Delineation Of Taxa Research Articles

Putative genome contamination has minimal impact on the GTDB taxonomy.

The Genome Taxonomy Database (GTDB) provides a species to domain classification of publicly available genomes based on average nucleotide identity (ANI) (for species) and a concatenated gene phylogeny normalized by evolutionary rates (for genus to phylum), which has been widely adopted by the scientific community. Here, we use the Genome UNClutterer (GUNC) software to identify putatively contaminated genomes in GTDB release 07-RS207. We found that GUNC reported 35,723 genomes as putatively contaminated, comprising 11.25 % of the 317,542 genomes in GTDB release 07-RS207. To assess the impact of this high level of inferred contamination on the delineation of taxa, we created 'clean' versions of the 34,846 putatively contaminated bacterial genomes by removing the most contaminated half. For each clean half, we re-calculated the ANI and concatenated gene phylogeny and found that only 77 (0.22 %) of the genomes were not consistent with their original classification. We conclude that the delineation of taxa in GTDB is robust to the putative contamination detected by GUNC.

Leaf heteroblasty and morphotypes of Acer monspessulanum (Djurdjura, Algeria) as revealed by traditional and geometric morphometrics

The genus Acer which is largely distributed in the Northern hemisphere is represented in Algeria by four species of which Acer monspessulanum is the most common. These four species and their putative hybrids are coexisting in many parts of their Algerian distribution range but their leaf morphology has not been the subject of quantitative analyses despite the interest of such kind of data in taxon delineation particularly in the case of interspecific hybridization. The present work is the first step towards a quantitative analysis of leaf morphology in Acer species in Algeria. We presently relied on traditional and geometric morphometrics methods in the study of Acer monspessulanum subsp. monspessulanum leaf morphology in two sites of the Djurdjura Mountain with consideration of tree and within-shoot effects. The results showed congruence between the two methods which both highlighted the presence of a marked heteroblasty. Basal leaves are twice as large as apical ones for all measured leaf features. The petiole is longer than the blade in basal leaves and inversely in apical ones. The median leaves have intermediate values. The results revealed also the coexistence of two contrasted leaf morphotypes on distinct trees of both sites.

When less is more and more is less: the impact of sampling effort on species delineation

AbstractTaxonomy is the very first step of most biodiversity studies, but how confident can we be in the taxa delineation? One may hypothesize that the more abundant the material, the more accurate the description of morphological variability and hence the better the taxonomic delineation. Yet, as we shall see, in the case of numerous transitional forms, this hypothesis may prove wrong. Similarly to rarefaction curves that assess the degree of knowledge on taxonomic diversity through sampling effort, we aim to test the impact of sampling effort on species delineation by subsampling a given assemblage. To do so, we use an abundant and morphologically diverse conodont fossil assemblage from the Smithian of Oman. We first recognize four well established morphospecies but about 80% of the specimens are transitional forms. We quantify the diagnostic characters in a sample of 159 P1 elements using geometric morphometrics and assess, via gradually subsampling the assemblage, the number of morphometric groups (i.e. morphospecies) using ordination and clustering analyses. Four morphospecies were detected when less than 20% of the specimens were considered. The number of detected clusters dropped to two when including more than 30% of the specimens. Such influence of sampling effort on species delineation highlights the complexity of taxonomic work, especially when transitional forms are more abundant than typical specimens. These results should encourage researchers to extensively illustrate, measure and quantitatively compare their material to better constrain the morphological variability and delineation of taxa.

Molecular Evaluation of the Fairy Shrimp Family Branchinectidae (Crustacea: Anostraca) Supports Peripatric Speciation and Complex Divergence Patterns.

The Branchinectidae is a diverse and widely distributed group of anostracans. The majority of work on the group has focused on the morphological delineation of taxa and biogeography. Here we present a molecular phylogeny for select members of the family to better understand the distribution of morphological variation among species, and test biogeographic models of speciation for the group. Although we conducted both molecular and morphological phylogenies for the Branchinectidae, the morphological analysis did not support our molecular phylogeny and it did not support previous species group concepts based on geography. Our molecular phylogenetic analysis suggests that the family may have originated in Eurasia and found support for numerous species groups. These phylogenetic groups assisted in delineating species groups that are all definable morphologically and/or ecologically. The peripatric speciation model was supported from our analysis, offering credence to previously published speciation models in anostracans. This suggests that these processes may be important in other Branchiopoda and should be rigorously evaluated when delineating species.

Targeted amplicon sequencing of 40 nuclear genes supports a single introduction and rapid radiation of Hawaiian Metrosideros (Myrtaceae)

Compared to traditionally used plastid or ribosomal markers, nuclear gene markers provide many advantages for molecular systematics of plants, and high-throughput sequencing technologies are making large nuclear datasets available at an unprecedented rate. We used targeted amplicon sequencing of 44 nuclear genes to construct a time-calibrated phylogeny of genus Metrosideros (Myrtaceae), evaluate recent systematic revisions, and assess whether phylogenetic signal within the Hawaiian Archipelago is correlated with island biogeography or morphological diversification. We generated a final dataset of 40 nuclear genes for 187 specimens, used multiple search heuristics and species-tree analysis to estimate a phylogeny, and incorporated new fossils for the genus to estimate divergence times across the dataset. All analyses supported the monophyly of Metrosideros, including Carpolepis and Tepualia. Hawaiian Metrosideros were monophyletic and dated to 3.1 MYA using new fossils for the genus, which is intermediate to previous estimates based on nuclear ribosomal/chloroplast loci and calibrated with island ages. Within the Hawaiian Metrosideros clade, we observed short branch lengths and unresolved relationships, and phylogenetic patterns were not concordant with biogeographic hypotheses of island progression, or the delineation of taxa or morphotypes. Average nucleotide diversity was relatively consistent across the Hawaiian Islands with the exception of slightly lower diversity on Kauaʻi. These results provide a data-rich estimate of the timing of a single introduction of Metrosideros to Hawai‘i and highlight the need for molecular markers with higher evolutionary rates for resolution of relationships within this recent radiation.

Catenula exigua sp. nov., a new marine diatom (Bacillariophyta) species from the Adriatic Sea

During a survey of the epizoic diatom flora on carapaces of loggerhead sea turtles (Caretta caretta) from the Adriatic Sea, a relatively small population of an unknown amphoroid species was observed, presenting some morphological similarities with the genera Lunella and Catenula. Due to the small size and the lack of distinct structures visible in LM, the observations of its ultrastructure under very high magnification in the scanning electron microscopy (SEM) proved to be critical in taxon delineation and indicated that it should be separated as a new Catenula species. The new species is characterized by very small, dorsiventral valves, striae composed of 1–3 small, marginally placed areolae, a very simple raphe structure lacking terminal raphe fissures, straight simple central raphe endings, the absence of helictoglossa and a distinct central nodule, the latter often discernible in LM. Typical ghost striae, reported to be one of the main morphological features of the genus Catenula, are very faintly present and only visible in the valve interior. The new species is illustrated using both LM and SEM observations and compared with all other known Catenula and Lunella species.

Delineating species along shifting shorelines: Tropheus (Teleostei, Cichlidae) from the southern subbasin of Lake Tanganyika

BackgroundSpecies delineation is particularly challenging in taxa with substantial intra-specific variation. In systematic studies of fishes, meristics and linear measurements that describe shape are often used to delineate species. Yet, little is known about the taxonomic value of these two types of morphological characteristics. Here, we used Tropheus (Teleostei, Cichlidae) from the southern subbasin of Lake Tanganyika to test which of these types of characters best matched genetic lineages that could represent species in this group of stenotypic rock-dwelling cichlids. We further investigated intra-population variation in morphology. By linking this to a proxy of a population’s age, we could assess the evolutionary stability of different kinds of morphological markers.ResultsMorphological data was collected from 570 specimens originating from 86 localities. An AFLP approach revealed the presence of five lineages in the southern subbasin: T. moorii, T. brichardi, T. sp. ‘maculatus’, T. sp. ‘Mpimbwe’ and T. sp. ‘red’, which we consider to represent distinct species. Although both types of morphological data supported this classification, a comparison of PST-values that describe inter-population morphological differentiation, revealed a better correspondence between the taxon delineation based on AFLP data and the patterns revealed by an analysis of meristics than between the AFLP-based taxon delineation and the patterns revealed by an analysis of shape. However, classifying southern populations of Tropheus was inherently difficult as they contained a large amount of clinal variation, both in genetic and in morphological data, and both within and among species. A scenario is put forward to explain the current-day distribution of the species and colour varieties and the observed clinal variation across the subbasin’s shoreline. Additionally, we observed that variation in shape was larger in populations from shallow shores whereas populations from steep shores were more variable in meristics. This difference is explained in terms of the different timescales at which small and large scale lake level fluctuations affected populations of littoral cichlids at steep and shallow shores.ConclusionsOur results showed meristics to be more evolutionary stable, and of higher taxonomic value for species delimitation in Tropheus, than linear measurements that describe shape. These results should be taken into account when interpreting morphological differences between populations of highly stenotypic species, such as littoral cichlids from the Great East African Lakes.

Resolving relationships and phylogeographic history of the Nyssa sylvatica complex using data from RAD-seq and species distribution modeling

Nyssa sylvatica complex consists of several woody taxa occurring in eastern North America. These taxa were recognized as two or three species including three or four varieties by different authors. Due to high morphological similarities and complexity of morphological variation, classification and delineation of taxa in the group have been difficult and controversial. Here we employ data from RAD-seq to elucidate the genetic structure and phylogenetic relationships within the group. Using the genetic evidence, we evaluate previous classifications and delineate species. We also employ Species Distribution Modeling (SDM) to evaluate impacts of climatic changes on the ranges of the taxa and to gain insights into the relevant refugia in eastern North America. Results from Molecular Variance Analysis (AMOVA), STRUCTURE, phylogenetic analyses using Maximum likelihood, Bayesian Inference, and Splittree methods of RAD-seq data strongly support a two-clade pattern, largely separating samples of N. sylvatica from those of N. biflora-N. ursina mix. Divergence time analysis with BEAST suggests the two clades diverged in the mid Miocene. The ancestor of the present trees of N. sylvatica was suggested to be in the Pliocene and that of N. biflora-N. ursina mix in the end of the Miocene. Results from SDM predicted a smaller range in the southern part of the species present range of each clade during the Last Glacial Maximum (LGM). A northward expansion of the ranges during interglacial period and a northward shift of the ranges in the future under a model of global warming were also predicted. Our results support the recognition of two species in the complex, N. sylvatica and N. biflora, following the phylogenetic species concept. We found no genetic evidence supporting recognitions of intraspecific taxa. However, we propose subsp. ursina and subsp. biflora within N. biflora due to their distinction in habits, distributions, and habitats. Our results further support movements of trees in eastern North America in response to climatic changes. Finally, our study demonstrates that RAD-seq data and a combination of population genomics and SDM are valuable in resolving relationship and biogeographic history of closely related species that are taxonomically difficult.

Evaluation of Morphological Characteristics to Delineate Taxa of the Genus Trigonopyxis (Amoebozoa, Arcellinida)

Morphological features are often the only characteristics suitable for identification of taxa in testate amoebae, especially in ecological and palaeoecological studies. However, whereas the morphology of some species is rather stable it may vary considerably in others. Within the order Arcellinida the genus Trigonopyxis with the type species Trigonopyxis arcula is morphologically highly variable. To identify reliable characteristics for morphology-based taxon delineation we investigated variations in shell size, pseudostome diameter and pseudostome form in T. arcula from three different sites of the Ecuadorian Andes, where these characteristics vary even more than previously described. Further, we investigated if morphological characteristics in Trigonopyxis varied with changes in environmental factors. We studied 951 shells of Trigonopyxis collected along an altitudinal gradient with varying abiotic factors. We established a method for characterization of the pseudostome form, which lead to five different morphotypes. Our results suggest that shell size alone is not an appropriate character for taxon delineation but can be used as an indicator for changes in environmental conditions. In contrast, the pseudostome form might be used for taxon delineation, but likely also varies considerably within taxa. Overall, the study provides an overview of the morphological variability of the genus Trigonopyxis.

Molecular Phylogeny and Ultrastructure of Aphelidium desmodesmi, a New Species in Aphelida (Opisthosporidia).

Aphelids are a diverse group of intracellular parasitoids of algae and diatoms, and are sister to true fungi. Included in four genera, the 14 described species utilize phagocytosis as their mode of nutrition, and the life cycles of these taxa are remarkably similar. However, their putative specificity of host, morphological and ultrastructural features, and genetic divergence have been considered in taxon delineation. Here, we examine the host specificity, morphology, ultrastructure, and molecular 18S gene sequence of a new species in Aphelida, Aphelidium desmodesmi sp. nov. This taxon is in a well-supported clade with two other species of Aphelidium, and this lineage is sister to Amoeboaphelidium and Paraphelidium. Of interest, the mitochondrial structure of Aph.desmodesmi is more like that of Paraphelidium than that of Aphelidium aff. melosirae, the only other species of Aphelidium to have been examined ultrastructurally. This research examines and expands our understanding of host range, morphological diversity, and genetic divergence of the aphelids.

Families of Sordariomycetes

Sordariomycetes is one of the largest classes of Ascomycota that comprises a highly diverse range of fungi characterized mainly by perithecial ascomata and inoperculate unitunicate asci. The class includes many important plant pathogens, as well as endophytes, saprobes, epiphytes, coprophilous and fungicolous, lichenized or lichenicolous taxa. They occur in terrestrial, freshwater and marine habitats worldwide. This paper reviews the 107 families of the class Sordariomycetes and provides a modified backbone tree based on phylogenetic analysis of four combined loci, with a maximum five representative taxa from each family, where available. This paper brings together for the first time, since Barrs’ 1990 Prodromus, descriptions, notes on the history, and plates or illustrations of type or representative taxa of each family, a list of accepted genera, including asexual genera and a key to these taxa of Sordariomycetes. Delineation of taxa is supported where possible by molecular data. The outline is based on literature to the end of 2015 and the Sordariomycetes now comprises six subclasses, 32 orders, 105 families and 1331 genera. The family Obryzaceae and Pleurotremataceae are excluded from the class.

Hypothesized evolutionary trends in zoospore ultrastructural characters in Chytridiales (Chytridiomycota)

Chytridiales is an order of zoosporic fungi currently comprising species representing 19 genera. Although morphologically and genetically diverse, these taxa have in common a zoospore with a suite of ultrastructural characters unique among Chytridiomycota. However, multiple states have been reported for almost every character that defines the Chytridiales zoospore. Two zoospore types have been recognized, each corresponding to a family. Here we examine zoospore ultrastructure of 52 isolates in Chytridiales and assess states for six characters to hypothesize evolutionary trends, using parsimony ancestral state reconstruction for evolutionary analysis. Based on suites of character states, we describe four additional zoospore types in Chytridiales. Five of the six characters ([i] location of the nucleus, [ii] morphology of the kinetosome-associated structure, [iii] complexity of the microtubular root, [iv] microbody-lipid globule complex cisterna structure and [v] thickness of the flagellar plug) revealed ancestral and derived states. The sixth character, structure of the paracrystalline inclusion, did not resolve ancestral and derived states. In each of the lineages within Chytridiales, the evolutionary trend appears to have been from a more complex zoospore to a less complex zoospore with reduced features. As we isolate and analyze additional taxa, we discover new ultrastructural character states that assist in taxon delineation and phylogenetic interpretation.

‘Candidatus Phytoplasma pruni’, a novel taxon associated with X-disease of stone fruits, Prunus spp.: multilocus characterization based on 16S rRNA, secY, and ribosomal protein genes

X-disease is one of the most serious diseases known in peach (Prunus persica). Based on RFLP analysis of 16S rRNA gene sequences, peach X-disease phytoplasma strains from eastern and western United States and eastern Canada were classified in 16S rRNA gene RFLP group 16SrIII, subgroup A. Phylogenetic analyses of 16S rRNA gene sequences revealed that the X-disease phytoplasma strains formed a distinct subclade within the phytoplasma clade, supporting the hypothesis that they represented a lineage distinct from those of previously described 'Candidatus Phytoplasma' species. Nucleotide sequence alignments revealed that all studied X-disease phytoplasma strains shared less than 97.5 % 16S rRNA gene sequence similarity with previously described 'Candidatus Phytoplasma' species. Based on unique properties of the DNA, we propose recognition of X-disease phytoplasma strain PX11CT1(R) as representative of a novel taxon, 'Candidatus Phytoplasma pruni'. Results from nucleotide and phylogenetic analyses of secY and ribosomal protein (rp) gene sequences provided additional molecular markers of the 'Ca. Phytoplasma pruni' lineage. We propose that the term 'Ca. Phytoplasma pruni' be applied to phytoplasma strains whose 16S rRNA gene sequences contain the oligonucleotide sequences of unique regions that are designated in the formally published description of the taxon. Such strains include X-disease phytoplasma and--within the tolerance of a single base difference in one unique sequence--peach rosette, peach red suture, and little peach phytoplasmas. Although not employed for taxon delineation in this work, we further propose that secY, rp, and other genetic loci from the reference strain of a taxon, and where possible oligonucleotide sequences of unique regions of those genes that distinguish taxa within a given 16Sr group, be incorporated in emended descriptions and as part of future descriptions of 'Candidatus Phytoplasma' taxa.

Mitochondrial DNA sequences suggest unexpected phylogenetic position of Corso-Sardinian grass snakes (Natrix cetti) and do not support their species status, with notes on phylogeography and subspecies delineation of grass snakes

We supplement a previously published mitochondrial DNA data set of grass snake sequences (ND1, ND2, ND4, cyt b, in total 3,806 bp) with sequences of Corso-Sardinian and Tuscan specimens and infer their phylogeny using Bayesian, maximum likelihood and maximum parsimony methods. In addition, we estimate divergence times of grass snake clades using a relaxed molecular clock calibrated with fossil evidence, and, in a second approach, the post-Messinian reopening of the Strait of Gibraltar. Recently it was suggested that Corso-Sardinian grass snakes represent a distinct species: Natrix cetti. All tree-building methods revealed well-supported branching patterns and deep divergences among grass snakes. However, sequences of N. natrix were consistently paraphyletic with respect to Corso-Sardinian sequences. The sister group of Corso-Sardinian grass snakes is a clade embracing N. n. helvetica and N. n. lanzai. Extensive gene flow between N. n. helvetica and a more distantly related subspecies (N. n. natrix) is well known, which is why we conclude that the status of Corso-Sardinian grass snakes as subspecies of N. natrix should be reinstated. Many currently recognized grass snake subspecies conflict with mitochondrial clades, suggestive of inappropriate morphological taxon delineation and mitochondrial introgression. Divergences among grass snakes are old, and the results of the two independent dating approaches are largely congruent. Accordingly, the Alpine orogenesis seems to have caused the origin of the oldest clade, corresponding to Iberian N. n. astreptophora. The formation of Corso-Sardinian grass snakes was dated to the Early Pliocene and could result from post-Messinian flooding of the Mediterranean Basin. Another deeply divergent clade of approximately the same age, endemic in central and northern Europe, suggests the Pleistocene survival of grass snakes north of the Alps. At least one glacial refuge in which old lineages survived Pleistocene cold periods was located on each of the three major southern European peninsulas and in Anatolia. Due to pronounced sequence divergences among Italian and southern Swiss grass snakes, we hypothesize multiple refugia south of the Alps and in the Apennine Peninsula, and there is evidence for two refuges on the Balkan Peninsula.

An assessment of fungal wall heteromannans as a phylogenetically informative character in ascomycetes

The fungal wall contains a small proportion of alkali-extractable water-soluble heteromannans (F1SS). They are the glycidic moiety of glycoproteins that have important roles in the biology of fungi. A considerable number of these polysaccharides has been described, differing in composition or linkage types. Their structure is similar in all species of a well-delimited genus, and teleomorphs and their corresponding anamorphs. Therefore, these polysaccharides have been used as chemotaxonomic markers at the genus level. Here we review cases where they have been found to resolve relationships around the genus level, and assess their phylogenetic informativeness in the delineation of taxa at family and higher ranks in the ascomycetes by comparison with molecular trees. Generally, the correlation is extremely good, from the species to the class level, though there are some divergences. In particular, comparisons suggest that the concept of the Sordariomycetes may eventually require revision as more molecular data become available. An analysis of the different chemical structures of these polysaccharides can lead to the proposal and testing of phylogenetic hypotheses, in a parallel manner to those generated from molecular trees. These molecules serve as an independent character similar to morphological or molecular characters.

DNA phylogeny reveals polyphyly of Phoma section Peyronellaea and multiple taxonomic novelties

Species of the anamorph genus Phoma are commonly isolated from a wide range of ecological niches. They are notoriously difficult to identify due to the paucity of morphological features and the plasticity of these when cultivated on agar media. Species linked to Phoma section Peyronellaea are typified by the production of dictyochlamydospores and thus have additional characters to use in taxon delineation. However, the taxonomy of this section is still not fully understood. Furthermore the production of such chlamydospores also is known in some other sections of Phoma. DNA sequences were generated from three loci, namely ITS, actin, and β-tubulin, to clarify the phylogeny of Phoma taxa that produce dictyochlamydospores. Results were unable to support section Peyronellaea as a taxonomic entity. Dictyochlamydospore formation appears to be a feature that developed, or was lost, many times during the evolution of Phoma. Furthermore, based on the multigene analyses, five new Phoma species could be delineated while a further five required taxonomic revision to be consistent with the genetic variation observed.

Proposal for a New Hierarchic Classification System, Actinobacteria classis nov.

A new hierarchic classification structure for the taxa between the taxonomic levels of genus and class is Proposed for the actinomycete line of descent as defined by analysis of small subunit (16S) rRNA and genes coding for this molecule (rDNA). While the traditional circumscription of a genus of the actinomycete subphylum is by and large in accord with the 16S rRNA/rDNA-based phylogenetic clustering of these organisms. most of the higher taxa proposed in the past do not take into account the phylogenetic clustering of genera. The rich chemical, morphological and physiological diversity of phylogenetically closely related genera makes the description of families and higher taxa so broad that they become meaningless for the description of the enclosed taxa. Here we present a classification system in which phylogenetically neighboring taxa at the genus level are clustered into families, suborders, orders, subclasses, and a class irrespective of those phenotypec characteristics on which the delineation of taxa has been based in the past. Rather than being based on a listing of a wide array of chemotaxonomic, morphological, and physiological properties, the delineation is based solely on 16S rDNA/rRNA sequence-based phylogenetic clustering and the presence of taxon-specific 16S rDNA RNA signature nucleotides.

Diversity of glucose entry routes in the Enterobacteriaceae

One of the purposes of bacterial taxonomy is to delineate groups (taxa) about which one can generalize knowledge obtained by the study of a few strains in these groups. The delineation of taxa is now achieved using molecular methods such as DNA-DNA hybridization, rRNA-DNA hybridization or rRNA sequencing. However, the description of these carefully delineated taxa rests on empirical 'biochemical' tests. Obviously, the fantastic diversity of metabolic pathways in bacteria is insufficiently used in taxonomy. Our group is engaged in taxonomic research and we want to define bacterial taxa on a sound biochemical and physiological basis. Our first project in that area concerned glucose entry routes in the Enterobacteriaceae. There are currently about a hundred genomic species (i.e. species defined by DNA relatedness, whether named or not) in the family Enterobacteriaceae. Most of our knowledge concerning the physiology and genetics of this family is limited to Escherichia coli K-12 and Salmonella typhimuri-

Diversity of glucose entry routes in the Enterobacteriaceae

One of the purposes of bacterial taxonomy is to delineate groups (taxa) about which one can generalize knowledge obtained by the study of a few strains in these groups. The delineation of taxa is now achieved using molecular methods such as DNA-DNA hybridization, rRNA-DNA hybridization or rRNA sequencing. However, the description of these carefully delineated taxa rests on empirical 'biochemical' tests. Obviously, the fantastic diversity of metabolic pathways in bacteria is insufficiently used in taxonomy. Our group is engaged in taxonomic research and we want to define bacterial taxa on a sound biochemical and physiological basis. Our first project in that area concerned glucose entry routes in the Enterobacteriaceae. There are currently about a hundred genomic species (i.e. species defined by DNA relatedness, whether named or not) in the family Enterobacteriaceae. Most of our knowledge concerning the physiology and genetics of this family is limited to Escherichia coli K-12 and Salmonella typhimuri-

Reproductive Isolation in Sedum sect. Gormania (Crassulaceae)

Crossing and cytological studies demonstrate the nature and extent of reproductive isolation among the 12 taxa of Sedum sect. Gormania. Artificial hybridizations between the taxa show that crossing barriers are few or essentially nonexistent. An analysis of the synthesized hybrids, including assessments of interfertilities, pollen stainabilities, and chromosomal configurations, con- firms that genetic isolation of the taxa operates at the F1 generation. That intertaxon hybrids are sterile or scarcely interfertile supports the present delineation of taxa and provides clues to their evolutionary pathways. Hypotheses regarding the origin of the polyploid taxa are supported: 1) that S. laxum subsp. eastwoodiae (4x) arose from S. laxum subsp. laxum (2x) and/or S. laxum subsp. heckneri (2x); 2) that S. laxum subsp. flavidum (4x) arose from S. laxum subsp. heckneri (2x) and S. obtusatum subsp. obtusatum (2x); 3) that S. obtusatum subsp. retusum (4x) arose from S. obtusatum subsp. obtusatum (2x) and/or S. obtusatum subsp. boreale (2x); and 4) that S. oregonense (6x) arose via alloploidy from S. obtusatum subsp. retusum (4x) and S. laxum subsp. laxum (2x).