Origins and introgression of polyploid species in Mentzelia section Trachyphytum (Loasaceae)

? Premise of the study: Polyploid speciation has been important in plant evolution. However, the conditions that favor the origination and persistence of polyploids are still not well understood. Here, we examine origins of 16 polyploid species in Mentzelia section Trachyphytum. ? Methods: We used phylogeny reconstructions based on DNA sequences from plastid regions and the nuclear gene isocitrate dehydrogenase (idh) to construct hypotheses of introgression and polyploidization. ? Key results: Molecular data suggest that homoploid hybridization has been surprisingly common in Trachyphytum. Diploid species had unequal involvement in polyploid origins, but most polyploid taxa had allopolyploid origins from extant progenitors. A few polyploids with extreme phenotypes did not appear to have extant progenitors. We infer that the progenitors of these species were derived from extinct diploid lineages or ancestral lineages of multiple extant diploids. In agreement with other recent studies, we recovered molecular evidence of multiple phylogenetically distinct origins for several polyploid taxa, including the widespread octoploid M. albicaulis. ? Conclusions: Evidence of high levels of introgression and allopolyploidy suggests that hybridization has played an important role in the evolution of Trachyphytum. Although idh sequences exhibited complicated evolution, including gene duplication, deletion, and recombination, they provided a higher percentage of informative characters for phylogeny reconstruction than the most variable plastid regions, allowing tests of hypotheses regarding polyploid origins. Given the necessity for rapidly evolving low-copy nuclear genes, researchers studying hybridization and polyploidy may increasingly turn to complex sequence data.     

Polyploids with different origins and ancestors form a single sexual polyploid species

Polyploidization is one of the few mechanisms that can produce instantaneous speciation. Multiple origins of tetraploid lineages from the same two diploid progenitors are common, but here we report the first known instance of a single tetraploid species that originated repeatedly from at least three diploid ancestors. Parallel evolution of advertisement calls in tetraploid lineages of gray tree frogs has allowed these lineages to interbreed, resulting in a single sexually interacting polyploid species despite the separate origins of polyploids from different diploids. Speciation by polyploidization in these frogs has been the source of considerable debate, but the various published hypotheses have assumed that polyploids arose through either autopolyploidy or allopolyploidy of extant diploid species. We utilized molecular markers and advertisement calls to infer the origins of tetraploid gray tree frogs. Previous hypotheses did not sufficiently account for the observed data. Instead, we found that tetraploids originated multiple times from extant diploid gray tree frogs and two other, apparently extinct, lineages of tree frogs. Tetraploid lineages then merged through interbreeding to result in a single species. Thus, polyploid species may have complex origins, especially in systems in which isolating mechanisms (such as advertisement calls) are affected directly through hybridization and polyploidy.     

Polyploid evolution and biogeography in Chelone (Scrophulariaceae): morphological and isozyme evidence

Chelone is a genus of perennial herbs comprising three diploid species (C. cuthbertii, C. glabra, and C. lyonii) and a fourth species (C. obliqua) that occurs as tetraploid and hexaploid races. To assess patterns of isozyme and morphological variation, and to test hypotheses of hybridization and allopolyploidy, we analyzed variation among 16 isozyme loci from 61 populations and 16 morphological characters from 33 populations representing all taxa and ploidy levels. Based on morphological analyses using clustering (unweighted pair group method using an arithmetic average) and ordination (principal components analysis and canonical variance analysis) methods, we recognize three diploid species without infraspecific taxa. Polyploids in the C. obliqua complex were most similar morphologically to diploid populations of C. glabra and C. lyonii. Patterns of isozyme variation among polyploids, which included fixed heterozygosity and recombinant profiles of alleles present in diploids, suggested polytopic origins of tetraploids and hexaploids. Our data indicate independent origins of polyploids in or near the southern Blue Ridge, Interior Highlands and Plains, and Atlantic Coastal Plain regions from progenitors most similar to C. glabra and C. lyonii. Extant tetraploids were not implicated in evolution of hexaploids, and plants similar to C. cuthbertii appeared unlikely as diploid progenitors for polyploids. We propose multiple differentiation and hybridization/polyploidization cycles in different geographic regions to explain the pattern of allopatry and inferred polytopic origins among polyploids.     

Relationship between species in the genus Rosa, section Pimpinellifoliae

The morphology of twelve species of Rosa is described and similarities between these species are assessed. Possible origins of the tetraploid species from diploid species are indicated on grounds of comparative morphology.
The wild origins of living and herbarium specimens are given in order to supplement published data on geographical distribution.
Meiosis in pollen mother cells, viability of pollen grains at anthesis and ability to set seed was studied in several F₁ hybrids: no indication of complete or even partial sterility was found. Reproductive isolation is therefore unlikely to be maintained by reduced fertility of interspecific hybrids.
Three species are reduced to synonymy with three other species, being retained as subspecific taxa. Two species are transferred from section Pimpinellifoliae to section Cinnamomeae.     

Incorporating allelic variation for reconstructing the evolutionary history of organisms from multiple genes: An example from Rosa in North America

Allelic variation within individuals holds information regarding the relationships of organisms, which is expected to be particularly important for reconstructing the evolutionary history of closely related taxa. However, little effort has been committed to incorporate such information for reconstructing the phylogeny of organisms. Haplotype trees represent a solution when one nonrecombinant marker is considered, but there is no satisfying method when multiple genes are to be combined. In this paper, we propose an algorithm that converts a distance matrix of alleles to a distance matrix among organisms. This algorithm allows the incorporation of allelic variation for reconstructing the phylogeny of organisms from one or more genes. The method is applied to reconstruct the phylogeny of the seven native diploid species of Rosa sect. Cinnamomeae in North America. The glyceralgehyde 3-phosphate dehydrogenase (GAPDH), the triose phosphate isomerase (TPI), and the malate synthase (MS) genes were sequenced for 40 individuals from these species. The three genes had little genetic variation, and most species showed incomplete lineage sorting, suggesting these species have a recent origin. Despite these difficulties, the networks (NeighborNet) of organisms reconstructed from the matrix obtained with the algorithm recovered groups that more closely match taxonomic boundaries than did the haplotype trees. The combined network of individuals shows that species west of the Rocky Mountains, Rosa gymnocarpa and R. pisocarpa, form exclusive groups and that together they are distinct from eastern species. In the east, three groups were found to be exclusive: R. nitida-R. palustris, R. foliolosa, and R. blanda-R. woodsii. These groups are congruent with the morphology and the ecology of species. The method is also useful for representing hybrid individuals when the relationships are reconstructed using a phylogenetic network.     

Anthocyanins in flowers of genus Rosa, sections Cinnamomeae (=Rosa), Chinenses, Gallicanae and some modern garden roses

Forty-four taxa of three sections (Cinnamomeae (=Rosa) 26, Chinenses 8 and Gallicanae 10) and eight modern garden roses in the genus Rosa were surveyed for their floral anthocyanins. Eleven anthocyanins: 3-glucosides and 3,5-diglucosides of cyanidin (Cy), pelargonidin (Pg) and peonidin (Pn), 3-rutinosides and 3-rho-coumaroylglucoside-5-glucosides of Cy and Pn, and Cy 3-sophoroside, were isolated from flowers of these taxa and identified by chemical and spectroscopic techniques. Four anthocyanins: Cy 3-rutinoside, Pn 3-rutinoside, Pn 3-rho-coumaroylglucoside-5-glucoside and Cy 3-sophoroside were found for the first time in Rosa flowers.Investigated sections of wild roses showed characteristic distribution of anthocyanins. Cy 3,5-diglucoside was the dominant anthocyanin detected in all three sections, but accumulation of Pn 3,5-diglucoside distinguished sections Cinnamomeae from other sections, and the occurrence of Cy 3-glucoside separates section Chinenses from others.Cy 3-sophoroside was detected in large amount in some taxa of section Cinnamomeae: e.g., R. moyesii and its related cultivars, and R. rugosa cv. Salmon Pink. The acylated Cy glycoside was found in all sections and also in some modern garden roses, while the acylated Pn glycoside was detected in the section Cinnamomeae, but not in sections Chinenses and Gallicanae. According to anthocyanin distribution patterns, eight groups were classified chemotaxonomically in genus Rosa.     

Polyploidy associated with altered and broader ecological niches in the Claytonia perfoliata (Portulacaceae) species complex

? Premise of the study: Polyploids are often hypothesized to have distinct and broader niches than their diploid progenitors. Differences in geographic distributions between diploid and polyploids are frequently used to infer niche differentiation and increased breadth, but they are seldom used to test these hypotheses explicitly. ? Methods: Niche overlap and breadth were compared for diploids, tetraploids, and hexaploids of three taxa in the Claytonia perfoliata complex (C. parviflora, C. perfoliata, and C. rubra) with the use of species distribution models. Resampling and randomization approaches were used to test hypotheses of niche differentiation, breadth, and conservatism. ? Key results: Niche differentiation was detected between polyploid and diploid cytotypes assigned to the same taxon (e.g., C. parviflora 2× vs. 4×) but not between hexaploids and tetraploids within a taxon (e.g., C. parviflora 4× vs. 6×). Individual hexaploid cytotypes had broader ecological niches than individual diploid cytotypes. However, as a group the three hexaploid taxa did not exceed the combined niche breadth of the three diploids, suggesting that polyploidy does not result in transgressive niche breadth for this group. Niche overlap was lowest among diploids and was highest among the three hexaploid cytotypes, consistent with introgression associated with polyploidy resulting in greater ecological similarity. Although cytotypes possessed nonidentical niches, after accounting for environmental differences among ranges, cytotypes were more similar than expected, suggesting niche conservatism and similar responses to environmental characteristics. ? Conclusions: These results suggest that polyploids occupy distinct and broader niches relative to diploids but that cytotypes also share fundamentally similar responses to environmental variation across ploidy levels.     

Multiple origins of polyploidy and the geographic structure of Heuchera grossulariifolia

Multiple origins of polyploidy from an ancestral diploid plant species were investigated using restriction site polymorphism and sequence variation in the chloroplast DNA (cpDNA) of Heuchera grossulariifolia (Saxifragaceae). Phylogenetic analysis indicated that autopolyploidy has arisen at least twice in the evolutionary history of this species and potentially up to as many as seven times. These results suggest a greater range of independent polyploid origins as compared to a previous study of H. grossulariifolia using cpDNA restriction sites that indicated a minimum of three independent origins. Moreover, most polyploid populations did not contain cpDNA haplotypes from a single origin, but rather combined haplotypes from at least two polyploid origins. Past migration among polyploid populations of independent origin or localized polyploid formation may explain the distribution of polyploid haplotypes within and among populations. The analysis also revealed a discrepancy between relatedness and geographical location. In nearly all sympatric populations of diploids and polyploids, polyploids had the same cpDNA haplotypes as diploids from a geographically remote population. This geographical discordance has several possible explanations, including small sample sizes, extinction of parental diploid haplotypes, chloroplast introgression, and homoplasy in the cpDNA sequence data. We conclude that the recurrent formation of polyploids is an important evolutionary mechanism in the diversification of H. grossulariifolia .     

PLANT POLYPLOIDY AND POLLINATION: FLORAL TRAITS AND INSECT VISITS TO DIPLOID AND TETRAPLOID HEUCHERA GROSSULARIIFOLIA

In many polyploid species, polyploids often have different suites of floral traits and different flowering times than their diploid progenitor species. We hypothesized that such differences in floral traits in polyploids may subsequently affect their interactions with pollinating and other insect visitors. We measured floral morphology and flowering phenology in 14 populations of diploid and autotetraploid Heuchera grossulariifolia Rydb. (Saxifragaceae), determined if repeated evolution of independent polyploid lineages resulted in differentiation in floral morphology among those lineages, and ascertained if there was a consistent pattern of differentiation among genetically similar diploid and autotetraploid populations. In addition, we evaluated the differences in suites of floral visitors within a natural community where diploids and autotetraploids occur sympatrically. Overall, flowers of autotetraploid plants were larger and shaped differently than those of diploids, had a different flowering phenology than that of diploids, and attracted different suites of floral visitors. In comparison with flowers of diploids, tetraploid floral morphology varied widely from pronounced differences between cytotypes in some populations to similar flower shapes and sizes between ploidal levels in other populations. Observations of floral visitors to diploids and autotetraploids in a natural sympatric population demonstrated that the cytotypes had different suites of floral visitors and six of the 15 common visitors preferentially visited one ploidy more frequently. Moreover, we also found that floral morphology differed among independent autotetraploid origins, but there was no consistent pattern of differentiation between genetically similar diploid and autotetraploid populations. Hence, the results suggest that the process of polyploidization creates the potential for attraction of different suites of floral visitors. Multiple origins of polyploidy also presents the opportunity for new or different plant-insect interactions among independent polyploid lineages. These differences in turn may affect patterns of gene flow between diploids and polyploids and also among plants of independent polyploid origin. Polyploidy, therefore, may result in a geographic mosaic of interspecific interactions across a species' range, contributing to diversification in both plant and insect groups.     

On the origin of <Emphasis Type="Italic">Solanum</Emphasis><Emphasis Type="Italic">nigrum</Emphasis>: can networks help?

Black nightshades are a group of species best known for their ‘poisonous’ or noxious weedy reputation. It is not so well known that species of this group serve as emerging food source in many countries worldwide especially in the African continent. Despite the fact that the section has recently been studied extensively, taxonomy is still unsettled and debated because of inter- and intraspecific hybridization, phenotypic plasticity and polyploidization. In this study we analyze the genetic relationships among diploid, tetraploid and hexaploid species of sect. Solanum, which have possibly taken part in the formation of Solanum nigrum, utilizing multi-locus (SCoT, ISSR) markers combined with chloroplast trnL-F sequence data and morphological characters. We scored 51 morphological characters united with SCoT (171), ISSR (224) and trnL-F (1042), for simultaneous analysis of 49 terminals and 1488 characters. The topology of the tree is concordant with the results of the network analysis. In the phylogenetic networks, all the accessions of the diploid species shared a split with all of the polyploid species. This reflected a high portion of shared ISSR and SCoT bands between diploids and polyploids. In addition, a strong split divided the diploid species. The history of S. nigrum might be reticulate with hybrid speciation playing an important rule. Genetically differentiated diploids in few combinations have created a series of genetically distinct polyploid populations. The insufficient isolation that permitted further recombination between ancient polyploids and diploids have resulted in high level of genotypic and phenotypic polymorphism. This high level of novel genomic variability obviously enabled species to succeed in their new environment.     

Origin and evolution of North American polyploid Silene (Caryophyllaceae)

Nuclear DNA sequences from introns of the low-copy nuclear gene family encoding the second largest subunit of RNA polymerases and the ribosomal internal transcribed spacer (ITS) regions, combined with the psbE-petL spacer and the rps16 intron from the chloroplast genome were used to infer origins and phylogenetic relationships of North American polyploid Silene species and their closest relatives. Although the vast majority of North American Silene species are polyploid, which contrasts to the diploid condition dominating in other parts of the world, the phylogenetic analyses rejected a single origin of the North American polyploids. One lineage consists of tetraploid Silene menziesii and its diploid allies. A second lineage, Physolychnis s.l., consists of Arctic, European, Asian, and South American taxa in addition to the majority of the North American polyploids. The hexaploid S. hookeri is derived from an allopolyploidization between these two lineages. The tetraploid S. nivea does not belong to any of these lineages, but is closely related to the European diploid S. baccifera. The poor resolution within Physolychnis s.l. may be attributed to rapid radiation, recombination among homoeologues, homoplasy, or any combination of these factors. No extant diploid donors could be identified in Physolychnis s.l.     

Genomes,multiple origins,and lineage recombination in the Glycine tomentella (Leguminosae) polyploid complex: histone H3-D gene sequences

Relationships among the various diploid and polyploid taxa that comprise Glycine tomentella have been hypothesized from crossing studies, isozyme data, and repeat length variation for the 5S nuclear ribosomal gene loci. However, several key questions have persisted, and detailed phylogenetic evidence from homoeologous nuclear genes has been lacking. The histone H3-D locus is single copy in diploid Glycine species and has been used to elucidate relationships among diploid races of G. tomentella, providing a framework for testing genome origins in the polyploid complex. For all six G. tomentella polyploid races (T1-T6), alleles at two homoeologous histone H3-D loci were isolated and analyzed phylogenetically with alleles from diploid Glycine species, permitting the identification of all of the homoeologous genomes of the complex. Allele networks were constructed to subdivide groups of homoeologous alleles further, and two-locus genotypes were constructed using these allele classes. Results suggest that some races have more than one origin and that interfertility within races has led to lineage recombination. Most alleles in polyploids are identical or closely related to alleles in diploids, suggesting recency of polyploid origins and spread beyond Australia. These features parallel the other component of the Glycine subgenus Glycine polyploid complex, G. tabacina, one of whose races shares a diploid genome with a G. tomentella polyploid race.     

Phylogenetic reconstruction based on low copy DNA sequence data in an allopolyploid: the B genome of wheat.

Study of bread wheat (Triticum aestivum) may help to resolve several questions related to polyploid evolution. One such question regards the possibility that the component genomes of polyploids may themselves be polyphyletic, resulting from hybridization and introgression among different polyploid species sharing a single genome. We used the B genome of wheat as a model system to test hypotheses that bear on the monophyly or polyphyly of the individual constituent genomes. By using aneuploid wheat stocks, combined with PCR-based cloning strategies, we cloned and sequenced two single-copy-DNA sequences from each of the seven chromosomes of the wheat B genome and the homologous sequences from representatives of the five diploid species in section Sitopsis previously suggested as sister groups to the B genome. Phylogenetic comparisons of sequence data suggested that the B genome of wheat underwent a genetic bottleneck and has diverged from the diploid B genome donor. The extent of genetic diversity among the Sitopsis diploids and the failure of any of the Sitopsis species to group with the wheat B genome indicated that these species have also diverged from the ancestral B genome donor. Our results support monophyly of the wheat B genome.     

Molecular phylogeny and reticulate origins of the polyploid Bromus species from section Genea (Poaceae)

The origin of polyploid Bromus species of section Genea was investigated using molecular data. This group of annual species native from the Old-World is composed of three diploids, two tetraploids, one hexaploid, and one octoploid. Molecular cloning, sequencing, and phylogenetic analyses were performed on several accessions per species. We used the low copy nuclear gene Waxy, repeated rDNA spacers ITS1 and ITS2 and chloroplast spacers trnT-trnL and trnL-trnF. Our analyses revealed four different lineages involved in the parentage of the polyploids and confirmed their reticulate origin. Three of these lineages are closely related to the diploid species B. sterilis, B. tectorum, and B. fasciculatus. The fourth lineage could not be related to any diploid according to the available data. Our data gave insights on the origin of all the polyploids of section Genea, and chloroplast data allowed us to identify the maternal lineages. The Waxy gene was the most informative regarding origin of the polyploids. The Waxy copies duplicated by polyploidy appear selectively maintained in the polyploid species. No sequence heterogeneity was encountered in the ITS region, where concerted evolution seems to have occurred toward either maternal or paternal repeats. These results provide new information about the origin and molecular evolution of these polyploids and will allow a more accurate taxonomic treatment of the concerned species, based on their evolutionary history.     

A CYTOGEOGRAPHIC STUDY OF CHAENACTIS DOUGLASII (COMPOSITAE,HELENIEAE)

Analysis of 512 plants derived from 200 populations shows that the widely distributed western North American Chaenactis douglasii species-complex consists of diploids (n = 6), triploids, tetraploids, and hexaploids. Microsporocytes were the source of most of the chromosome counts. About 9% of all plants examined had one or more full-sized extra chromosomes. Multivalents, usually a ring or chain of four chromosomes, were almost entirely restricted to polyploids, where one or more were identified in 38% of the tetraploids and 33% of the hexaploids. With two exceptions, diploids and polyploids were not found in the same population. Frequencies of diploid, triploid, tetraploid, and hexaploid populations were, respectively, 34, 1.5, 55 and 9.5%. With significant exceptions, diploid populations predominate in the Pacific and Rocky Mountain Systems, whereas polyploid ones are most frequent in the intervening plateaus. Ploidy level is correlated with age of substrate, rather than with climate, elevation, vegetation, or soil type. Range, morphology, ploidy level, and meiotic behavior suggest that var. achilleifolia tetraploids and hexaploids are descendents of hybrids between other variants of the complex. The diploid-tetraploid-hexaploid geographic distribution and the age of the substrates where each tends to occur suggest that the complex evolved in late Cenozoic time in response to major climatic and geologic changes that induced migration and hybridization. The hybrid derivatives, stabilized by polyploidy and tolerant of increasing aridity, came to occupy newly available habitats in areas disturbed by volcanic activity and glacial or glacial-related processes.     

Breeding Systems in American and Asian Trillium Species by Means of Chromosome Analyses

Abstract Species of Trillium have a disjunct distribution occurring in both North America and eastern Asia. In North America all 36 species are diploid. The 11 species of eastern Asia, however, include only a single diploid with all the other species being polyploids. Why do different patterns of speciation develop in North America and in eastern Asia? The breeding systems of populations in the North American T. erectum, T. grandiflorum and T. ovatum , and in Asian T. kamtschaticum were investigated by estimating the inbreeding coefficient from cold-induced banding patterns which reveal homozygotes and heterozygotes. From the analyses of the inbreeding coefficients, T. erectum, T. grandiflorum and the Pacific coastal species, T. ovatum are predominantly inbreeding species. T. ovatum populations from the Rocky Mountain region are outbreeders. However the Japanese species, T. kamtschaticum has a mixture of outbreeding and inbreeding among populations. The development of polyploid systems in Asia is possibly the result of the diversity of the breeding systems among the populations. The shift from outbreeding to inbreeding appears to be an important key step in the occurrence of poliploids by hybridization between the different species.     

Evolution and polyploid origins in North American Arctic Puccinellia (Poaceae) based on nuclear ribosomal spacer and chloroplast DNA sequences

The proportion of polyploid plant species increases at higher latitudes, and it has been suggested that original postglacial Arctic immigrants of some large groups, including grasses, were polyploid. We analyzed noncoding nuclear and chloroplast DNA of all North American diploid Puccinellia (Poaceae) and a subset of arctic polyploids to hypothesize evolutionary relationships among diploids and to evaluate the parentage of polyploids. Diploids formed three lineages: one uniting arctic species P. arctica and P. banksiensis; a second comprising arctic species P. tenella, P. alaskana, P. vahliana, and P. wrightii; and a third uniting the two temperate species P. lemmonii and P. parishii. The arctic species P. angustata (hexaploid) and P. andersonii (primarily octoploid) apparently derive from the P. arctica-P. banksiensis lineage based on ITS and chloroplast sequences, and share an ancestor with arctic triploid/tetraploid P. phryganodes based on nrDNA sequences. Sequence comparisons also suggest tetraploid P. bruggemannii evolved from two arctic lineages: P. vahliana-P. wrightii and P. arctica-P. banksiensis. These patterns and the predominance of arctic rather than temperate diploid species support the idea that diploid Puccinellia recolonized the Arctic from northern glacial refugia like Beringia, and also formed stabilized polyploid hybrids during these refugial events or subsequently during postglacial colonization.     

Other tetraploid species and conspecific diploids as sources of genetic variation for an autotetraploid

? Premise of the study: Most plants are polyploid and have more than two copies of the genome. The evolutionary success of polyploids is often attributed to their potential to harbor increased genetic variation, but it is poorly understood how polyploids can attain such variation. Because of their formation bottleneck, newly formed tetraploids start out with little variation. Tetraploids may attain genetic variation through a combination of new mutations, recurrent formation, and gene exchange with diploid ancestors or related tetraploid species. We explore the role of gene exchange and introgression in autotetraploid Rorippa amphibia, a species that harbors more genetic variation than its diploid ancestors. ? Methods: We crossed autotetraploid R. amphibia to diploid conspecifics and tetraploid R. sylvestris and backcrossed resulting F(1) hybrids. We used flow cytometry to determine the ploidy of all progeny. ? Key results: Tetraploids of R. amphibia and R. sylvestris were interfertile; F(1) hybrids were fertile and could backcross. Crosses between diploids and tetraploids yielded a small number of viable, often tetraploid progeny. This indicates that unreduced gametes can facilitate gene flow from diploids to tetraploids. We detected a frequency of unreduced gametes of around 2.7 per 1000, which was comparable between diploids and tetraploids. ? Conclusions: Introgression from tetraploid R. sylvestris provides a realistic source of variation in autotetraploid R. amphibia. Only in a scenario where other compatible partners are absent, for example immediately after tetraploidization, gene flow through unreduced gametes from diploids could be an important source of genetic variation for tetraploids.     

SYSTEMATIC INFERENCES FROM VARIATION IN ISOZYME PROFILES OF ARCTIC AND ALPINE CESPITOSE FESTUCA (POACEAE)

Variation in isozyme patterns was used to assess species boundaries in North American arctic and alpine representatives of the Festuca ovina L. complex. Isozyme profiles, in combination with chromosome number, delimit four discrete entities within the complex: F. brevissima Jurtzev (diploid); F. aggr. auriculata Drobov (diploid); F. brachyphylla Schultes (hexaploid); and tetraploid populations corresponding in morphology to F. baffinensis Polunin (arctic Canada) and F. minutiflora Rydberg (alpine United States). Although no fixed difference was detected between isozyme profiles of the latter two taxa, they are morphologically distinct. Thus variation in isozymes, morphology, and chromosome number delimits five taxa within the F. ovina complex in North America. Some alleles observed in the polyploid taxa were not detected among the diploids, and some observed in F. brachyphylla, the hexaploid taxon, were not detected in either the diploid or the tetraploid species. One possible explanation for these occurrences is that the North American polyploids originated in Eurasia, where many other potential diploid and tetraploid progenitors occur.     

Life history divergence of sympatric diploid and polyploid populations of brine shrimp Artemia parthenogenetica

In order to study how polyploidy affects life history patterns in animals, we have examined sympatric diploid and polyploid brine shrimp (Artemia parthenogenetica) from China, Italy and Spain under laboratory conditions. At optimal temperature and salinity (25°C and 90 ppt), diploids from the three populations had much higher intrinsic rates of increase, higher fecundity, faster developmental rates, and larger brood sizes than their sympatric polyploids. The Chinese and Italian populations were selected for further analysis to determine the life history responses of diploids and polyploids to temperature and salinity changes. Under intermediate and high salinities, Chinese and Italian polyploids produced most of their offspring as dormant cysts while their sympatric diploids produced most of their offspring as nauplii. This relationship is reversed in the Spanish diploid-polyploid complex. For the Chinese population at 25° C, pentaploid clones had higher developmental rates than diploid clones at 35 ppt; at 90 ppt, diploid clones had higher developmental rates than the pentaploids. Italian diploids and tetraploids had different responses to variation in both temperature (25° C and 31° C) and salinity (30 ppt and 180 ppt). Our results demonstrate that relative fitness of the two cytotypes is a function of environmental conditions and that sympatric diploids and polyploids respond differently to environmental changes. Chinese and Italian polyploids are expected to have lower fitness than their sympatric diploids when the physical environment is not stressful and when intraspecific competition is important. However, polyploids may have advantages over sympatric diploids in stressful habitats or when they encounter short-term lethal temperatures. These results suggest that polyploid Artemia have evolved a suite of life-history characteristics adapting them to environments that contrast to those of their sympatric diploids.