Investigations into the evolutionary history of the polyploid complexAntennaria neodioica (Asteraceae: Inuleae)

TheAntennaria neodioica polyploid agamic complex is a polymorphic species occurring across North America mainly north of the terminal margin of the Wisconsin glacier. This taxonomically difficult group has recently been treated as consisting of the four subspeciesA. neodioica subsp.canadensis, subsp.howellii, subsp.neodioica, and subsp.petaloidea. TheA. neodioica agamic complex has been considered of hybrid origin with several sexual diploid species constituting its parentage. Crosses were made among five sexual diploid species ofAntennaria, morphologically similar toA. neodioica s.l., in an attempt to discover its origins. Representative specimens of the five diploid species,A. neodioica s. l., and the synthetic interspecific F₁ hybrids were subjected to various analyses including PCA, cluster (UPGMA), and discriminant analyses. Results suggest that theA. neodioica complex is of multiple hybrid origin involving the four diploid speciesA. neglecta, A. plantaginifolia, A. racemosa, andA. virginica. BecauseA. neodioica is the result of diverse origin it is more desirable to consider the agamic complex as a separate, distinct species from its sexual diploid relatives. Several morphological characters in the diploid species were determined to be polygenically inherited.Investigations into the Evolutionary History of the Polyploid Complexes inAntennaria (Asteraceae: Inuleae). I.     

ALLOZYME DIVERGENCE AMONG FIVE DIPLOID SPECIES OF ANTENNARIA (ASTERACEAE: INULEAE) AND THEIR ALLOPOLYPLOID DERIVATIVES

The Antennaria parlinii and A. neodioica agamic complexes are widely distributed across North America. Morphological data have suggested that these dioecious, perennial, entire-leaved herbs are of multiple hybrid origin from among five sexual diploid species. Antennaria neglecta, A. plantaginifolia, A. racemosa, and A. virginica are hypothesized to be the diploid progenitors of the A. neodioica complex, whereas A. parlinii sensu lato is thought to include the genomes of A. plantaginifolia, A. racemosa, and A. solitaria. An electrophoretic study was initiated to assess the degree of divergence among the five diploid species and to test the hypotheses of the hybrid (allopolyploid) origins of A. parlinii and A. neodioica. Twenty genetic loci were surveyed in 76 populations of the diploid and polyploid taxa. The diploid species are well defined morphologically, although the genetic basis of differences distinguishing them have not been determined. The species exhibit little divergence at genes specifying soluble enzymes, however each species has unique alleles in highest frequency at one or two genes. Allozymes indicate that gene diversity in the obligately outcrossing diploids occurs primarily within rather than among populations. The shale barren endemic, A. virginica, is as genetically diverse as the more edaphically diverse and widespread species. Tetraploid cytotypes of diploid (2n = 28) A. virginica possess the same allozymes as the diploids and these cytotypes appear to be of autopolyploid (non-hybrid) origin. Enzyme electrophoresis is concordant with morphological data in suggesting that A. neodioica contains the genomes of A. neglecta, A. virginica, A. plantaginifolia, A. racemosa and perhaps A. solitaria whereas the latter three species are the progenitors of A. parlinii.     

PATTERNS OF ISOZYME VARIATION IN WESTERN NORTH AMERICAN ANTENNARIA (ASTERACEAE: INULEAE) II. DIPLOID AND POLYPLOID SPECIES OF SECTION ALPINAE

Antennaria is a genus of dioecious, perennial, herbaceous Composites that are especially diverse in the cordillera of western North America. Section Alpinae consists of about nine taxa, among them A. aromatica, A. densifolia, A. media, A. pulchella, and A. umbrinella. Although diploids are morphologically distinct, the polyploid derivatives of the diploids obscure the morphological distinctness of the groups. A survey of 19 putative isozyme loci indicates that the diploids have diverged only moderately from one another with respect to biochemical genetics (I = 0.838 to 0.961). Additionally, only moderate amounts of genetic diversity were detected. Isozyme data are supportive of a hypothesis of a rapid mode of speciation in Antennaria, where morphological differentiation has been accompanied by small amounts of allozyme divergence. Polyploids have significantly higher amounts of heterozygosity than diploids and tetrasomic inheritance is inferred. Evidence from morphology and biochemical genetics suggests that the polyploids represent a continuum between interracial autopolyploids and segmental allopolyploids. In light of the relatively low degree of genetic and morphological divergence among many Antennaria species, taxonomic conservatism is advocated.     

ISOZYME NUMBER IN SUBTRIBE ONCIDIINAE (ORCHIDACEAE): AN EVALUATION OF POLYPLOIDY

The Oncidiinae has attracted attention because of the variation it exhibits in chromosome number, n = 5–30, which is greater than the range in the rest of the Orchidaceae. The genus Psygmorchis, with n = 5 and 7, has been a particular focus of controversy, and many authors have suggested that 5 and 7 are the base numbers for the subtribe. The other taxa in the subtribe presumably evolved through hybridization and polyploidy. Other workers have found that the lowest counts correlate with derived morphological conditions and have hypothesized that these low numbers result from aneuploid reductions, while higher numbers are associated with ancestral morphologies and are not the result of polyploidy. These two hypotheses were evaluated by determining isozyme numbers for 13 enzymes in species that span the chromosomal range known for the Oncidiinae (n = 5–30). Isozyme number has been shown to be a reliable indicator of polyploidy in angiosperms because polyploids display isozyme multiplicity relative to diploids. This analysis revealed no differences among species in isozyme number for the enzymes examined. Therefore, our data reject the hypothesis that species with higher chromosome numbers are polyploid.     

Polyploidy in Asteraceae of the xerophytic scrub of the Ecological Reserve of the Pedregal of San Angel,Mexico City

Asteraceae species diversity is high in the xerophytic scrub of the Ecological Reserve of the Pedregal of San Angel (REPSA), located in the southern part of the Basin of the Valley of Mexico. Here we determined whether the frequency of polyploidy is high in the reserve, given the enhanced ability of polyploids to colonize new habitats. In addition, we compared the frequency of polyploidy in Asteraceae in the reserve with the frequency in three oceanic archipelagos and two continental areas in Mexico. This was done to see how the ‘virtual’ island of the open lava flow in the reserve compares with volcanic islands at different distances from source areas. Chromosome numbers for 75 species of Asteraceae were obtained from published literature. Based on the possession of three or more basic chromosome sets in a nucleus, 33% were polyploids. If taxa with haploid chromosome numbers of n ≥ 14 or n ≥ 11 were considered to be polyploids, the proportion of polyploids rose to 57 and 75%, respectively. When using a phylogenetic approach, the highest percentage of polyploids (84%) was obtained and it could be inferred whether they are palaeo‐ or neopolyploids; thus, we consider that this criterion better reflects the events of polyploidy in Asteraceae. A high frequency of polyploid species in Asteraceae in REPSA suggests that polyploids may have contributed to the species diversity and the vegetation structure of the xerophytic scrub of this reserve. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 211–229.     

EVOLUTIONARY STUDIES OF ATRIPLEX: CHROMOSOME RACES OF A. CONFERTIFOLIA (SHADSCALE)

Atriplex confertifolia (Torr. and Frem.) S. Wats., a species endemic to western United States, occurs as 2n, 4n, 6n, 8n and 10n chromosome races. In the Great Basin, diploids occur altitudinally above the upper levels of Pleistocene lakes; the polyploids are in valley bottoms and sides. Diploids are larger in all respects than tetraploids, hexaploids, and octoploids. Decaploids are the most robust of all. Diploids can be distinguished in early seedling stages by having broader leaves and shorter internodes than polyploids. Polyploid plants are currently produced sporadically in natural populations and appear to be easily generated by unreduced gametes. Most polyploid populations in the Great Basin apparently arose recently and erupted into extensive stands because of the sudden availability of large, relatively uniform domains provided by the disappearance of Pleistocene lakes.     

Light reduction predicts widespread patterns of dominance between asters and goldenrods

Here we investigate the long-cited pattern that throughout the eastern United States, Solidago species (goldenrods), and in particular S. canadensis displace Aster species and dominate old-field communities. Theory predicts that such a ubiquitous pattern of repeated dominance should be linked to competitive ability for a limiting resource. However, no one has investigated this possibility in old-fields, representing a potentially significant gap in our understanding of a common human-altered environment. We tested the hypothesis that S. canadensis is the superior competitor for light compared to other common co-occurring goldenrod species, and that the goldenrods in general are the superior competitors for light compared to coexisting aster species, which are typically less abundant. We tested this hypothesis by comparing the light attenuation abilities of four goldenrod species, S. canadensis, S. rugosa, S. gigantea, and Euthamia graminifolia, and three aster species, Aster novae-angliae, A. pilosus, and A. prenanthoides. Consistent with our hypothesis, S. canadensis had a greater ability to attenuate light than any of the other goldenrods at higher densities, and the goldenrods overall had a greater ability to attenuate light than the asters. By conducting a census in our study area, we verified that S. canadensis is locally the most abundant goldenrod and that goldenrods are more locally abundant than asters. Furthermore, by conducting a literature survey we found evidence that S. canadensis replaces A. pilosus through time. Thus we found a close correspondence between relative abundance in the field and light attenuation ability in field experiments. These results are consistent with theory predicting that competition for limiting resources, in this case light, explains patterns of dominance and relative abundance in old-field plant communities.     

ELECTROPHORETIC ENZYME ANALYSIS OF NORTH AMERICAN AND EASTERN ASIAN POPULATIONS OF AGASTACHE SECT. AGASTACHE (LABIATAE)

Agastache sect. Agastache consists of seven species in North America and one disjunct in eastern Asia. Starch-gel electrophoresis of enzymatic proteins was employed to assess genetic relationships among these species and to estimate the amount of genetic divergence between the North American and Asian populations. Species of the western United States appear to be better adapted for outcrossing than are the others and are much more genetically variable, with higher levels of heterozygosity per individual, more alleles per species, and higher percentages of polymorphic loci per population. Nonmetric multidimensional scaling of Nei's genetic distances among 32 populations partitioned the section into four discrete groups: 1) A. nepetoides (eastern North America), 2) A. scrophulariifolia and A. foeniculum (eastern and central North America), 3) the four species of the western United States (A. urticifolia, A. occidentalis, A. parvifolia and A. cusickii) and 4) A. rugosa (eastern Asia). Asian Agastache, separated from its American congeners for over twelve million years, differed from American populations at two of fifteen loci surveyed. Nei's genetic distances between Asian and North American populations ranged from 0.2877 to 0.6734.     

Widespread inbreeding and unexpected geographic patterns of genetic variation in eastern hemlock (Tsuga canadensis), an imperiled North American conifer

Eastern hemlock (Tsuga canadensis [L.] Carr.) is an ecologically important tree species experiencing severe mortality across much of its eastern North American distribution, caused by infestation of the exotic hemlock woolly adelgid (Adelges tsugae Annand). To guide gene conservation strategies for this imperiled conifer, we conducted a range-wide genetic variation study for eastern hemlock, amplifying 13 highly polymorphic nuclear microsatellite loci in 1,180 trees across 60 populations. The results demonstrate that eastern hemlock exhibits moderate inbreeding, possibly a signature of a prehistoric decline associated with a widespread insect outbreak. Contrary to expectations, populations in formerly glaciated regions are not less genetically diverse than in the putative southern refugial region. As expected, peripheral disjunct populations are less genetically diverse than main-range populations, but some are highly genetically differentiated or contain unique alleles. Spatially explicit Bayesian clustering analyses suggest that three or four Pleistocene glacial refuges may have existed in the Southeastern United States, with a main post-glacial movement into the Northeast and the Great Lakes region. Efforts to conserve eastern hemlock genetic material should emphasize the capture of broad adaptability that occurs across the geographic range of the species, as well as genetic variability within regions with the highest allelic richness and heterozygosity, such as the Southern Appalachians and New England, and within disjunct populations that are genetically distinct. Much genetic variation exists in areas both infested and uninfested by the adelgid.     

BIOSYSTEMATIC INVESTIGATIONS IN SOUTH FLORIDA POPULATIONS OF RUELLIA (ACANTHACEAE)

Relationships of the putative Ruellia species, R. caroliniensis, R. ciliosa, R. heteromorpha, and R. succulenta were analyzed by population sampling, investigation of chromosome numbers, garden transplants, and hybridization experiments. Character data from 25 local, Florida populations and from transplants were evaluated. Some interfertility of these “taxa” was demonstrated, as was some interfertility with other species occurring in the eastern United States. Chromosome numbers are n = 17 and 2n = 34. The effect of different breeding systems on population structure is discussed. Total evidence suggests that the 4 “taxa” in question be considered ecological races of a single species widespread in the eastern United States, namely R. caroliniensis.     

CHEMICAL CHANGES IN LESPEDEZA ASSOCIATED WITH INDUCED POLYPLOIDY

Hanson , C. H. (U.S.D.A., Beltsville, Md.), W. A. Cope and R. M. Brinkley . Chemical changes in Lespedeza associated with induced polyploidy. Amer. Jour. Bot. 46(1): 36-39. 1959.— Changes in the quantities of 5 chemical constituents which were associated with induced polyploidy were studied using leaves of the following polyploids: Autotetraploids—Lespedeza cuneata, L. latissima, L. stipulacea, and L. striata; amphidiploids—L. hedysaroides × L. cuneata and L. latissima × L. cuneata. L. stipulacea and L. striata were annuals with low concentrations of tannin; the other species were perennials and high in tannin content. The polyploids were generally lower in crude protein, P₂O₅, and CaO, and higher in K₂O and total tannin. The greatest and most significant shift was in total tannin. Induced polyploids of Lespedeza did not appear promising as forage plants.     

Polyploidy in arctic plants

The Arctic is an excellent model system for the study of polyploidy. It is one the Earth's most polyploid‐rich areas, in particular of high‐level and recently evolved polyploids. Here we re‐address previous hypotheses on arctic polyploidy based on a new analysis of the circumarctic flora, and review recent molecular, cytological and reproductive studies. The frequency and level of polyploidy strongly increase northwards within the Arctic. We found no clear‐cut association between polyploidy and the degree of glaciation for the arctic flora as a whole, which contains many widespread species. However, for ‘arctic specialist’ taxa with restricted distributions, the frequency of diploids is much higher in the Beringian area, which remained largely unglaciated during the last ice age, than in the heavily glaciated Atlantic area. This result supports the hypothesis that polyploids are more successful than diploids in colonizing after deglaciation. There is abundant molecular evidence for recurrent formation of arctic polyploids at different scales in time and space. Examples are given of low‐level polyploids formed after the last glaciation and of repeated and successively more high‐level polyploidizations throughout the Quaternary. Recurrent polyploid origins, followed by interbreeding within and across ploidal levels, provide a major explanation for the taxonomic complexity of the arctic flora. In the well‐studied, recently deglaciated archipelago of Svalbard, most species are mainly self‐fertilizing or clonal. All Svalbard polyploids examined so far are genetic allopolyploids with fixed heterozygosity at isozyme loci. The level of heterozygosity in 65 taxa increases dramatically from diploids to high‐level polyploids. In the circumarctic area, there is evidence for numerous recently evolved sibling species within diploid taxonomic species. Rapid evolution of crossing barriers at the diploid level promotes further diversification after expansion from different refugia, and may provide new raw materials for allopolyploid formation. We conclude that the evolutionary success of polyploids in the Arctic may be based on their fixed‐heterozygous genomes, which buffer against inbreeding and genetic drift through periods of dramatic climate change. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 521–536.     

Phylogenetic inferences in Antennaria (Asteraceae: Gnaphalieae: Cassiniinae) based on sequences from Nuclear Ribosomal DNA internal transcribed spacers (ITS)

The phylogenetic relationships among sexually reproducing species of Antennaria (Asteraceae) are poorly understood. An earlier cladistic analysis based on morphology did not fully resolve the phylogeny of these taxa and therefore a different approach using molecular data was explored. The internal transcribed spacer regions (ITS-1 and ITS-2) of nuclear ribosomal DNA were sequenced for 30 species of Antennaria and one species from each of the outgroup genera Anaphalis, Ewartia, Leontopodium, and Pseudognaphalium. The ITS-1 sequence in Antennaria ranged from 253 to 260 base pairs (bp) in length, and the proportion of nucleotide differences between pairs of species of Antennaria ranged from 1 to 14%. For ITS-2, the divergence between pairs of species of Antennaria ranged from 0 to 8%. ITS-2 is shorter than ITS-1, ranging from 213 to 219 bp. Phylogenetic analysis indicates that, relative to the outgroups included, Antennaria is a well-supported monophyletic group. Based on the genera surveyed, Leontopodium appears to be the sister genus of Antennaria. The general topology of the molecular trees agrees with that based on previous morphological analyses and indicates that Antennaria is composed of six clades of equal rank, corresponding to the traditionally recognized informal groups, the Geyeriae, Argenteae, Arcuatae, Dimorphae, Pulcherrimae, and Catipes. Sequence and morphological data indicate that the Alpinae and Dioicae are unnatural, polyphyletic units that should be abandoned and redefined as the monophyletic Catipes group. Phylogenetic analysis of ITS sequences also suggests the dissociation of A. stenophylla from the Dimorphae, where it is traditionally placed, and its affiliation with the Argenteae, as well as the placement of A. arcuata in its own group.     

CHROMOSOME NUMBERS IN SPHAERALCEA SECTION FENDLERIANAE

Chromosome numbers are reported for ten taxa in Sphaeralcea section Fendlerianae (Malvaceae). New ploidy levels are reported for six taxa, with one species not previously reported, and extensive polyploidy at all taxonomic levels is documented. The geographic and taxonomic distribution of polyploids suggests that polyploidy arose many times in the taxa of Sphaeralcea. Sphaeralcea fendleri var. venusta, S. polychroma, and S. wrightii populations have yielded exclusively tetraploid counts. Tetraploidy is correlated with taxa having lavender petals. Polyploidy has also allowed the taxa to expand their distributions without resulting in speciation.     

GEOGRAPHY OF CLAYTONIA VIRGINICA CYTOTYPES

Three widespread ploidy levels have evolved in Claytonia virginica, a diploid represented by n = 8, 7, and 6 races, a triploid primarily by n = 12 and 11, and a tetraploid by n = 16, 15, 14, and other aneutetraploids. In addition, sporadic higher polyploids (5x–24x) occur throughout eastern North America. These data are considered briefly in relation to meiotic behavior, to coexistence of major cytotypes, and to speciation.     

CYTOGEOGRAPHY OF PANICUM VIRGATUM IN CENTRAL NORTH AMERICA

Mc Millan , Calvin . (U. Texas, Austin.), and John Weiler . Cytogeography of Panicum virgatum in central North America. Amer. Jour. Bot. 46(8): 590–593. Illus. 1959.—For 124 clones of Panicum virgatum L. representing 44 local populations from Manitoba and eastern Montana to Texas, the chromosome number was determined. Most of the clones were grown in a transplant garden at Lincoln, Nebraska. Among the earlier-flowering clones from the northern and western area, a preponderance were tetraploids (n = 18). Among the later-flowering clones, the Iowa material was predominantly tetraploid, while those from Nebraska presented a diversity of types (n = 18, 27, 36) within each population. A series of polyploids was common in population samples from southern Kansas, Oklahoma, and Missouri. Through the complex breeding patterns resulting from the common occurrence of a range of polyploidy and the possible action of apomixis, variability within the population is maintained, and with it, genetic insurance for survival in highly unpredictable prairie habitats.     

ON THE METHODS OF EXPERIMENTAL TAXONOMY

By studying seedling progenies from individual plants it may be decided whether the material under investigation is allogamous, autogamous or apomictic. Chromosome counts disclose whether the material is cytologically stable or variable. If variation occurs it may be a question of polyploidy or aneuploidy. Aneuploidy may either be an occasional consequence of polyploidy or it may represent dysploidy and the formation of new basic numbers. Chromosome counts combined with measurements of chromosome size may reveal cases of pseudopolyploidy of the kind first observed in Luzula. If chromosome morphology is different between two taxa this indicates reproductive isolation. Different species usually differ with regard to their chromosome structure. Studies of meiosis are desirable in polyploids in order to distinguish between auto- and alloploidy. Cases of genic control of meiosis in polyploids of Phleum and Triticum demonstrate, however, that partial or complete homology between genomes does not always result in multivalent formation. Apparent alloploids may in reality be more or less autoploid. Incompatibility is not a reliable criterion of species differentiation. Strong or absolute barriers between diploids and autotetraploids may have purely quantitative causes. Hybrid sterility is not always a reliable measure of the degree of genetic difference between the parents, but may be caused by heterozygosity for chromosomal rearrangements. Cytological and experimental methods are, nevertheless, indispensable tools in plant taxonomy.     

CYTOTAXONOMY OF THE ELEOCHARIS TENUIS COMPLEX

Chromosome numbers of 2n = 20, 24, 36, and 38 were found in the various taxa comprising the Eleocharis tenuis complex in the United States. Eleocharis tenuis var. verrucosa, the only taxon in the complex with 2n = 20 is given the new status of a full species [E. verrucosa (Svens.) Harms] and seems most closely allied to two southern members of the subseries Truncatae which are not members of the tenuis complex s. s. Karyotypes and meiotic pairing in artificial hybrids indicate that the remainder of the taxa are derivatives of X = 6 with E. tenuis and one of two cytotypes of E. compressa with the lowest numbers of 2n = 24, although evidence is presented to indicate that they are of amphidiploid origin. A second cytotype of E. compressa with 2n = 36 is a segmental autoallohexaploid, whereas E. elliptica and E. elliptica var. pseudoptera (Svens.) Harms, comb. nov. with 2n = 38 are hyperploid derivatives of 2n = 36.