute to its invasive success.

The current range of Amur sleeper Perccottus glenii Dybowski, 1877 (Odontobutidae, Pisces) in Eurasia

The native range of the fish Amur sleeper Perccottus glenii is in the Far East of the Russian Federation, in China and North Korea. From 1916 to 2008 Amur sleeper has spread widely through North Eurasia, causing local declines of many populations of indigenous invertebrates, fishes and amphibians. The extent of its non-native range has been determined by analysis of literature, museum collections, specialist responses to a questionnaire and data of four expeditions to West and East Siberia. These sources yielded 876 records of Amur sleeper. Several new non-native parts of the rang e were described, but some previously reported parts of the range were considered unfounded. Overall, the current distribution of P. glenii is wider than previously believed. Up to the present time, this fish has been found outside its native range in 48 regions (36 provinces, 9 republics, and 3 territories) of the Russian Federation as well as in Latvia, Lithuania, Estonia, Byelorussia, Ukraine, Moldova, Kazakhstan (north of the country), Mongolia, Poland, Slovakia, Hungary, Serbia, Bulgaria, and Romania. The first indications of the possible appearance of this species have been received from Moldova and Italy. The non-native distribution of Amur sleeper covers now more than 100° West to East and 20° South to North. The detailed map of the current invasive range is presented. The map of native range is corrected.

Assessing the biosecurity risk from pathogens and herbivores to indigenous plants: lessons from weed biological control

Some potentially invasive herbivores/pathogens in their home range may attack plants originating from another geographic area. Methods are required to assess the risk these herbivores/pathogens pose to these plants in their indigenous ecosystems. The processes and criteria used by weed biological control researchers to assess the impact of potential biological control agents on a plant species in its non-native range provide a possible framework for assessing risks to indigenous plants. While there are similarities between these criteria such as the need for clear objectives, studies in the native range of the herbivore/pathogen, good knowledge of the ecology of the target plant and taxonomy of the plant and herbivore/pathogen, and modelling of the interaction between the two organisms, there are some important differences in approach. These include the need to consider the threat classification of the plant, the likely greater risk from polyphagous herbivores/pathogens than oligophagous or monophagous species, and the need to consider the impact of an additional natural enemy in conjunction with a suite of existing natural enemies. The costs of conducting a risk assessment of a herbivore/pathogen in another country that damages plants indigenous to another geographic area means that criteria will be needed for deciding which foreign herbivore/pathogen species should be assessed. These criteria could include the threat classification of the plant, the amount of damage to the particular plant organs affected, and the importance in key ecosystems.

Virulence of soil‐borne pathogens and invasion by Prunus serotina

*Globally, exotic invaders threaten biodiversity and ecosystem function. Studies often report that invading plants are less affected by enemies in their invaded vs home ranges, but few studies have investigated the underlying mechanisms. *Here, we investigated the variation in prevalence, species composition and virulence of soil-borne Pythium pathogens associated with the tree Prunus serotina in its native US and non-native European ranges by culturing, DNA sequencing and controlled pathogenicity trials. *Two controlled pathogenicity experiments showed that Pythium pathogens from the native range caused 38-462% more root rot and 80-583% more seedling mortality, and 19-45% less biomass production than Pythium from the non-native range. DNA sequencing indicated that the most virulent Pythium taxa were sampled only from the native range. The greater virulence of Pythium sampled from the native range therefore corresponded to shifts in species composition across ranges rather than variation within a common Pythium species. *Prunus serotina still encounters Pythium in its non-native range but encounters less virulent taxa. Elucidating patterns of enemy virulence in native and nonnative ranges adds to our understanding of how invasive plants escape disease. Moreover, this strategy may identify resident enemies in the non-native range that could be used to manage invasive plants.

A multi‐species experiment in their native range indicates pre‐adaptation of invasive alien plant species

*To understand prerequisites of biological invasions, it is imperative to know whether species have traits that pre-adapt them to become invasive elsewhere. However, few experimental studies have explicitly tested this by comparing traits between invasive and noninvasive species in their native range instead of in the nonnative range. *We used native plant material of 14 European congeneric pairs of herbaceous species that were all introduced to North America, and of which one species per pair is invasive. *In our germination and common garden experiment with and without fertilizer addition, the invasive species germinated faster, produced more biomass and had a higher proportion of flowering plants than the noninvasive congeners. *Our results indicate that species traits, which lead to a high plant performance in the native range, can confer pre-adaptation to become invasive. We suggest that such traits may be especially relevant for use in risk-assessment protocols before introduction elsewhere.

Bioclimatic predictions of habitat suitability for the biofuel switchgrass in North America under current and future climate scenarios

Abstract Dedicated biofuel crops, while providing economic and other benefits, may adversely impact biodiversity directly via land use conversion, or indirectly via creation of novel invasive species. To mitigate negative impacts bioclimatic envelope models (BEM) can be used to estimate the potential distribution and suitable habitat based on the climate and distribution in the native range. We used CLIMEX to evaluate the regions of North America suitable for agronomic production, as well as regions potentially susceptible to an invasion of switchgrass (Panicum virgatum) under both current and future climate scenarios. Model results show that >8.7 million km2 of North America has suitable to very favorable habitat, most of which occurs east of the Rocky Mountains. The non-native range of western North America is largely unsuitable to switchgrass as a crop or potential weed unless irrigation or permanent water is available. Under both the CGCM2 and HadCM3 climate models and A2 and B2 emissions scenarios, an overall increase in suitable habitat is predicted over the coming century, although the western US remains unsuitable. Our results suggest that much of North America is suitable for switchgrass cultivation, although this is likely to shift north in the coming century. Our results also agree with field collections of switchgrass outside its native range, which indicate that switchgrass is unlikely to establish unless it has access to water throughout the year (e.g., along a stream). Thus, it is the potential invasion of switchgrass into riparian habitats in the West that requires further investigation.

Rethinking the common garden in invasion research

In common garden experiments, a number of genotypes are raised in a common environment in order to quantify the genetic component of phenotypic variation. Common gardens are thus ideally suited for disentangling how genetic and environmental factors contribute to the success of invasive species in their new non-native range. Although common garden experiments are increasingly employed in the study of invasive species, there has been little discussion about how these experiments should be designed for greatest utility. We argue that this has delayed progress in developing a general theory of invasion biology. We suggest a minimum optimal design (MOD) for common garden studies that target the ecological and evolutionary processes leading to phenotypic differentiation between native and invasive ranges. This involves four elements: (A) multiple, strategically sited garden locations, involving at the very least four gardens (2 in the native range and 2 in the invaded range); (B) careful consideration of the genetic design of the experiment; (C) standardization of experimental protocols across all gardens; and (D) care to ensure the biosafety of the experiment. Our understanding of the evolutionary ecology of biological invasions will be greatly enhanced by common garden studies, if and only if they are designed in a more systematic fashion, incorporating at the very least the MOD suggested here.

Patterns of selection of two North American native and nonnative populations of monkeyflower (Phrymaceae)

To better understand invasion dynamics, it is essential to determine the influence of genetics and ecology in species persistence in both native and nonnative habitats. One approach is to assess patterns of selection on floral and growth traits of individuals in both habitats. Mimulus guttatus (Phrymaceae) has a mixed mating system and grows under variable water conditions across its native and nonnative range in North America. Field investigations of patterns of selection of floral and plant size traits were conducted in two native and two nonnative populations. Field-collected seed was grown and crossed in the glasshouse using a paternal half-sib design. The resulting offspring were grown in saturated and dry-down low-water conditions and the same traits were measured in both environments. Patterns of selection varied across years in the native range. Nonnative populations exhibited selection for increased floral size, consistent with the hypothesis that selection favors larger size in nonnative habitats. In the glasshouse, we detected genetic variation for traits across population/treatment combinations. However, size hierarchy in the glasshouse was dependent on water conditions. Our results suggest that both variable selection pressures and local adaptation probably influence the persistence of both native and nonnative populations.

Enhanced fitness due to higher fecundity, increased defence against a specialist and tolerance towards a generalist herbivore in an invasive annual plant

Aims: The superior performance of many non-indigenous species in a new range can be attributed to different factors such as pre-adaptation to environmental conditions in new areas or to factors inherent to dis- placement mechanisms such as loss of co-evolved pathogens and herbivores that increase the speed of evolutionary change towards a shift in allocation from defence to growth and reproduction. To as- sess the importance of the different mechanisms governing the suc- cess of Conyza canadensis, a globally successful invader, we simultaneously tested several recent hypotheses potentially explain- ing the factors leading to biological invasion. Methods: We tested (i) whether plants from the non-native range showed a higher fitness than plants from the native North American range, (ii) whether they differed in resistance against an invasive generalist herbivore, the slug Arion lusitanicus and against a recently estab- lished specialist aphid herbivore, Uroleucon erigeronense and (iii) experimentally assessed whether C. canadensis releases allelopathic chemicals that have harmful effects on competing species in the non- native range. We compared populations along a similar latitudinal gradient both in the native North American and invasive European range and analysed patterns of adaptive clinal variation in biomass production.

An explicit test for the contribution of environmental maternal effects to rapid clinal differentiation in an invasive plant

Population differentiation of alien invasive plants within their non-native range has received increasingly more attention. Common gardens are typically used to assess the levels of genotypic differentiation among populations.However, in such experiments, environmental maternal effects can influence phenotypic variation among individuals if seed sources are collected from field populations under variable environmental regimes. In the present study, we investigated the causes of an altitudinal cline in an invasive plant. Seeds were collected from Senecio inaequidens (Asteraceae) populations along an altitudinal gradient in southern France. In addition, seeds from the same populations were generated by intra-population crossings in a climatic chamber. The two seed lots were grown in a common garden in Central Belgium to identify any evidence of environmentally induced maternal effects and ⁄ or an altitudinal cline in a suite of life-history traits. Results failed to detect any environmental maternal effects. However, an altitudinal cline in plant height and aboveground biomass was found to be independent of the maternal environment.

Germination responses of an invasive species in native and non-native ranges

Studying germination in the native and non-native range of a species can provide unique insights into processes of range expansion and adaptation; however, traits related to germination have rarely been compared between native and nonnative populations. In a series of common garden experiments, we explored whether differences in the seasonality of precipitation, specifically, summer drought vs summer rain, and the amount and variation of annual and seasonal precipitation affect the germination responses of populations of an annual ruderal plant, Centaurea solstitialis, from its native range and from two non-native regions with different climates. We found that seeds from all native populations, irrespective of the precipitation seasonality of the region in which they occurred, and non-native populations from regions with dry summers displayed similarly high germination proportions and rates. In contrast, genotypes from the non-native region with predominantly summer rain exhibited much lower germination fractions and rates. Also, percent germination was strongly correlated with variation in precipitation in winter, the season that follows germination for C. solstitialis. Specifically, germination was lower for native and non-native populations experiencing greater variation in winter precipitation. This correlation, however, was greatly influenced by the non-native region with summer rain, which also exhibited the greatest variation in winter precipitation among studied regions. These results suggest that rather than general climatic patterns, the degree of risk experienced at early developmental stages could exert an important control over the germination strategy of C. solstitialis populations in both native and non-native ranges. Also, these findings reveal a largely unique germination response in C. solstitialis genotypes growing in the non-native region with summer rain and high variation in winter precipitation. Our work raises the possibility that rapid adaptive changes in germination strategies may contribute to the success of globally distributed invaders.

Ecophysiological characteristics of invasive Spartina alterniflora and native species in salt marshes of Yangtze River estuary, China

Biological invasions represent one of the significant components of global change. A comparative study of invaders and co-occurring natives is a useful approach to gaining insights into the invasiveness of exotic plants. Spartina alterniflora, a C 4 grass, is a widespread invader in the coastal wetlands in China and other regions of the world. We conducted a comparative study of S. alterniflora and native C 3 species, Phragmites australis and Scirpus mariqueter, in terms of their gas exchange and efficiencies in resource utilization. We tested the hypothesis that S. alterniflora has growth-related ecophysiological advantages over the natives in its non-native range, which result in its rapid growth and enhance its invasiveness. Photosynthesis, leaf area index (LAI), specific leaf area (SLA), and the efficiency of resource use (light, water, and nitrogen) were examined monthly for eight months in 2004. Overall, S. alterniflora had greater LAI, higher maximal net photosynthetic rate ( A max), and longer growing season than those of the native species. On average, the efficiencies of S. alterniflora in light, water, and nitrogen utilization were respectively 10.1%, 26.1%, and 33.1% higher than those of P. australis, and respectively 70.3%, 53.5%, 28.3% higher than those of S. mariqueter. However, SLA of S. alterniflora was significantly lower than those of P. australis and S. mariqueter. Although there was no general pattern in the relationship between invasiveness and plant photosynthetic types, in this study, most of the ecophysiological characteristics that gave S. alterniflora a competitive advantage in the Yangtze River estuary were associated with photosynthetic pathways. Our results offer a greater understanding of the relationship between invasiveness and plant photosynthetic type. Our results also indicate that LAI and the length of the photosynthetic season, which vary with habitats, are also important in invasion success.

The distribution and abundance of the Neogobius fishes in their native range (Bulgaria) with notes on the non-native range in the Danube River

The recent distribution and abundances of the genus Neogobius were studied (using electrofishing and beach seining) within (Bulgaria) and outside (Austria, Slovakia, Croatia) its native range along the longitudinal profile of the Danube River in 2004, 2005 and 2006. The sampling was conducted under normal autumn water levels. CPUE values (number of fish per 100 m shoreline) were calculated and compared between sampling sites. The presence of four Neogobius species - N. kessleri, N. fluviatilis, N. melanostomus, and N. gymnotrachelus -was confirmed in Bulgaria and Croatia. Though the latter two species did occur in Croatia, they occurred at very low densities (comparing to the densities in the upper and lower Danube), indicating newly established populations. Our results support the hypothesis of disjunct spreading of these species. The presence of N. fluviatilis in Austria was not confirmed. In general, the Neogobius spp. showed the lowest density within their native range (6 individuals per 100 m shoreline) while their densities were considerably larger outside their native range (18-28 individuals per 100 m shoreline). In addition. N. kessleri and N. melanostomus were significantly larger at sites of non-native distribution compared to native sites.

Multiple Sources, Admixture, and Genetic Variation in IntroducedAnolisLizard Populations

Invasive species are classically thought to suffer from reduced within-population genetic variation compared to their native-range sources due to founder effects and population bottlenecks during introduction. Reduction in genetic variation in introduced species may limit population growth, increase the risk of extinction, and constrain adaptation, hindering the successful establishment and spread of an alien species. Results of recent empirical studies, however, show higher than expected genetic variation, rapid evolution, and multiple native-range sources in introduced populations, which challenge the classical scenario of invasive-species genetics. With mitochondrial DNA (mtDNA) sequence data, we examined the molecular genetics of 10 replicate introductions of 8 species of Anolis lizards. Eighty percent of introductions to Florida and the Dominican Republic were from multiple native-range source populations. MtDNA haplotypes restricted to different geographically distinct populations in the native range of a species commonly occurred as intrapopulation polymorphisms in introduced populations. Two-thirds of introduced populations had two or more sources, and admixture elevated genetic variation in half of the introduced populations above levels typical of native-range populations. The mean pairwise sequence divergence among haplotypes sampled within introduced populations was nearly twice that within native-range populations (2.6% vs. 1.4%). The dynamics of introductions from multiple sources and admixture explained the observed genetic contrasts between native and introduced Anolis populations better than the classical scenario for most introduced populations. Elevated genetic variation through admixture occurred regardless of the mode or circumstances of an introduction. Little insight into the number of sources or amount of genetic variation in introduced populations was gained by knowing the number of physical introductions, the size of a species' non-native range, or whether it was a deliberate or accidental introduction. We hypothesize that elevated genetic variation through admixture of multiple sources is more common in biological invasions than previously thought. We propose that introductions follow a sequential, two-step process involving a reduction in genetic variation due to founder effects and population bottlenecks followed by an increase in genetic variation if admixture of individuals from multiple native-range sources occurs.

Microhabitat Use by Introduced Hemidactylus turcicus (Mediterranean Geckos) in North Central Florida

We examined the relationship between seven wall-microhabitat features and the occurrence of the nonindigenous Hemidactylus turcicus (Mediterranean Gecko) in north-central Florida using a repeatable technique. We characterized 160 one-story walls by age of the building, cardinal orientation, color, length, presence or absence of a light source, building material, and vegetation level, and recorded the presence or absence of H. turcicus for each wall during two separate nocturnal visits. The occurrence of H. turcicus was only dependent on wall surface color and length. Both the lack of significance of the majority of the microhabitat variables investigated and the fact that H. turcicus was found on all wall types suggest that this gecko is capable of inhabiting a wide variety of wall environments. This habitat flexibility may be a key factor in the prolific expansion of this gecko's nonnative range.

Forecasting the Spread of Invasive Rainbow Smelt in the Laurentian Great Lakes Region of North America

Rainbow smelt (Osmerus mordax) have invaded many North American lakes, often resulting in the extirpation of native fish populations. Yet, their invasion is incipient and provides the rationale for identifying ecosystems likely to be invaded and where management and prevention efforts should be focused. To predict smelt presence and absence, we constructed a classification-tree model based on habitat data from 354 lakes in the native range for smelt in southern Maine. Maximum lake depth, lake area, and Secchi depth (surrogate measure of lake productivity) were the most important predictors. We then used our model to identify lakes vulnerable to invasion in three regions outside the smelt's native range: northern Maine (52 of 244 lakes in the non-native range), Ontario (4447 of 8110), and Wisconsin (553 of 5164). We further identified a subset of lakes with a strong potential for impact (potential-impact lakes) based on the presence of fish species that are affected by rainbow smelt. Ninety-four percent of vulnerable lakes in the non-native range in Maine are also potential-impact lakes, as are 94% and 58% of Ontario and Wisconsin's vulnerable lakes, respectively. Our modeling approach can be applied to other invaders and regions to identify invasion-prone ecosystems, thus aiding in the management of invasive species and the efficient allocation of invasive species mitigation and prevention resources.

Production Ecology of the Non-indigenous Seagrass, Dwarf Eelgrass (Zostera japonica Ascher. & Graeb.), in a Pacific Northwest Estuary, USA

The non-indigenous seagrass Zostera japonica Ascher. & Graeb. (dwarf eelgrass) was first identified in central Oregon (USA) estuaries about 30 years ago. The autecology of this species is poorly described at the southern end of its non-native range although several process oriented studies have been conducted. I examined the production ecology of Z. japonica in the Yaquina Bay estuary. Strong seasonal patterns in light and temperature appeared to control the seasonal variations in biomass and growth. Above- and below-ground biomass ranged between 40–100 and 70–170 gdw m−2 respectively and seasonal changes in the root:shoot ratio were controlled by above-ground biomass dynamics. Shoot density ranged between 4000 and 11 000 shts m−2. Areal leaf growth ranged between 0.1 and 1.7 gdw m−2 d−1 and annual production was about 314 ± 60 gdw m−2 y−1 (mean ± SD). Nutrients were not limiting in this system as a result of coastal upwelling and watershed inputs. The Z. japonica population studied in Oregon exhibited different patterns of persistence, phenology and flowering intensity relative to other populations along its native and non-native range. These differences suggest that management policies developed for one site may not be appropriate for other sites. The data presented here greatly expands our knowledge base on Z. japonica and provides insight to the processes controlling the dynamics and spread of this non-indigenous seagrass.

EXPERIMENTAL STUDIES OF ADAPTATION IN clarkia xantiana:II. FITNESS VARIATION ACROSS A SUBSPECIES BORDER

Because the range boundary is the locale beyond which a taxon fails to persist, it provides a unique opportunity for studying the limits on adaptive evolution. Adaptive constraints on range expansion are perplexing in view of widespread ecotypic differentiation by habitat and region within a species' range (regional adaptation) and rapid evolutionary response to novel environments. In this study of two parapatric subspecies, Clarkia xantiana ssp. xantiana and C. x. ssp. parviflora, we compared the fitness of population transplants within their native region, in a non-native region within the native range, and in the non-native range to assess whether range expansion might be limited by a greater intensity of selection on colonists of a new range versus a new region within the range. The combined range of the two subspecies spans a west-to-east gradient of declining precipitation in the Sierra Nevada of California, with ssp. xantiana in the west being replaced by ssp. parviflora in the east. Both subspecies had significantly higher fitness in the native range (range adaptation), whereas regional adaptation was weak and was found only in the predominantly outcrossing ssp. xantiana but was absent in the inbreeding ssp. parvifilora. Because selection intensity on transplants was much stronger in the non-native range relative to non-native regions, there is a larger adaptive barrier to range versus regional expansion. Three of five sequential fitness components accounted for regional and range adaptation, but only one of them, survivorship from germination to flowering, contributed to both. Flower number contributed to regional adaptation in ssp. xantiana and fruit set (number of fruits per flower) to range adaptation. Differential survivorship of the two taxa or regional populations of ssp. xantiana in non-native environments was attributable, in part, to biotic interactions, including competition, herbivory, and pollination. For example, low fruit set in ssp. xantiana in the east was likely due to the absence of its principal specialist bee pollinators in ssp. parviflora's range. Thus, convergence on self-fertilization may be necessary for ssp. xantiana to invade ssp. parviflora's range, but the evolution of outcrossing would not be required for ssp. parviflora to invade ssp. xantiana's range.