Essential oil composition of Bothriochloa Kuntze (Poaceae) from South America and their chemotaxonomy

The essential oils compositions of the 13 taxa (twelve species and two varities) of Bothriochloa have been studied. Two entities are endemics: Bothriochloa eurylemma and Bothriochloa meridionalis. The other nine taxa have disjunt distribution between North and South America and are representatives of two complexes: the Bothriochloa barbinodis complex and the Bothriochloa saccharoides complex. Multivariated statistical analysis (Principal Component Analysis, Hierarchical Cluster Analysis) applied to GC–MS data seems to have systematic significance for the delimitation of the species, independently of their morphological characteristics. However, some affinities with chromosome complement and mode of reproduction (amphimixis or sexuality) can be recognized.     

Reproductive biology of South American Bothriochloa (Poaceae: Andropogoneae)

The reproductive biology of 13 South American taxa of Bothriochloa was studied. All surveyed species reproduce sexually and are self-compatible. Several elements also point to self-pollination as their main breeding system: (i) bud pollination that occurs in most species enhancing cleistogamy, (ii) presence of pits in the glumes that impede the opening of flowers and (iii) reduction in the number of stamens, a trend associated with outbreeding.     

高粱属植物的地理分布

为探讨高粱属(Sorghum Moench)的系统发育关系,通过野外调查及查阅标本和文献资料,对高粱属植物的地理分布进行了整理和研究。高粱属植物约有29种,分布于全世界热带到温带地区,其中澳大利亚22种,亚洲15种,非洲9种,欧洲3种,地中海2种,美洲6种。中国有5种,分布在东北、西南到华南各省(区)。高粱属有5亚属,仅高粱亚属(subgen.Sorghum)延伸至新世界,其他亚属均分布在旧世界,高粱亚属覆盖非洲并扩散到全世界热带到温带地区;拟高粱亚属(subgen.Parasorghum)分布在非洲、亚洲、澳大利亚;有柄高粱亚属(subgen.Stiposorghum)主要分布在澳大利亚,个别种分布到亚洲;多毛高粱亚属(subgen.Chaetosorghum)分布在澳大利亚;异高粱亚属(subgen.Heterosorghum)分布在澳大利亚和亚洲。这表明澳大利亚东北部是高粱属的现代分布中心和多样化中心,非洲东北部和热带亚洲是否是高粱属的起源地尚需确证。     

Allelic variation in the amphitropical disjunct Muhlenbergia torreyi (Poaceae: Muhlenbergiinae)

Muhlenbergia torreyi occurs in the southwestern United States, northern Mexico, and northwestern Argentina. Allozyme data were used to evaluate genetic diversity within and among populations of this amphitropical disjunct species. Electrophoretic examination of 22 putative enzyme loci in 15 populations revealed high levels of genetic variation (P ranging from 64.7 to 82.4;H from 0.527 to 0.757) and high levels of genetic diversity (F ranging from −0.584 to −0.939). All populations possess high levels of heterogeneity (F _1S approaching −1, mean of −0.837) and exhibit lower levels of genetic fixation among populations (F _ST mean of 0.127). A comparison of genetic identity values among populations from North and South America indicates that the genetic variation is slightly greater (I=0.93) in North America than in South America (I=0.96). A total of 51 alleles were shared among all populations, and four unique alleles were detected: two from North American populations and two from South America. It seems likely thatMuhlenbergia torreyi has dispersed to South America recently, because the populations there are less variable. Chromosome numbers, not determined by the authors, of 2n=20, 21 forM. torreyi and 2n=80, 82 forM. arenicola are reported.Muhlenbegia arenicola is first reported from South America.

---

Resumen   Muhlenbergia torreyi habita en el suroeste de Estados Unidos, norte de México, y noreste de Argentina. Mediante el análisis de alozimas se evaluó la diversidad genética dentro y entre poblaciones de esta especie, que es disjunctiva anfitropical. El exámen electroforético de 22 loci putativos enzimáticos en 15 poblaciones, reveló altos niveles de variación genética (P varía de 64.7 a 82.4;H varía de 0.527 a 0.757) y altos niveles de diversidad genética (F varía de −0.584 a −0.939). Todas las poblaciones poseen altos niveles de heterogeneidad dentro de las mismas poblaciones (F _IS cerca −1, media de −0.837) y exhiben bajos niveles de fijación genética entre poblaciones (F _ST media de 0.127). Una comparación de valores de identidad genética entre poblaciones de Norte y Sudamérica indican que la variación genética es un poco mayor (I=0.93) en Norte América que en Sudamérica (I=0.96). Las poblaciones poseían un total de 51 alelos en común y se detectaron cuatro alelos únicos; dos en las poblaciones norteamericanas y dos en las poblaciones sudamericanas. Probablemente,Muhlenbergia torreyi ha sido dispersada recientemente hacia Sudamérica porque esta poblacion es menos variable. El número de cromosomas, no determinado por los autores, de 2n=20, 21 paraM. torreyi and 2n=80, 82 paraM. arenicola son reportados.Muhlenbergia arenicola es reportada por primera vez en Sudamérica.

     

Floral development and the formation of unisexual spikelets in the Andropogoneae (Poaceae)

We investigated spikelet development in four distantly related species of the grass tribe Andropogoneae to determine whether spikelet development and the formation of unisexual florets are uniform throughout the tribe. We studied development in Bothriochloa bladhii, Coelorachis aurita, Heteropogon contortus, and Hyparrhenia hirta, and compared these with Panicum, a member of the sister tribe Paniceae. Many aspects of spikelet development in the species we have studied correlate with what is already known for Tripsacum and maize (both Andropogoneae), despite variation in how unisexual florets are distributed on the plant. The formation of unisexual spikelets is also uniform. All florets initiate both pistil and stamen primordia. In florets destined to be male, cell death occurs in the subepidermal layers of the gynoecium after the formation of a gynoecial ridge. In florets destined to be female, there is no apparent cell death in the stamens, but growth ceases after anther formation. The similarity in spikelet development and the formation of unisexual florets point to a common genetic mechanism for sex determination throughout the Andropogoneae and possibly the entire Panicoideae. Use of a cell death pathway to cause gynoecial abortion may be the basis of one morphological character that defines the subfamily.     

Chromosome number in South American species of Bothriochloa (Poaceae: Andropogoneae) and evolutionary history of the genus

Mitotic chromosome number of 14 taxa of Bothriochloa native to Argentina, Brazil and Uruguay were surveyed. Chromosome numbers of B. eurylemma, B. meridionalis and B. velutina are reported for the first time, with 2n = 6x = 60, and this ploidy level is the most common among the studied taxa. In addition, new cytotypes were found for B. alta (2n = 60), B. barbinodis (2n = 60), B. exaristata (2n = 80), B. laguroides var. torreyana (2n = 80), B. longipaniculata (2n = 60 and 80), B. perforata (2n = 60) and B. springfieldii (2n = 60). These numbers differ from those reported in the literature.     

Breeding system diversification and evolution in American Poa supersect. Homalopoa (Poaceae: Poeae: Poinae)

Background and Aims Poa subgenus Poa supersect. Homalopoa has diversified extensively in the Americas. Over half of the species in the supersection are diclinous; most of these are from the New World, while a few are from South-East Asia. Diclinism in Homalopoa can be divided into three main types: gynomonoecism, gynodioecism and dioecism. Here the sampling of species of New World Homalopoa is expanded to date its origin and diversification in North and South America and examine the evolution and origin of the breeding system diversity.Methods A total of 124 specimens were included in the matrix, of which 89 are species of Poa supersect. Homalopoa sections Acutifoliae, Anthochloa, Brizoides, Dasypoa, Dioicopoa, Dissanthelium, Homalopoa sensu lato (s.l.), Madropoa and Tovarochloa, and the informal Punapoa group. Bayesian and parsimony analyses were conducted on the data sets based on four markers: the nuclear ribosomal internal tanscribed spacer (ITS) and external transcribed spacer (ETS), and plastid trnT-L and trnL-F. Dating analyses were performed on a reduced Poa matrix and enlarged Poaceae outgroup to utilize fossils as calibration points. A relaxed Bayesian molecular clock method was used.Key Results Hermaphroditism appears to be pleisiomorphic in the monophyletic Poa supersect. Homalopoa, which is suggested to have originated in Eurasia 8·4–4·2 million years ago (Mya). The ancestor of Poa supersect. Homalopoa radiated throughout the New World in the Late Miocene–Early Pliocene, with major lineages originating during the Pliocene to Pleistocene (5–2 Mya). Breeding systems are linked to geographic areas, showing an evolutionary pattern associated with different habitats. At least three major pathways from hermaphroditism to diclinism are inferred in New World Homalopoa: two leading to dioecism, one via gynodioecism in South America and another directly from hermaphroditism in North America, a result that needs to be checked with a broader sampling of diclinous species in North America. A third pathway leads from hermaphroditism to gynomonoecism in Andean species of South America, with strictly pistillate species evolving in the highest altitudes.Conclusions Divergence dating provides a temporal context to the evolution of breeding systems in New World Poa supersect. Homalopoa. The results are consistent with the infrageneric classification in part; monophyletic sections are confirmed, it is proposed to reclassify species of sect. Acutifoliae, Dasypoa and Homalopoa s.l. and it is acknowledged that revision of the infrageneric taxonomy of the gynomonoecious species is needed.     

Taxonomy ofHordeum caespitosum,H. jubatum,andH. lechleri (Poaceae: Triticeae)

Hordeum caespitosum Scribner,H. jubatum L., andH. lechleri (Steudel)Schenck are very similar in appearance and therefore until recently were mostly not recognized as separate entities. The first two are tetraploid and natives to North America, but the second occurs naturally in eastern Siberia and has been introduced in Europe and South America and may become a cosmopolitan weed. The third is hexaploid and South American. This study analyses their morphological diversity by means of selected multivariate techniques in order to determine if there is justification to recognize them as three separate morphological species. Logistic discrimination, although based on a reduced set of characters, yielded the highest percent of correct assignments. A linear discriminant function is provided and validated by 100 bootstrap repeats. Canonical discriminant analysis indicated three groups. It is subsequently concluded that the three are separate morphological species. Although a linear discriminant function is given, a traditional identification key is provided based on the palea length and triad (the group of three spikelets at each rachis node) length.     

Natural hybridization in the Andropogon lateralis complex (Andropogoneae,Poaceae) and its impact on taxonomic literature

In north‐eastern Argentina, Paraguay and south‐eastern Brazil, morphologically intermediate plants involving Andropogon lateralis, A. bicornis, A. glaziovii, A. arenarius and A. hypogynus were found. The possibility that they were natural hybrids was tested in two ways: (1) where they were sterile, their morphology was compared with that of the putative parents, and their meiosis and reproductive behaviour were studied; (2) where they were fertile, studies of artificial hybrids were also made. Most of the hybrids were sterile. The only fully fertile combination, generating recombination and hybrid swarms, was A. lateralis × A. hypogynus. In spite of apparently normal chromosome pairing, fertility was low in all other combinations on both the male and female sides. Sterility is probably a result of ‘cryptic’ or ‘gametic sterility’, which produces complete sterility of the gametes. Many of the hybrids survive and compete successfully with the parental species in natural populations, but their sterility maintains the genetic isolation of the majority of the taxa involved. Meiotic chromosome behaviour in all the hybrids indicates that the group of species shares slightly different forms of three basic genomes. Several specimens of natural hybrids were found in historical herbarium collections. In the past, they were given the status of type specimens of at least five taxonomic entities (A. lindmanii, A. coloratus, A. lateralis var. subtilior, A. multiflorus and A. lateralis var. bogotensis). The taxonomic consequences of these findings are discussed. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 136–154.     

Delimitation of the genus Schizachyrium (Poaceae,Andropogoneae) based on molecular and morphological data

Schizachyrium (Poaceae, Andropogoneae) includes about 60 species distributed in tropical and subtropical regions of the world. In all recent molecular phylogenies of Andropogoneae, representatives of Schizachyrium appear closely related to Andropogon species. The objective of this study was to contribute to the delimitation of Schizachyrium. We performed a phylogenetic study including 38 taxa (>63%) of Schizachyrium, along with representatives of related genera, mainly of Andropogon, yielding a total of 49 taxa. This is the first phylogenetic analysis to include the type species of Schizachyrium, S. condensatum. DNA sequences of two plastid markers (ndhF and trnL-F) were analyzed under Bayesian methods. The results indicate that Schizachyrium is not monophyletic: 26 of the 38 Schizachyrium taxa analyzed are placed in a Schizachyrium s.s. clade that includes the type species of the genus, while 10 taxa are related to Andropogon species and two other species, S. delavayi (from China and India) and S. jeffreysii (from Africa), appear clearly separated. Additionally, 58 morphological characters (41 qualitative and 17 quantitative) were scored for the same 49 taxa and analyzed under the parsimony criterion. Character optimizations showed that (i) the reduced pedicellate spikelets, (ii) with lower glume less than or equal to 0.5 mm wide, (iii) awned, and (iv) without lemma and palea support the Schizachyrium s.s. clade. We propose these four characters as diagnostic features for the delimitation of Schizachyrium s.s.     

Chromosome number and ploidy level of some South American species of Schizachyrium (Poaceae, Andropogoneae)

Chromosome numbers are reported here for 53 accessions belonging to 13 taxa of the genus Schizachyrium. The chromosome counts for the following seven species are recorded here for the first time: S. bimucronatum (2n?=?20), S. gracilipes (2n?=?20), S. hatschbachii (2n?=?20), S. lactiflorum (2n?=?20), S. plumigerum (2n?=?20), S. salzmannii (2n?=?20; 2n?=?40) and S. spicatum (2n?=?20). Counts differing from those previously reported are given for two species: S. scabriflorum (2n?=?20 and 80) and S. tenerum (2n?=?40). Most of the analyzed species of this genus are diploids. The occurrence of polyploidy in S. glaziovii, S. lactiflorum, S. salzmanni, S. scabriflorum and S. tenerum is reported here for the first time. Data on chromosome numbers and ploidy levels are discussed in relation to the taxonomy and evolution of the genus.     

Allozyme variation within and between populations inLolium (Poaceae)

Isozyme analysis was used to determine genetic variation within and between populations of sevenLolium species. All populations from the inbreeding species (L. temulentum, L. remotum, L. loliaceum, andL. persicum) were completely fixed for all enzymes scored. They also contained, for four of the five enzyme systems studied, exactly the same allelic variant. The three cross-breeding species showed large within-population variation and much less between-population variation. The great similarity of the allozymic variants found in all species, made the division of the genusLolium into species on basis of allozymic data difficult. It was not possible to separate the different inbreeding species from each other. Within the cross-breeding groupL. multiflorum andL. rigidum could be distinguished fromL. perenne. L. multiflorum, andL. rigidum could, with more difficulty, also be separated from each other. Allelic variation could have more relation with the provenance of the populations than with taxonomic classification.Genetic variation inLolium spp. II. For first part see Pl. Syst. Evol. 188: 87–99.     

The phylogeny of the genus Ischioscia Verhoeff, 1928, with redescriptions of three species (Crustacea: Isopoda: Oniscidea)

The genus Ischioscia Verhoeff, 1928 is reviewed. 26 species are considered valid. A key for their identification is given, as well as a map showing the geographic distribution. The known range of the genus covers a large area from the central Amazon region to the mountains of Guatemala. The species of Ischioscia have a typical “philosciid” habitus (“runner” type); they can be distinguished from other Neotropical species with similar habitus by the following apomorphies: (1) male pereiopod 1 carpus enlarged to a plate-like extension, (2) scale field on male pereiopod 1 covering entire frontal side of the carpus, (3) male pereiopod 7 ischium with a ventral scale field, (4) dactylus in both sexes with a long inner claw. The groundpattern of Ischioscia is reconstructed, and an analysis of the phylogenetic relations within the genus is made on the basis of morphological data. The species are very similar to each other, most differences are found in the male structures of sexually dimorphic features. Ischioscia sturmi (Vandel, 1972), I. amazonica Lemos de Castro, 1955 and I. bolivari Vandel, 1968 are redescribed in detail.     

Genome analysis ofElymus angulatus andE. patagonicus (Poaceae) and their hybrids with N. and S. AmericanHordeum spp.

The results of genome analysis of five hybrids, viz.Elymus patagonicus ×Hordeum procerum, E. patagonicus ×H. tetraploidum, E. angulatus ×H. jubatum, E. angulatus ×H. lechleri, andE. angulatus ×H. parodii, are reported. The genomic constitution ofHordeum tetraploidum andH. jubatum is best given as H₁H₁H₂H₂, ofH. lechleri andH. parodii as H₁H₁H₂H₂H₄H₄, ofH. procerum as H₁H₁H₂H₂H₃H₃, and ofElymus patagonicus andE. angulatus as SSH₁H₁H₂H₂.     

Morphological variation inLolium (Poaceae) as a measure of species relationships

Analysis of morphological and phenological data for determining the genetic variation within sevenLolium species led to the recognition of two groups within this genus. One group, containing the two inbreeding speciesL. temulentum andL. persicum, was clearly distinct from all other species. Strong morphological and phenological intergradation was found between both species. The cross-breeding species,L. perenne, L. rigidum, andL. multiflorum, formed another group. Little differentiation was found between these species, though they were distinct. Two inbreeding species,L. loliaceum andL. remotum, were clearly distinct from each other and the two groups.L. loliaceum had an isolated position and was most related toL. rigidum. L. remotum had an intermediate position between the cross-breeding and inbreeding species, and was almost equally distant from all three cross-breeding species.Genetic variation inLolium spp. I.     

Hybridisation,genomic constitution and generic delimitation inElymus s. l. (Poaceae: Triticeae)

Intergeneric crosses were made between representatives of the genomically-defined generaElymus, Agropyron, Elytrigia, Pseudoroegneria, andThinopyrum. The genomic constitution ofElytrigia repens, the type species ofElytrigia, is shown to be SSH, a genomic combination otherwise found only inElymus. The S genome ofPseudoroegneria has almost always a dominant influence on the morphology of the taxa of which it is a component.Wang (1989) showed that the J genome inThinopyrum and the S genome have considerable homoeology, with a mean c-value of 0.35 in diploid SJ hybrids. A genetic coherence from S to SJ^e, J^e, J^eJ^b, and J^b can be expected, agreeing with the continuous morphologic variation pattern observed. Because of the absence of morphological discontinuities between the taxa,Pseudoroegneria (S),Elymus (SH, SY, sometimes with additional genomes),Elytrigia (SSH, SSHX), andThinopyrum (SJ, SJJ, J) are best treated as a single genus,Elymus, following the generic concept ofMelderis in Flora Europaea and Flora of Turkey. The basic genomic constituents ofElymus will then be the S and/or J genomes.Agropyron, with diploids, tetraploids, and hexaploids based on the P genome is morphologically distinct from other genera inTriticeae. In a few species ofElymus andPseudoroegneria, a P genome is an additional constituent. In these cases the P genome has a negligible morphological influence. Therefore, it seems reasonable to maintainAgropyron as a separate genus.     

Differentiation of fourHordeum sect.Stenostachys spp. (Poaceae: Triticeae) using multivariate morphometrics

Field collections and 296 herbarium sheets were examined for 27 morphometric variables. A priori species identifcation was based on geographical distribution except forH. californicum, a diploid species primarily occurring in California and differing from the much more widespread tetraploidH. brachyantherum that thrives in N. America and N.E. Asia;H. capense grows in S. Africa andH. secalinum mainly in Europe. Various cluster analyses were used followed by cluster recovery verification. Classificatory discriminant analysis and validation by the bootstrap yielded 85–90% overall total correct classification of the four species. Canonical analysis revealed thatH. californicum occupies an intermediate phenetic position among the other three distinct species. Factors of shape differences were unravelled and portrayed by shearing. A revised key to species was drawn up.