A biotechnology strategy for medium- and long-term conservation of cycads

It has been possible to regenerate a few cycad species in vitro by somatic embryogenesis, either from zygotic embryos (Ceratozamia hildae, C. mexicana, Encephalartos cycadifolius,E. dyerianus, E. natalensis, Zamia fischeri, Z. furfuracea, andZ. pumila) or from leaves of mature phase trees (C. euryphyllidia, Ceratozamia hildae, andC. mexicana). This strategy has great potential for the commercial vegetative propagation of certain highly endangered species (e.g.,C. euryphyllidia) and should indirectly protect wild populations of these species by discouraging collection in situ. Embryogenic cultures of several cycad species have grown vigorously and are highly morphogenic more than 11 years after induction. The long-term conservation of cycad genetic resources can also be addressed for species that can be regenerated by somatic embryogenesis. Preliminary studies indicate that embryogenic cultures that have been pretreated on plant growth medium containing 0.75 M sucrose for two days, encapsulated in sodium alginate, and desiccated for six hours can survive immersion in liquid nitrogen (−196°C).     

INSECT POLLINATION IN THE CYCAD ZAMIA PUMILA (ZAMIACEAE)

A wind and insect exclusion pollination experiment was conducted in a wild population of the cycad Zamia pumila L. in Florida. Cones from which insects but not wind were excluded produced no viable seeds, while cones from which wind but not insects were excluded produced abundant viable seeds. Two beetle species have been identified which may be effecting pollination. Adults and larvae of both beetles are found abundantly on the male cones, and adults of both species and larvae of one species are found occasionally on female cones. Z. pumila produces sugar and amino acid-rich micropyle droplets which may serve as pollinator rewards. Mimicry of resource-rich male cones by female cones and the use of cones as refuges between mating and feeding bouts may account for beetle movement from male to female cones.     

Comparison of genetic population structure between two cyprinids, <Emphasis Type="Italic">Hemigrammocypris rasborella</Emphasis> and <Emphasis Type="Italic">Pseudorasbora pumila</Emphasis> subsp., in the Ise Bay basin,central Honshu,Japan

Two small cyprinid fishes, Hemigrammocypris rasborella and Pseudorasbora pumila subsp. (sensu Nakamura 1963), inhabit similar habitats and often occur sympatrically in the Ise Bay basin, central Honshu Island, Japan. Their genetic population structures were revealed, using sequence data from the mitochondrial cytochrome b gene, and then compared. Hemigrammocypris rasborella populations in the Ise Bay area formed a monophyletic group that has been isolated from eastern (Tenryu River) and western (Lake Biwa–Yodo River) populations at least for several hundred thousand years. Pseudorasbora pumila subsp., endemic to the Ise Bay area, was estimated to have become isolated from its sister subspecies, P. p. pumila, about 5 million years ago. Both H. rasborella and P. pumila subsp. had centers of genetic diversity around the Okazaki Plain in the eastern part of the basin and showed trans-bay distribution of haplotypes or haplotype groups. Their common population structure was explained by geological features in the Ise Bay area, in which a large paleo-river system developed in regression periods, suggesting gene flow among populations of each species in the mid to lower reaches of the paleo-river. Based on the estimated expansion or divergence time, however, not all populations experienced gene flow during the Last Glacial. In contrast to the maintenance of high genetic diversity in H. rasborella, almost all populations of P. pumila subsp. have lost mitochondrial DNA genetic diversity. This implies that effective population size of P. pumila subsp. tended to be smaller, probably because of differences in reproductive ecology, even though the two species have been exposed to similar environmental changes. For conservation of the two species, genetic and adaptive differentiation among local populations should be considered, and attention should be paid to inbreeding depression, especially in P. pumila subsp. An erratum to this article can be found at     

Somatic embryogenesis and organogenesis in Zamia fischeri,Z. furfuracea and Z. pumila

Morphogenesis from cultured megagametophyte, nucellus and immature embryos of three Zamia species on modified B5 medium containing 2,4-d and kinetin was compared. Organogenesis and somatic embryogenesis occurred from the megagamethophyte and zygotic embryo explants of Zamia pumila and Z. furfuracea tissue cultures, but only from the megamametophyte of Z. fischeri. Nucellar callus of Z. pumila produced globular structures that failed to develop further. Plantlets were recovered from somatic embryos of Z. pumila.Abbreviations 2,4-d 2,4-dichlorophenoxyacetic acid     

When North and South don’t mix: genetic connectivity of a recently endangered oceanic cycad,Cycas micronesica,in Guam using EST‐microsatellites

Subject to environmental changes and recurrent isolation in the last ca. 250 Ma, cycads are often described as relicts of a previously common lineage, with populations characterized by low genetic variation and restricted gene flow. We found that on the island of Guam, the endemic Cycas micronesica has most of the genetic variation of 14 EST‐microsatellites distributed within each of 18 genetic populations, from 24 original sampling sites. There were high levels of genetic variation in terms of total number of alleles and private alleles, and moderate levels of inbreeding. Restricted but ongoing gene flow among populations within Guam reveals a genetic mosaic, probably more typical of cycads than previously assumed. Contiguous cycad populations in the north of Guam had higher self‐recruitment rates compared to fragmented populations in the south, with no substantial connection between them except for one population. Guam’s genetic mosaic may be explained by the influence of forest continuity, seed size, edaphic differences, and human transport of cycads. Also important are the extent of synchrony among flushes of reproductive female seed‐bearing sporophylls and restricted pollen movement by an obligate mutualist and generalist insects. An NADH EST‐locus under positive selection may reflect pressure from edaphic differences across Guam. This and three other loci are ideal candidates for ecological genomic studies. Given this species’ vulnerability due to the recent introduction of the cycad aulacaspis scale, we also identify priority populations for ex situ conservation, and provide a genetic baseline for understanding the effects of invasive species on cycads in the Western Pacific, and islands in general.     

Conserved genetic regions across angiosperms as tools to develop single‐copy nuclear markers in gymnosperms: an example using cycads

Several individuals of the Caribbean Zamia clade and other cycad genera were used to identify single‐copy nuclear genes for phylogeographic and phylogenetic studies in Cycadales. Two strategies were employed to select target loci: (i) a tblastX search of Arabidopsis conserved ortholog sequence (COS) set and (ii) a tblastX search of Arabidopsis‐Populus‐Vitis‐Oryza Shared Single‐Copy genes (APVO SSC) against the EST Zamia databases in GenBank. From the first strategy, 30 loci were selected, and from the second, 16 loci. In both cases, the matching GenBank accessions of Zamia were used as a query for retrieving highly similar sequences from Cycas, Picea, Pinus species or Ginkgo biloba. After retrieving and aligning all the sequences in each locus, intron predictions were completed to assist in primer design. PCR was carried out in three rounds to detect paralogous loci. A total of 29 loci were successfully amplified as a single band of which 20 were likely single‐copy loci. These loci showed different diversity and divergence levels. A preliminary screening allowed us to select 8 promising loci (40S, ATG2, BG, GroES, GTP, LiSH, PEX4 and TR) for the Zamia pumila complex and 4 loci (COS26, GroES, GTP and HTS) for all other cycad genera.     

Hybrid speciation in sparrows II: a role for sex chromosomes?

Homoploid hybrid speciation in animals is poorly understood, mainly because of the scarcity of well‐documented cases. Here, we present the results of a multilocus sequence analysis on the house sparrow (Passer domesticus), Spanish sparrow (P. hispaniolensis) and their proposed hybrid descendant, the Italian sparrow (P. italiae). The Italian sparrow is shown to be genetically intermediate between the house sparrow and Spanish sparrow, exhibiting genealogical discordance and a mosaic pattern of alleles derived from either of the putative parental species. The average variation on the Z chromosome was significantly reduced compared with autosomal variation in the putative parental species, the house sparrow and Spanish sparrow. Additionally, divergence between the two species was elevated on the Z chromosome relative to the autosomes. This pattern of variation and divergence is consistent with reduced introgression of Z‐linked genes and/or a faster‐Z effect (increased rate of adaptive divergence on the Z). F_ST‐outlier tests were consistent with the faster‐Z hypothesis: two of five Z‐linked loci (CHD1Z and PLAA) were identified as candidates for being subject to positive, divergent selection in the putative parental species. Interestingly, the two latter genes showed a mosaic pattern in the (hybrid) Italian sparrow; that is, the Italian sparrow was found to be fixed for Spanish sparrow alleles at CHD1Z and to mainly have house sparrow alleles at PLAA. Preliminary evidence presented in this study thus suggests that sex chromosomes may play a significant role in this case of homoploid hybrid speciation.     

VRN1 genes variability in tetraploid wheat species with a spring growth habit

Background

Vernalization genes VRN1 play a major role in the transition from vegetative to reproductive growth in wheat. In di-, tetra- and hexaploid wheats the presence of a dominant allele of at least one VRN1 gene homologue (Vrn-A1, Vrn-B1, Vrn-G1 or Vrn-D1) determines the spring growth habit. Allelic variation between the Vrn-1 and vrn-1 alleles relies on mutations in the promoter region or the first intron. The origin and variability of the dominant VRN1 alleles, determining the spring growth habit in tetraploid wheat species have been poorly studied.

Results

Here we analyzed the growth habit of 228 tetraploid wheat species accessions and 25 % of them were spring type. We analyzed the promoter and first intron regions of VRN1 genes in 57 spring accessions of tetraploid wheats. The spring growth habit of most studied spring accessions was determined by previously identified dominant alleles of VRN1 genes. Genetic experiments proof the dominant inheritance of Vrn-A1d allele which was widely distributed across the accessions of Triticum dicoccoides. Two novel alleles were discovered and designated as Vrn-A1b.7 and Vrn-B1dic. Vrn-A1b.7 had deletions of 20 bp located 137 bp upstream of the start codon and mutations within the VRN-box when compared to the recessive allele of vrn-A1. So far the Vrn-A1d allele was identified only in spring accessions of the T. dicoccoides and T. turgidum species. Vrn-B1dic was identified in T. dicoccoides IG46225 and had 11 % sequence dissimilarity in comparison to the promoter of vrn-B1. The presence of Vrn-A1b.7 and Vrn-B1dic alleles is a predicted cause of the spring growth habit of studied accessions of tetraploid species. Three spring accessions T. aethiopicum K-19059, T. turanicum K-31693 and T. turgidum cv. Blancal possess recessive alleles of both VRN-A1 and VRN-B1 genes. Further investigations are required to determine the source of spring growth habit of these accessions.

Conclusions

New allelic variants of the VRN-A1 and VRN-B1 genes were identified in spring accessions of tetraploid wheats. The origin and evolution of VRN-A1 alleles in di- and tetraploid wheat species was discussed.

     

Geographic distribution and multilocus organization of isozyme variation of rice (Oryza sativa L.)

Genetic organization of isozyme variation in rice (Oryza sativa L.) was investigated based on 17 polymorphic isozyme loci using a sample of 511 accessions of worldwide origin. The genetic diversity within the species was very high (H=0.36 with 4.82 alleles per locus), as compared with most selfing plant species. Three diversity centers were detected for isozyme variation including South Asia, China and Southeast Asia. The accessions were classified into three well-differentiated cultivar groups corresponding to the indica and japonica subspecies, and a new unnamed group. Variation within the cultivar groups accounted for 80% of the total isozyme variation. Within-country variation accounted for 58% of the total variation while among-region and among-country variation within the cultivar groups accounted for only 14% and 8% of the total variation. Analyses using log-linear models revealed that pronounced non-random associations between and among alleles at many unlinked isozyme loci were organized in a non-hierarchical pattern, and subspecific and macro-geographic differentiation was much more pronounced in multilocus phenotype frequencies than in allelic frequencies at individual loci. These results suggest that selection on multilocus gene complexes was largely responsible for the maintenance of the extensive isozyme variation within the species and the indica-japonica differentiation. Our results further suggest the independent domestication of indica and japonica, the dual origins of the indica rice from China and South Asia (India), and the differentiation of the ecotypes ’javanica’ and the ’temperate japonica’ within the japonica subspecies. Received: 5 August 1999 / Accepted: 13 December 1999     

Mast-seeding in the cycad genus Encephalartos: a test of the predator satiation hypothesis

Mast-seeding behaviour was monitored in 18 populations of eight species of the African cycad genus Encephalartos between 1988 and 1991. The coefficient of variation (V) in annual cone production for each population ranged between 88 and 200, indicating large fluctuations in reproductive effort between years. Data were collected to determine whether mast-seeding reduced levels of predispersal seed predation by satiating seed predators in mast years and whether it resulted in a reproductive advantage over plants which reproduced more frequently. Masting intensity was greatest in those populations in which individual plants suffered the highest levels of predispersal seed predation in years when only a few plants produced seeds. The principal seed predators were two congeneric weevil species, Antliarhinus zamiae and A. signatus, which develop exclusively on cycad seeds. The lowest intensity of mast-seeding was recorded for cycad populations with low levels of seed predation and in which A. zamiae and A. signatus occurred only in low numbers or were entirely absent. Larger seed crops appeared to result in lower levels of seed predation by A. zamiae and A. signatus in four populations of E. altensteinii, and differences in seed crop size accounted for 48–66% of variation in levels of seed predation in populations of five cycad species. In one population of E. altensteinii, lower levels of seed predation in plants reproducing periodically resulted in a reproductive advantage over plants reproducing more frequently. These results are consistent with the predator satiation hypothesis. However, in most cycad populations, numbers of seed predators did not appear to decrease significantly after a period of 2–8 years between reproductive episodes and, in two of three populations examined, periodic reproduction did not increase the number of seeds surviving to dispersal over a 4-year period. These results are interpreted to mean that periodic reproduction has not evolved in response to selection imposed by seed predators, but that selection may favour those plants which experience lower levels of seed predation by coning in synchrony with the majority of plants in the population.     

Genetic structure of Glycine canescens,a perennial relative of soybean

Summary Allozyme variation as detected by starch gel electrophoresis was used to assess the extent and spatial organization of genetic variation across the entire range of Glycine canescens sensu lato. Eleven enzyme systems were assayed in 116 accessions of this taxon and 102 alleles were detected at a total of 31 loci. Eighty-one percent of loci were polymorphic. Most of this variation occurred between and very little within accessions. Three major groupings were detected. These groupings (groups 1, 2, and 3) also differed with respect to mean seed size and their geographic distribution. A further ten accessions stood out from these distinct groups. These accessions were most closely related to group 3 but were variable among themselves. In general, they were collected from highly dissected terrain, often in the remote interior of the continent. A final group of 18 problematic accessions (group X), originally tentatively identified as G. canescens on morphological grounds, was shown to be isozymically distinct from this species and was reclassified as one form of the polytypic species G. clandestina.     

ALLOZYME VARIATION IN OLD WORLD RACES OF SORGHUM BICOLOR (POACEAE)

A survey of allozyme variation in cultivated races of sorghum (Sorghum bicolor) was undertaken. Eight plants each of 83 accessions representing the five primary races and five of the intermediate races of sorghum were analyzed for 15 enzyme systems encoded by 27 loci. Low levels of variation were found within accessions, which is typical of self-pollinating species. Little variation was also found among accessions. Compared with other cereals, S. bicolor is depauperate in allozyme variation. We found an average of 1.81 alleles per locus with a mean expected heterozygosity of 0.008 for the accessions and total panmictic heterozygosity of 0.093. Only 9% of the variation present was found within accessions, while 91% was among accessions. Most of the variation present is attributable to differences in geographic origin of the accessions rather than racial differences. Western and eastern Africa have the highest levels of total heterozygosity (0.108 and 0.088, respectively), while southern Africa has the lowest (0.008). Principal component analysis revealed continuous variation among races and geographic regions with the accessions failing to segregate into discrete racial or geographic clusters.     

GENETIC DIVERSITY,INTROGRESSION, AND INDEPENDENT DOMESTICATION OF OLD WORLD CULTIVATED COTTONS

Gossypium arboreum L. and G. herbaceum L. are the diploid species of cultivated cotton. Little is known regarding the time and place of domestication of either species. Because G. arboreum is known only as a cultigen, others have proposed that it arose from domesticated G. herbaceum during the more than 5,000-year history of Old World cotton cultivation, with wild G. herbaceum subsp. africanum (Watt) Mauer as the putative ancestor of both species. An alternative hypothesis is that the two species have independent origins from progenitors that diverged prior to domestication. The relative merits of these opposing hypotheses were evaluated using data derived from starch gel electrophoresis of enzymes. One hundred and three accessions of G. arboreum and 31 accessions of G. herbaceum were examined for allelic variation at 40 allozyme loci. All measures of genetic variability demonstrated that G. arboreum contains greater diversity than G. herbaceum, although both species have relatively low levels of allozyme variation. In contrast to expectations based on morphology and other chemical data sets, the two species are highly differentiated with respect to allozyme composition. Gossypium arboreum and G. herbaceum each contain a large number of unique alleles and are fixed or nearly fixed for alternate alleles at 8 loci. Five allozyme loci have alleles that are rare in one of the two species but common in the other. Based on restricted occurrence of these alleles to areas with a long history of sympatric cultivation and the geographic distribution of a null allele, we suggest that a significant portion of the allelic diversity in both species results from historical, bidirectional interspecific introgression. The interspecific genetic identity estimate (0.74) is markedly lower than for documented progenitor-derivative and crop-ancestor species pairs. Based on these data, as well as previous cytogenetic data and the observation of F₂ breakdown in interspecific crosses, we suggest that cultivated G. arboreum and G. herbaceum were independently domesticated from divergent ancestors.     

Very close relationship of the chloroplast genomes among Saccharum species

We recently determined the complete sequence of the sugarcane chloroplast genome. Here, we have used the information for a comprehensive phylogenetic analysis of the genus Saccharum, using all six species (13 accessions). The polymorphisms between sugarcane and maize in 26 chloroplast genome regions were used for the analysis. In 18 of the 26 regions (a total of 5,381 bp), we found 41 mutations involving 17 substitutions, three inversions, six insertion/deletion mutations, and 15 simple sequence repeat length polymorphisms. Based on these results, we calculated a phylogenetic tree of the genus Saccharum, in which all six species are clearly separated. By the analysis, (1) S. sinense and S. barberi, which have identical sequences, belong to the same clade, whereas the other four species, S. officinarum, S. robustum, S. edule, and S. spontaneum, form an independent clade; (2) S. spontaneum has a paraphyletic relationship with the other five species; and (3) no or very low intraspecific variation was observed in S. officinarum, S. robustum, S. sinense, S. barberi, and S. edule, whereas higher intraspecific variation was observed in S. spontaneum. Based on the number of nucleotide substitutions, the divergence time between S. officinarum and S. spontaneum, and between S. officinarum and maize were calculated to be about 730–780 thousand years ago and about 5.9 million years ago, respectively. These results suggest that the cytoplasm of Saccharum species are very closely related.     

Comparison of DNA restriction fragment length polymorphisms of Nostoc strains in and from cycads

DNA was prepared from cyanobacteria freshly isolated from coralloid roots of natural populations of five cycad species: Ceratozamia mexicana mexicana (Mexico), C. mexicana robusta (Mexico), Dioon spinulosum (Mexico), Zamia furfuraceae (Mexico) and Z. skinneri (Costa Rica). Using the Southern blot technique and cloned Anabaena PCC 7120 nifK and glnA genes as probes, restriction fragment length polymorphisms of these cyanobacterial symbionts were compared. The five cyanobacterial preparations showed differences in the sizes of their DNA fragments hybridizing with both probes, indicating that different cyanobacterial species and/or strains were in the symbiotic associations. On the other hand, a similar comparison of cyanobacteria freshly collected from a single Encephalartos altensteinii coralloid root and from three independently subcultured isolates from the same coralloid root revealed that these were likely to be one and the same organism. Moreover, the complexity of restriction patterns shows that a mixture of Nostoc strains can associate with a single cycad species although a single cyanobacterial strain can predominate in the root of a single cycad plant. Thus, a wide range of Nostoc strains appear to associate with the coralloid roots of cycads.Non-standard abbreviations bp base pairs - kbp kilobase pairs - RFLP's restriction fragment length polymorphisms     

Temporal changes in genetic diversity of common bean (Phaseolus vulgaris L.) accessions cultivated between 1800 and 2000

Fourteen microsatellite markers were used to describe genetic diversity in a sample of 128 common bean (Phaseolus vulgaris L.) accessions cultivated within the territory of Slovenia and its nearby regions between 1800 and 2000. The accessions were grouped into three periods: period I comprising accessions from the beginning of the 19th century, while the other two periods included accessions from the middle (period II) and the end of the 20th century (period III). Seven control accessions of known Mesoamerican and Andean origin were also included in the study. A total of 130 alleles were generated. Allelic richness, in terms of number of alleles per locus, was 6.07 for period I, 6.71 for period II, and 6.07 for period III. In the UPGMA dendrogram, all studied accessions were intermixed in three main clusters, indicating that the diversity in the time periods overlapped. Two clusters consisted of accessions of Andean and Mesoamerican origin, while the third represents additional variation, which existed in this area already 200 years ago. The analysis of molecular variance showed that a great part of genetic diversity has been preserved till today, confirming the results of cluster analysis. The calculation of number of alleles per locus revealed no significant quantitative change in genetic diversity over the last 200 years of common bean cultivation. However, the calculation of genetic distances indicated slight qualitative shifts in genetic diversity of common bean germplasm over time, while the calculations of allelic frequency variation and polymorphic information content revealed recent decline of some alleles’ frequencies. These findings should stress the need for establishing an appropriate strategy of genetic resources management. The text was submitted by the authors in English.     

Clonal diversity and genetic differentiation revealed by SSR markers in wild Vaccinium macrocarpon and Vaccinium oxycoccos

American cranberry (Vaccinium macrocarpon) is a perennial, woody plant species, native to North American bogs and wetlands. Cranberries represent one of the few agriculturally important native plants in which wild gene pools are still readily available within the undeveloped wetlands of the northern US and Canada. Earlier studies have reported low genetic variation in V. macrocarpon at the species and population level. However, in this study, we characterised 229 individuals of wild V. macrocarpon and V. oxycoccos (small cranberry) from Wisconsin and 22 accessions using microsatellite markers and observed substantial genetic variation and differentiation within and among populations and species. While V. macrocarpon was analysed using 108 alleles from 11 microsatellite loci revealing 42 unique genotypes, V. oxycoccos was analysed using 156 alleles from eight loci revealing 28 unique genotypes. There were a total of 182 alleles found in both species combined with 156 of those alleles present in V. oxycoccos and 84 alleles found in V. macrocarpon. All eight loci possessed species‐specific alleles with V. oxycoccos possessing 98 private alleles versus 26 private alleles found V. macrocarpon, and 58 alleles were found in common between both species. Our data will be valuable not only for future wild cranberry diversity and population genetics research, but for other cranberry breeding and genetics studies.     

GENETIC DIVERGENCE IN GALVEZIA (SCROPHULARIACEAE): EVOLUTIONARY AND BIOGEOGRAPHIC RELATIONSHIPS AMONG SOUTH AMERICAN AND GALÁPAGOS SPECIES

Nineteen populations representing all species of Galvezia were examined for electrophoretic differentiation at 23 loci. Allozyme variation within G. leucantha is minimal, suggesting that a genetic bottleneck, possibly resulting from a single colonization event, accompanied establishment of Galvezia in the Galàpagos Islands less than 1.5 million years ago. South American species have higher levels of genetic polymorphism. Interspecific divergence patterns in Galvezia are characterized by many qualitative allelic differences, including one to six marker alleles for each species. Isozyme data support delimitation of four species. Geographic modes of speciation predominate. Allozyme data indicate that the Galàpagos endemic is not a recent derivative from a continental progenitor; rather, it is most likely the sister species to G. fruticosa or to a lineage of G. fruticosa and an undescribed species from southern Peru. It is hypothesized that the presence of six fixed, unique alleles in G. leucantha identifies a relictual genotype from an extinct South American progenitor. Biogeographic patterns and the presence of an allele shared between G. leucantha and two populations from the Sechura Desert indicate that northern Peru is the probable source region for the Galàpagos introduction. Estimates of elapsed divergence times and paleoecological data suggest that continental species diverged in the late Pliocene or during the Pleistocene as a result of geographic isolation caused by climatic and vegetational change.     

Transfer of Pterosiphonia pumila Yendo to the genus Symphyocladia (Rhodomelaceae, Rhodophyta)

The red alga Pterosiphonia pumila Yendo (Rhodomelaceae, Ceramiales) has flattened thalli in which there is complete lateral fusion between branches and their parental axes during early stages of development, the laterals only becoming distally free in reproduc-tively mature plants. This results in a leaf-like organization distinct from that displayed by all other species of the genus Pterosiphonia, the members of which have terete to compressed axes that are congenitally fused along only a few (<4.5) proximal segments of the laterals. The organization of P. pumila is, thus, similar to that of species of the genus Symphyocladia, to which it is here transferred as Symphyoctadia pumila (Yendo) Uwai et Masuda, comb. nov. Symphyoctadia pumila is restricted to Japan and Korea and is distinguished from the other three described species of Symphyocladia by its small (<3 cm long) ecorticate thalli and complete lack of vegetative trichoblasts. Symphyoctadia pennata Okamura has a similar suite of characteristics, has been successfully cross-bred with S. pumila in laboratory culture and is, thus, placed in synonymy with S. pumila.     

Allozyme variation among the spontaneous species of Sorghum section Sorghum (Poaceae)

Summary A survey of allozyme variation among the spontaneous taxa of Sorghum section Sorghum was undertaken. Eight plants each of 90 accessions representing the diploid S. bicolor (ssp. arundinaceum and drummondii) and the tetraploids S. almum and S. halepense were analyzed for 17 enzyme systems encoded by 30 loci. Low levels of variation were found within and among accessions, although there was more variation than is typical of inbreeding species. We found an average of 3.2 alleles per locus in ssp. arundinaceum, with a mean expected heterozygosity for the accessions of 0.034 and total panmictic heterozygosity of 0.154. An analysis of the apportionment of genetic variation among accessions of ssp. arundinaceum indicated that 26% of the variation occurs within accessions and 74% among accessions. Cultivated sorghum contains far less allozymic variation than ssp. arundinaceum, its presumed progenitor. This is consistent with the prediction that cultivated sorghum experienced a loss of genetic variation during domestication. For the most part, cultivated sorghum contains a subset of the allozymes found in ssp. arundinaceum. Principal component analysis revealed continuous variation among the accessions and geographic regions, with accessions failing to segregate into discrete clusters. However, accessions of race virgatum of ssp. arundinaceum occupied one end of the continuum and were, in that sense, distinguished from the other accessions. Similarly, most accessions of S. halepense and S. almum occupied the central portion of the continuum. The allozymic data presented here are consistent with the hypothesized origin of S. halepense via autopolyploidy or segmental allopolyploidy.