Christopher G. Eckert’s research while affiliated with Queen's University and other places

Adaptation to new habitats might facilitate species’ range shifts in response to climate change. In 2005, we transplanted experimental populations of coastal dune plant Camissoniopsis cheiranthifolia into four sites within and one site beyond its poleward range limit. Beyond-range transplants had high fitness and often delayed reproduction. To test for adaptation associated with experimental range expansion, we transplanted descendants from beyond and within-range populations after 10 generations in situ into two sites within the range, one at the range edge, and two sites beyond the range. We expected to detect adaptation to beyond-range conditions due to substantial genetic variation within experimental populations and environmental variation among sites. However, individuals from beyond-range experimental populations were not fitter than those from within the range when planted at either beyond-range site, indicating no adaptation to the beyond-range site or beyond-range environments in general. Beyond-range descendants also did not suffer lower fitness within the range. Although reproduction was again delayed beyond the range, late reproduction was not favored more strongly beyond than within the range, and beyond-range descendants did not delay reproduction more than within-range descendants. Persistence in beyond-range environments may not require adaptation, which could allow a rapid response to climate change.

For many plants, asexual, clonal reproduction appears to be the predominant form of reproduction at their geographic range edge, but how long clonal populations persist and their role in range dynamics are largely unknown. If asexuality enables well-adapted genotypes to persist, this may allow species to expand beyond their sexual niche. Under this scenario, genomic signatures of long-term asexuality should be detectable in range-edge populations. We investigated this hypothesis in the wetland plant, Decodon verticillatus (Lythraceae), which reproduces predominantly through clonal reproduction at its northern range limit. We assembled the transcriptome de novo, identified single nucleotide polymorphisms (SNPs), and compared patterns of genetic variation in sexual and asexual populations at and approaching the range limit. Asexual genotypes exhibited several signatures of long-term asexuality including higher heterozygosity, fewer unique homozygous SNPs (doubletons), and a breakdown of isolation by distance. They also tended to include more deleterious non-synonymous and radical amino acid altering mutations. However, the frequency of unique heterozygous SNPs (singletons) did not differ between sexual and asexual genotypes, and average genetic differentiation was unexpectedly higher among sexual than among asexual genotypes. Yet, overall, our results are consistent with the hypothesis that asexual reproduction enabled D. verticillatus to expand its range further north than would have been possible under sexual reproduction alone. Understanding the factors that influence range dynamics is becoming increasingly important to better anticipate the capacity of species to adapt and shift their ranges in response to anthropogenic environmental changes and to prioritize range-edge populations for conservation.

Understanding the causes and limits of population divergence in phenotypic traits is a fundamental aim of evolutionary biology, with the potential to yield predictions of adaptation to environmental change. Reciprocal transplant experiments and the evaluation of optimality models suggest that local adaptation is common but not universal, and some studies suggest that trait divergence is highly constrained by genetic variances and covariances of complex phenotypes. We analyze a large database of population divergence in plants and evaluate whether evolutionary divergence scales positively with standing genetic variation within populations (evolvability), as expected if genetic constraints are evolutionarily important. We further evaluate differences in divergence and evolvability-divergence relationships between reproductive and vegetative traits and between selfing, mixed-mating, and outcrossing species, as these factors are expected to influence both patterns of selection and evolutionary potentials. Evolutionary divergence scaled positively with evolvability. Furthermore, trait divergence was greater for vegetative traits than for floral (reproductive) traits, but largely independent of the mating system. Jointly, these factors explained ~40% of the variance in evolutionary divergence. The consistency of the evolvability-divergence relationships across diverse species suggests substantial predictability of trait divergence. The results are also consistent with genetic constraints playing a role in evolutionary divergence.

Evolution of self-fertilization may be initiated by a historical population bottleneck, which should diagnostically reduce lineage-wide genetic variation. However, selfing can also strongly reduce genetic variation after it evolves. Distinguishing process from pattern is less problematic if mating system divergence is recent and geographically simple. Dramatically reduced diversity is associated with the transition from outcrossing to selfing in the Pacific coastal endemic Abronia umbellata that includes large-flowered, self-incompatible populations (var. umbellata) south of San Francisco Bay and small-flowered, autogamous populations (var. breviflora) to the north. Compared to umbellata, synonymous nucleotide diversity across 10 single-copy nuclear genes was reduced by 94% within individual populations and 90% across the whole selfing breviflora lineage, which contained no unique polymorphisms. The geographic pattern of genetic variation is consistent with a single origin of selfing that occurred recently (7-28 kya). These results are best explained by a historical bottleneck, but the two most northerly umbellata populations also contained little variation and clustered with selfing populations, suggesting that substantial diversity loss preceded the origin of selfing. A bottleneck may have set the stage for the eventual evolution of selfing by purging genetic load that prevents the spread of selfing.

Whether geographically peripheral populations are worth conserving has been hotly debated yet remains unresolved. This is especially relevant in high-latitude countries where, within their political jurisdictions, many species reach their range limits as peripheral isolates and require conservation attention, even if they are common elsewhere. In Canada, ~ 77% of “at-risk” plant species are at their northern range limit in southern Canada but more common south of the Canada-USA border. Peripheral populations might contain little genetic variation, suffer low fitness and be prone to extinction, or they might be adapted to extreme range-edge environments and thus well-poised to participate in range shifts during climate change. Abronia umbellata is endemic to coastal dunes from Baja California, Mexico to Oregon, USA but also occurs as disjunct populations designated “at-risk” in Washington, USA and British Columbia, Canada. Based on sequence variation at nine single-copy nuclear genes assayed for 94 individuals from 25 populations across the species range, these disjunct populations were very similar to range edge populations 350 and 650 km to the south in Oregon, and likely arose through recent, long-distance dispersal or fragmentation of a recently expanded range. In contrast, southern-edge populations in Baja, though not disjunct, were genetically unique and unexpectedly diverse, may currently be in decline yet receive no conservation protection. In this case, the conservation significance of range edge populations depends on which edge, and the unprotected southern edge populations seem a higher priority than those benefitting from special status at the northern range limit.

Ecological experiments usually infer long‐term processes from short‐term data, and the analysis of geographic range limits is a good example. Species' geographic ranges may be limited by low fitness due to niche constraints, a hypothesis most directly tested by comparing the fitness of populations transplanted within and beyond the range. Such studies often fail to find beyond‐range fitness declines strong enough to conclude that geographic range limits are solely imposed by niche limits. However, almost all studies only follow transplants for a single generation, which will underestimate the importance of niche limitation because critical but infrequent range‐limiting events may be missed and methodological issues may artificially boost the fitness of beyond‐range transplants. Here, we present the first multi‐generation beyond‐range transplant experiment that involves adequate replication and proper experimental controls. In 2005, experimental populations of the coastal dune plant Camissoniopsis cheiranthifolia were planted at four sites within and one site beyond the northern limit. Fitness of initial transplants was high beyond the limit, suggesting that the range was limited by dispersal and not niche constraints. To better address the niche‐limitation hypothesis, we quantified density and fitness of descendant C. cheiranthifolia populations 12–14 yr (˜10 generations) after transplant. Average annual fruit production and density of reproductive individuals were as high beyond the range as at four comparable experimental populations and eight natural populations within the range, and the beyond‐range population had more than tripled in size since it was planted. This provides unprecedented support for the conclusion that northern range limit of C. cheiranthifolia results from something other than niche limitation, likely involving constraints on local dispersal.

Colonisation along ubiquitous gradients of growing season length should require adaptation of phenological traits, driven by natural selection. While phenology often varies with season length and genetic differentiation in phenological traits sometimes seems adaptive, few studies test whether natural selection is responsible for these patterns. The annual plant Rhinanthus minor is genetically differentiated for phenology across a 1000-m elevational gradient of growing season length in the Canadian Rocky Mountains. We estimated phenotypic selection on five phenological traits for three generations of naturally occurring individuals at 12 sites (n = 10112), and two generations of genetically and phenotypically more variable transplanted populations at nine of these sites (n = 24611). Selection was weak for most traits, but consistently favored early flowering across the gradient rather than only under short seasons. There was no evidence that apparent selection favoring early reproduction arose from failure to consider all components of fitness, or variation in other correlated phenological traits. Instead, selection for earlier flowering may be balanced by selection for strong cogradient phenological plasticity which indirectly favors later flowering. However, this probably does not explain the consistency of selection on flowering time across this steep elevational gradient of growing season length. This article is protected by copyright. All rights reserved

High-latitude countries tend to contain the polar range-edge of many species that are nationally rare but globally common. This can focus national conservation efforts toward range-edge populations, whose conservation needs and value are disputed. Using plants in Canada as a case study, we ask whether national species-conservation rankings prioritize range-edge populations, and whether conservation priority is matched by habitat protection and research effort. We found that >75% of federally protected plants only occur in Canada peripherally, at the northernmost 20% or less of their total range, and that the most imperilled taxa had the smallest percentage of their range in Canada (endangered plants: median = 1.0%). Occurring peripherally in Canada was associated with higher threat even after accounting for range area, potentially because range-edge taxa experienced 85% higher human population densities in their Canadian range than non-peripheral taxa. High conservation priority was not matched by habitat protection, as more imperilled and more peripheral taxa had smaller fractions of their Canadian range in protected areas. Finally, peer-reviewed research on plants at-risk in Canada was low. Only 42% of plants considered at-risk in Canada had been studied in Canada, and only 11% of species with broad distributions outside Canada had been studied in the context of their wider geographic range—information that is critical to establishing their relative conservation value. Our results illustrate that plant conservation in Canada is fundamentally linked to conserving range-edge populations, yet edge populations themselves are understudied, a research gap we must close to improve evidence-based conservation.

High-latitude countries tend to contain the polar range-edge of many species that are nationally rare but globally common. This can focus national conservation efforts toward range-edge populations, whose conservation needs and value are disputed. Using plants in Canada as a case study, we ask whether national species-conservation rankings prioritize range-edge populations, and whether conservation priority is matched by habitat protection and research effort. We found that >75% of federally-protected plants only occur in Canada peripherally, at the northernmost 20% or less of their total range, and that the most imperilled taxa had the smallest percentage of their range in Canada (endangered plants: median=1.0%). Occurring peripherally in Canada was associated with higher threat even after accounting for range area, potentially because range-edge taxa experienced 85% higher human population densities in their Canadian range than non-peripheral taxa. High conservation priority was not matched by habitat protection, as more imperilled and more peripheral taxa had smaller fractions of their Canadian range in protected areas. Finally, peer-reviewed research on plants at-risk in Canada was low. Only 42% of plants considered at-risk in Canada had been studied in Canada, and only 11% of species with large distributions outside Canada had been studied in the context of their wider geographic range information that is critical to establishing their relative conservation value. Our results illustrate that plant conservation in Canada is fundamentally linked to conserving range-edge populations, yet edge populations themselves are understudied, a research gap we must close to improve evidence-based conservation.

Premise: Interspecific hybridization can cause genetic structure across species ranges if the mating system and degree of sympatry/parapatry with close relatives varies geographically. The coastal dune endemic Camissoniopsis cheiranthifolia (Onagraceae) exhibits genetic subdivisions across its range, some of which are associated with shifts in mating system from outcrossing to selfing, while others are not. For instance, strong differentiation between large-flowered, self-incompatible (LF-SI) and large-flowered, self-compatible (LF-SC) populations occurs without much reduction in outcrossing or obvious barriers to gene flow. We hypothesized that LF-SI diverged from LF-SC via hybridization with the predominantly inland SI sister species C. bistorta. Methods: We analyzed spatial proximity using 1460 herbarium records and genetic variation at 12 microsatellites assayed for 805 and 404 individuals from 32 C. cheiranthifolia and 18 C. bistorta populations, respectively. We also assayed nine chloroplast microsatellites for 124 and 111 individuals from 27 and 19 populations, respectively. Results: Closer parapatry was associated with unexpectedly high genetic continuity between LF-SI C. cheiranthifolia and C. bistorta. LF-SI genotypes clustered with C. bistorta exclusive of other C. cheiranthifolia genotypes. Similarly, pairwise FST among SI C. cheiranthifolia and C. bistorta, adjusted for geographic proximity, was not higher between heterospecific than conspecific populations. Conclusions: The lack of genetic differentiation between LF-SI C. cheiranthifolia and C. bistorta populations, even those located away from the zone of parapatry, suggests that, instead of hybridizing with C. bistorta, LF-SI C. cheiranthifolia is rather an ecotype of C. bistorta that has adapted to coastal dune habitat independent of other lineages in C. cheiranthifolia proper.