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Establishing a baseline of current Arctic vascular plant diversity and distribution is critical, given the rapid and major environmental changes occurring in the Arctic ecozone in response to climate change. Here, we report the results of a floristic study of vascular plant diversity of Dorset and Mallik islands in the Canadian Arctic Archipelago, Nunavut, Canada. These two small islands lie off the coast of the Foxe Peninsula of southwestern Baffin Island, and they are part of the Circumpolar Arctic bioclimate Subzone C. The hamlet of Kinngait (formerly Cape Dorset) is located on Dorset Island, and Nunavut’s Mallikjuak Territorial Park encompasses all of Mallik Island. The study is based on a specimen-based dataset comprising 876 unique collections from the two islands gathered over the last century, including 268 new ones collected in 2015. Results are presented in an annotated checklist. The vascular plant flora of the study area comprises 26 families, 71 genera, 150 species and three infraspecific taxa; 139 species are recorded on Dorset Island and 102 on Mallik Island. Eleven taxa are newly recorded from the study area in six families: Carex rupestris, Eriophorum scheuchzeri subsp. scheuchzeri, E. triste (Cyperaceae); Diapensia lapponica (Diapensiaceae); Equisetum arvense subsp. alpestre (Equisetaceae); Oxytropis deflexa var. foliolosa (Fabaceae); Potentilla arenosa subsp. arenosa, P. hyparctica subsp. hyparctica (Rosaceae); Antennaria friesiana subsp. friesiana, Askellia pygmaea, and Taraxacum phymatocarpum (Asteraceae).

Premise of the study Autopolyploidy, or whole-genome duplication within a species, leads to closely related cytotypes in one geographic location. One hypothesized mechanism by which autopolyploids become established is climatic niche divergence from their diploid progenitor. Here we tested this hypothesis in diploid, triploid, and tetraploid Galax urceolata (Diapensiaceae) and predicted the effects of climate change on the relative distributions of these cytotypes. Methods We investigated whether climatic niche divergence has shaped the current distributions of Galax urceolata cytotypes in eastern North America using climatic niche modeling, multivariate analyses of environmental space, and geographic range analyses. We then projected the models of the three cytotypes onto an ensemble of future climate maps to determine how the distributions might be altered over time. Key results All cytotypes are geographically sympatric; however, climatic niche contraction and a slight niche shift of the tetraploids was observed relative to that of the diploids. Climate projections for all diploid and tetraploid cytotypes showed substantial range contraction without much, or any, range shift, suggesting that Galax urceolata will likely go extinct in nature as mountain refugia become warmer. Conclusions Galax urceolata tetraploids occupy a slightly wetter habitat than that of their diploid progenitors. While we cannot take into account future adaptation, our models suggest extensive decreases in range distributions for both diploid and tetraploid G. urceolata based on climate change projections. Galax urceolata may therefore be under extreme threat due to loss of suitable habitat, and conservation efforts will be needed.

Callmander, M.W., O.D. Durbin, H.W. Lack, P. Bungener, P. Martin & L. Gautier (2017). Etienne-Pierre Ventenat (1757–1808) and the gardens of Cels and Empress Josephine. Candollea 72 : 87–132. In English, English and French abstracts. DOI : http//dx.doi.org/10.15553/c2017v721a8The intensive geographical exploration of the world, starting in the mid eighteenth century, resulted in the discovery of numerous plant species new to science of which many were subsequently introduced into cultivation in Europe. Etienne-Pierre Ventenat (1757–1808) contributed widely to this process. After the end of his curatorial and teaching commitments at Sainte-Genevieve Abbey in Paris he became an active botanist following in the footprints of his mentor Charles Louis L'Heritier de Brutelle (1746–1800). Ventenat was attracted by a trend of his period : publishing magnificent botanical books splendidly illustrated. He published three flower books from 1800 to 1808 on exotic plants recently brought to France from all over the world by various expeditions. Two flower books describe plants cultivated by Jacques-Martin Cels (1740–1806), an influential horticulturist who had built one of the most impressive plant collections of his period at Montrouge, just outside Paris. Another plant lover was Empress Josephine (1763–1814). After her wedding with Napoleon Bonaparte (1769–1821), she bought in 1799 the splendid estate of Malmaison west of Paris. There the Empress assembled a remarkable collection of exotic plants, largely cultivated in her orangery and the “Grande Serre Chaude”. She engaged Ventenat to describe her luxurious collection and the artist Pierre-Joseph Redoute (1761–1841), later known as the “Raphael des fleurs”, to depict the plants. This resulted in the famous flower book Jardin de la Malmaison, a masterpiece of botanical illustration. Ventenat, exhausted, passed away in 1808. His personal herbarium was bought in 1809 by Benjamin Delessert (1773–1847) whose herbarium was later donated to the City of Geneva in 1869. In his three flower books Ventenat treated a total of 343 plant names. A search in the herbaria of Geneva, Paris, Berlin and Madrid resulted in the finding of nearly all the original material relative to these names. A comprehensive list of names is provided here with comments on the origin of the plant material in the context of the explorations of this period, as well as information on typification and currently accepted names. In his three flower books Ventenat validated the names of 208 taxa (207 species and one variety) new to science : of these 67 are still accepted today as published while 52 are basionyms of names currently placed in another genera. Of these 208 names, lectotypification was not needed for 116 and the holotype has been found in G. For the remaining 92 names, 21 previous lectotypifications have been published but 5 of them needed a second-step lectotypification ; 70 new lectotypes have been selected and one neotype. Further comments on the typification of ten species are also provided.

Premise of the study Whereas population genetic studies have examined allopolyploids, comparable studies of naturally occurring autopolyploids remain rare. To address fundamental questions regarding autopolyploidy, we undertook a detailed population genetic study of one of the classic examples of autopolyploidy, Galax urceolata (Diapensiaceae), which comprises diploid, triploid, and autotetraploid cytotypes. Galax is endemic to the Appalachian Mountains, the adjacent piedmont, sandhills, and coastal plain and represents perhaps the most widely known example of autopolyploidy in nature. Methods Flow cytometry was used to diagnose ploidal level of ∼1000 individuals across 71 populations. We used 10 microsatellite markers to examine genetic variation across the geographic range of Galax and assessed multiple origins though comparisons of diploid, triploid, and tetraploid accessions using multiple analytical approaches. Key results Tetraploids had higher levels of heterozygosity than diploids did. Genetic variation in diploid and tetraploid Galax is geographically structured among the ecoregions of the southeastern United States. Autotetraploidy in Galax urceolata has occurred independently at least 46 times, with triploidy having occurred a minimum of 31 times. Conclusions Genetic differentiation among ecoregions suggests historical patterns of local adaptation. The numerous independent origins of tetraploid Galax reported here are among the highest frequencies of independent polyploidizations ever reported for any polyploid (auto- or allopolyploid).

Relationships among the genera of the small, woody family Styracaceae and among families of the large, diverse order Ericales have resisted complete resolution with sequences from one or a few genes. We used plastome sequencing to attempt to resolve the backbone relationships of Styracaceae and Ericales and to explore plastome structural evolution. Complete plastomes for 23 species are newly reported here, including 18 taxa of Styracaceae and five of Ericales (including species of Sapotaceae, Clethraceae, Symplocaceae, and Diapensiaceae). Combined with publicly available complete plastome data, this resulted in a data set of 60 plastomes, including 11 of the 12 genera of Styracaceae and 12 of 22 families of Ericales. Styracaceae plastomes were found to possess the quadripartite structure typical of angiosperms, with sizes ranging from 155 to 159 kb. Most of the plastomes were found to possess the full complement of typical angiosperm plastome genes. Unusual structural features were detected in plastomes of Alniphyllum and Bruinsmia, including the presence of a large 20-kb inversion (14 genes) in the Large Single-Copy region, the loss or pseudogenization of the clpP and accD genes in Bruinsmia, and the loss of the first exon of rps16 in B. styracoides. Likewise, the second intron from clpP was found to be lost in Alniphyllum and Huodendron. Phylogenomic analyses including all 79 plastid protein-coding genes provided improved resolution for relationships among the genera of Styracaceae and families of Ericales. Styracaceae was strongly supported as monophyletic, with Styrax, Huodendron, and a clade of Alniphyllum + Bruinsmia successively sister to the remainder of the family, all with strong support. All genera of Styracaceae were recovered as monophyletic, except for Halesia and Pterostyrax, which were each recovered as polyphyletic with strong support. Within Ericales, all families were recovered as monophyletic with strong support, with Balsaminaceae sister to remaining Ericales. Most relationships recovered in plastome analyses are congruent with previous analyses based on smaller data sets. Our results demonstrate the power of plastid phylogenomics to improve phylogenetic hypotheses among genera and families, and provide new insight into plastome evolution across Ericales.

74. Solenandria cordifolia P. Beauv. ex Vent., Jard. Malmaison 2: tab. 69. 1804. Lectotypus (designated here): UNITED STATES : ��� Caroline ���, Palisot de Beauvois s.n. (G [G00341549]!; isolecto-: G [G00341547, G00341548]!). = Galax urceolata (Poir.) Brummitt Notes. ��� Three specimens of a single gathering collected by Ambroise Marie Fran��ois Joseph Palisot de Beauvois (1752- 1820) have been located in Ventenat���s herbarium at G. The more complete collection is here designated as the lectotype., Published as part of Callmander, Martin W., Durbin, Olivier D., Lack, Hans-Walter, Bungener, Patrick, Martin, Pascal & Gautier, Laurent, 2017, Etienne-Pierre Ventenat (1757 - 1808) and the gardens of Cels and Empress Jos��phine, pp. 87-132 in Candollea 72 (1) on page 116, DOI: 10.15553/c2017v721a8, http://zenodo.org/record/5721919

Molecular phylogenetics is commonly used to perform accurate taxonomic classification. Furthermore, nuclear multilocus sequences have been applied to resolve ambiguous phylogenetic relationships based on conventional markers such as chloroplast and nuclear ribosomal DNA. Here, we reconstructed the phylogenetic tree of Diapensiaceae to revise the traditional classification of the genus Shortia sensu lato (s.l.), which includes Shortia sensu stricto (s.s.) and Schizocodon; whether the latter taxon is treated as an independent genus or includes Shortia s.l. has been disputed since its first description. The phylogeny based on eight nuclear markers developed for this study indicated that Shortia s.l. is not a monophyletic group and that Schizocodon and Shortia s.s. are reciprocally monophyletic. In addition, Schizocodon would be sister to other genus Diapensia despite their morphological differentiation. Thus, our data suggest that Schizocodon represents an independent genus. Our study demonstrated that multiple nuclear loci can resolve obscure phylogenetic relationships determined using conventional markers.

Range shifts during the Pleistocene shaped the unique phylogeographical structures of numerous species. Accompanying species migration, sister taxa may have experienced multiple introgression events. Here, we report the signature of introgression events in multiple areas in Schizocodon, herbs endemic to Japan, using amplified fragment length polymorphism (AFLP) fingerprinting and plastid DNA haplotyping in 48 populations. Although the present distributions of S. soldanelloides and S. ilicifolius are mainly allopatric, the species share plastid DNA haplotypes in each region (north-eastern, north-central, south-central and south-western Japan); in contrast, the specific groups were highly supported by AFLP analyses. These results support the occurrence of multiple introgression events in Schizocodon. Notably, the disjunct plastid haplotypes found only in S. ilicifolius var. intercedens suggest complete plastid DNA replacement at local areas from S. soldanelloides into S. ilicifolius var. ilicifolius. Furthermore, we found that S. soldanelloides experienced range contraction and expansion during glacial and interglacial cycles based on mismatch distribution analysis and ecological niche modelling. Based on several pieces of evidence, our study supports the idea that historical range shifts associated with Pleistocene climatic oscillations favoured multiple and regional introgression events in Schizocodon. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 46–63.

We compared allozyme variation in the two arctic–alpine plants Diapensia lapponica var. obovata and Empetrum nigrum var. japonicum between Sakhalin Island in Russian Far East, within their range core, and the Korean island of Jeju, their world’s southernmost distribution. For D. lapponica var. obovata, Sakhalin populations harbored moderate levels of within-population genetic variation and low among-population divergence, whereas extremely low levels of within-population genetic diversity and high among-population differentiation were found in Jeju Island populations. In contrast, we found moderate levels of within-population variation and low among-population differentiation in E. nigrum var. japonicum in both northern populations (those of Sakhalin and an additional population from northern Japan) and Jeju Island populations. Under a similar scenario of immigration history of arctic–alpine plants on Jeju Island during the glacial periods of the Pleistocene and local persistence through glacial/interglacial cycles, the contrasting genetic structure between D. lapponica var. obovata and E. nigrum var. japonicum is mainly attributable to their different life-history, ecological, and demographic traits: (1) hermaphroditic versus monoecious, dioecious or polygamous, (2) seeds with no adaptations for long-distance dispersal versus berry-like drupes dispersed by animals and birds, and (3) a very small patch near the peak of Mt. Halla with a few hundred individuals versus a relatively continuous distribution around the peak of Mt. Halla with numerous individuals. From a conservation perspective, in situ and ex situ conservation measures should be strengthened for D. lapponica var. obovata on Jeju Island given their extreme rarity there.

Structure and fungal identities were examined in the mycorrhizal roots of Schizocodon soldanelloides var. magnus (Diapensiaceae) to determine the mycorrhizal category. Previous studies had suggested the mycorrhizae of Diapensiaceae could be categorized as ericoid, but the mycorrhizal fungi have never been identified. The diameter of the fine lateral roots, in which coiled hyphae were found in epidermal cells, was mostly less than 100 μm. Molecular analyses identified the fungal isolates to be Helotiales and Oidiodendron. From the structure and fungal identities, we confirmed that the mycorrhiza of S. soldanelloides is an ericoid mycorrhiza.

Galax urceolata (Diapensiaceae) is a common evergreen herb of southern Appalachian forests. During the fall and winter, leaves of plants in high light produce substantial amounts of anthocyanins. Oddly, rhizomes in these plants also accumulate anthocyanins. The purpose of this observational study was to identify seasonal trends in anthocyanin production in above- and below-ground tissues of Galax. We measured anthocyanins and chlorophyll in Galax using standard extraction and spectrophotometric procedures from plants in sun-exposed and shaded locations; one population at Mount Jefferson State Natural Area in fall 2007 and two at Grandfather Mountain in fall/winter 2008–09. Rhizome carbohydrates (soluble sugars and starch) were measured from the Grandfather populations using high performance liquid chromatography. We found significantly more anthocyanins in leaves and rhizomes of plants from sun-exposed locations compared to plants from shaded locations, but no differences in carbohydrate concentr...

Pyxidanthera brevifolia is an evergreen semi-woody cushion plant endemic to the Sandhills of North and South Carolina, with the majority of populations occurring on Fort Bragg Military Reservation in North Carolina. Currently the species is listed as Endangered in North Carolina and is designated as a Species at Risk (SAR) by the US Department of Defense. Previous studies have suggested that seeds may not be viable because they failed to germinate under controlled conditions. Our objectives in this study were to attempt germination of Pyxidanthera brevifolia seeds, determine the best temperature conditions for germination, and understand more about germination requirements to aid in future restoration efforts. Using seeds that had been stored at room temperature for six months, we performed a germination experiment at the NCSU Phytotron with six treatments, all combinations of three temperature regimes (low (18 °C day / 14 °C night), medium (22/18 °C), and high (26/22 °C)) and two light condition...

Continental island systems harbour relict biota and populations that might have migrated during glacial periods due to the formation of landbridges. Here we analysed the genetic structure of relict populations of the temperate plant Shortia rotundifolia on the subtropical island of Iriomotejima, Japan. This plant, which inhabits riparian environments, is designated “near threatened”. Only five extant populations have been found on the island. Our analyses of 10 nuclear microsatellite loci detected genetic diversity of H E = 0.488 and H O = 0.358 for all populations of S. rotundifolia on the island. A high inbreeding coefficient for all populations together (F IS = 0.316) and each population separately (F IS = 0.258–0.497) might be attributable to crossing among closely related descendants within a population, an idea that is supported by the relatedness coefficient. These results and an examination of the populations’ demographic histories suggest that the extant populations on Iriomotejima have not experienced a recent population bottleneck. The five extant populations were genetically differentiated (F ST = 0.283; P

Previous phylogeographic studies of arctic-alpine plants have partly elucidated that alpine areas at the southernmost edges of their distribution have been one of the most important areas for the survival of populations through Quaternary climatic oscillations. To reveal the genetic structure of arctic-alpine plants near the southern edge of their range in East Asia, 159 samples of Diapensia lapponica subsp. obovata (Diapensiaceae) were collected from 22 populations in Japan. Based on 1,134 bp of chloroplast DNA (cpDNA), we determined 10 haplotypes. Neighbor-joining (NJ) analysis, including D. lapponica from Sweden and Schizocodon soldanelloides, revealed that all haplotypes in Japan were monophyletic. Genetic differentiation between populations of central Honshu and northern Japan was strongly supported by the NJ tree (100%), analysis of molecular variance (AMOVA, 68%), and Monmonier’s algorithm (>95%). The most ancestral haplotype rarely occurred in Japan, which implied that the extant genetic structure was not caused by genetic divergence during colonization. Thus, these results suggest that the arctic-alpine plant D. lapponica subsp. obovata survived climatic oscillations in central Honshu. This finding supports the hypothesis that high mountains in temperate regions were important for survival of arctic-alpine plants during Quaternary climatic oscillations.

The distribution of aluminium (Al) accumulation in the Ericales is surveyed, based on semi-quantitative tests of 114 species and literature data. Al accumulation mainly characterises the families Diapensiaceae, Pentaphylacaceae, Symplocaceae, Ternstroemiaceae, and Theaceae. Al accumulation is consistently present or absent in most families examined, but the character appears to be more variable in a few taxa (e.g., Lecythidaceae, Myrsinaceae). Although the interfamilial relationships within the Ericales require further research, the ability to accumulate high levels of Al appears to show considerable taxonomic significance. While the majority of Al accumulating Ericales includes woody, tropical plants, the feature is remarkably present in several herbaceous Diapensiaceae, which have a distribution in cold to temperate areas. The association of different mycorrhizae types with plant roots is suggested to play a role in the exclusion of high Al levels from the shoot.

Polyploid plants are typically geographically separated from their diploid progenitors, but it is rarely known whether this segregation is maintained by habitat differentiation or by selection against triploid hybrids. We tested for habitat differentiation between diploid and tetraploid populations of Galax urceolata in the southeastern United States by comparing the abundance and richness of vegetation associated with their respective habitats and testing for spatial autocorrelation in the distributions of diploid and tetraploid plants. Diploid and tetraploid habitats did not differ with respect to plant species richness. Herbaceous plants were 500% more common in tetraploid than diploid populations, while shrubs and trees were more common in diploid populations. Differences were found for the relative abundances of the most common species, with three species being significantly more prevalent in tetraploid than diploid populations. Spatial patchiness was detected for diploids and tetraploids, on a scale...

Phylogenetic interrelationships in the enlarged order Ericales were investigated by jackknife analysis of a combination of DNA sequences from the plastid genes rbcL, ndhF, atpB, and the mitochondrial genes atp1 and matR. Several well-supported groups were identified, but neither a combination of all gene sequences nor any one alone fully resolved the relationships between all major clades in Ericales. All investigated families except Theaceae were found to be monophyletic. Four families, Marcgraviaceae, Balsaminaceae, Pellicieraceae, and Tetrameristaceae form a monophyletic group that is the sister of the remaining families. On the next higher level, Fouquieriaceae and Polemoniaceae form a clade that is sister to the majority of families that form a group with eight supported clades between which the interrelationships are unresolved: Theaceae-Ternstroemioideae with Ficalhoa, Sladenia, and Pentaphylacaceae; Theaceae-Theoideae; Ebenaceae and Lissocarpaceae; Symplocaceae; Maesaceae, Theophrastaceae, Primulaceae, and Myrsinaceae; Styracaceae and Diapensiaceae; Lecythidaceae and Sapotaceae; Actinidiaceae, Roridulaceae, Sarraceniaceae, Clethraceae, Cyrillaceae, and Ericaceae.

Determinants of reproductive success (measured as capsule mass) in Diapensia lapponica, an early-flowering pollen-risker dependent on insects for pollination, were evaluated in an exposed subarctic-alpine population in northern Sweden. The species was protogynous and self-compatible, but self-deposition of pollen was inefficient. Hand-pollination increased capsule mass significantly, indicating that reproductive success was severely pollen limited. An experiment designed to test the relative importance of insect pollination in comparison to water availability (precipitation) and shelter (temperature and wind), revealed that exclosure of insects decreased capsule mass significantly, as did watering, whereas shelter tended to increase capsule mass. Pollination was the most important determinant of reproductive success, and the study suggests that pollen limitation may be particularly severe for early-flowering species, i.e. pollen-riskers, in exposed locations.

The geographical distributions of diploid and polyploid Galax urceolata overlap in the Blue Ridge Mountains, USA. As part of an investigation into the evolutionary forces governing the establishment of polyploids and their coexistence with diploids, we examined the population frequencies of diploids, triploids and tetraploids in the area of overlap. Ploidy was inferred from estimates of DNA content, using flow cytometry, for 1570 individuals sampled from 42 populations. Across the entire sampling area, diploids and tetraploids were most abundant (55% and 34% of individuals, respectively), whereas triploids were least abundant (11%). Cytotype frequencies differed significantly among the northern, central and southern regions of the range (G = 649.02, d.f. = 4, P < 0.0001), with diploids most frequent in the north-east and least frequent in the south-west. Twenty-six per cent of the populations contained three cytotypes, 33% contained two, and 40% had a single cytotype. Populations with two cytotypes occurred in all possible cytotype combinations, but when triploids were present, they were always in the minority. Uniform populations were either diploid (81%) or tetraploid (19%), but never triploid. Overall, populations are predominantly diploid or tetraploid but rarely evenly mixed, suggesting disruptive selection for chromosome number in G. urceolata. The contribution of ecological sorting and frequency-dependent mating success to the distribution of polyploids and diploids is discussed.

Cladistic analyses of chloroplast DNA disagree with current classifications by placingPolemoniaceae near sympetalous families with two staminal whorls, includingFouquieriaceae andDiapensiaceae, rather than near sympetalous families with a single staminal whorl, such asHydrophyllaceae andConvolvulaceae. To explore further the affinities ofPolemoniaceae, we sequenced 18S ribosomal DNA for eight genera ofPolemoniaceae and 31 families representing a broadly definedAsteridae. The distribution of variation in these sequences suggest some sites are hypervariable and multiple hits at these sites have obscured much of the hierarchical structure present in the data. Nevertheless, parsimony, least-squares minimum evolution, and maximum likelihood methods all support a monophyleticPolemoniaceae that is placed nearFouquieriaceae, Diapensiaceae and related “ericalean” families.

Ten microsatellite loci were isolated and characterized for Shortia rotundifolia, a relict temperate plant on the subtropical Ryukyu Islands and Taiwan. The number of alleles ranged from 2 to 21. The expected (H E) and observed (H O) heterozygosities were 0.027–0.877 and 0.027–0.622, respectively, from 74 individuals on Iriomotejima Island of Ryukyus. One locus exhibited significantly fewer heterozygotes than expected under Hardy–Weinberg equilibrium (P

Although anthocyanin coloration in lower (abaxial) leaf cells has been documented for numerous species, the functional significance of this character has not been comprehensively investigated according to habitat or leaf orientation. Here, we demonstrate that abaxial anthocyanin may function as a photoprotectant, similarly to its purported role in upper (adaxial) cells, in leaves vulnerable to high irradiance incident on abaxial surfaces. Spectral scans were derived for Galax urceolata leaves with the following phenotypes: abaxial or adaxial anthocyanin only, abaxial and adaxial anthocyanin, and no anthocyanin. To determine whether anthocyanins conferred protection from photoinhibition, maximum photosystem II efficiencies of red (anthocyanic) and green (acyanic) surfaces were compared during and after exposure to photoinhibitory conditions. Leaves were either positioned with their adaxial surfaces facing the light source or inverted to expose abaxial surfaces. Spectral scans showed increased absorptance of 500–600 nm wavelengths by red surfaces (consistent with the absorbance spectrum of anthocyanin), regardless of whether that surface was abaxial or adaxial. Leaves with anthocyanin in either illuminated surface were also photoinhibited less than leaves lacking anthocyanin in that surface. These results suggest that anthocyanic layers reduce absorbed sunlight in the mesophyll not only for adaxial surfaces, but also for the abaxial. Adaxial/abaxial anthocyanin plasticity may therefore be adaptive in high-light environments or during light-sensitive developmental stages where leaf orientation and/or substrate albedo are variable.

PREMISE OF THE STUDY Microsatellite markers were developed for Galax urceolata to investigate genetic diversity, population structure, and polyploid origins (auto- vs. allopolyploid), and to estimate the minimum number of independent cytotype origins. METHODS AND RESULTS Ten primer sets have been developed, and preliminary study indicates that all loci are appropriate for population-level genetic investigations. All loci are polymorphic with 6 to 46 alleles per locus. Expected heterozygosity ranges from 0.1007 to 0.6085. CONCLUSIONS The microsatellite markers presented will facilitate analyses of polyploid origins, genetic diversity, geographic structure, and gene flow.