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.

     

Early embryogenesis in Tropaeolum majus L.: Diversification of plastids

Embryogeny in the nasturtium is characterized by the development of a large, tripartite suspensor and storing cotyledons. A light and electron microscopic study revealed an early diversification of the plastids in the various regions of the suspensor and the embryo proper. Amyloplasts are found in the developing cotyledons of the heart-like embryo, while chloroplasts occur within the meristematic part of the embryo and the adjacent portion of the suspensor. The cells between the meristem and the storing cotyledons display undifferentiated leukoplasts, whereas leukoplasts with an electron-dense matrix occur in the basal cell mass of the embryo-suspensor. Etioplasts develop in several cells of the placental haustorium of the suspensor. The carpel haustorium shows rather undifferentiated leukoplasts, which are transformed into electron-dense plastids during autolysis of the suspensor. This early plastidal differentiation in discussed with respect to its control and functional significance.     

Rumex subgen.Rumex sect.Axillares (Polygonaceae) in South America

Rumex sect.Axillares first described by the author in 1937 is known to be represented in N. America by 20 endemic species. From S. America only five species, all endemic, were previously known; this number is raised in the present revision to 15. Outside America there exist small groups of endemic species in S. Africa and in the islands of Hawaii as well as in Australia and scattered single species in C. Africa, on the remote Atlantic islands of Tristan de Cunha and Gough Island, and in eastern Asia.—In Europe the section is completely absent, except for three American species which occur as rare aliens. TheAxillares are not only morphologically well-characterized by their growth—no leaf rosettes, flowering period practically unlimited by formation of later flowering axillary shoots, flowers often monoecious—but also by their genetic structure expressed by a different behaviour in hybridization. Hybrids are frequent between species of sect.Rumex (syn.Simplices) and are easy to recognize by their sterility which results in a certain peculiarity in habit. In contrast, no spontaneous hybrids are known with certainty among species of sect.Axillares. Most artificial hybrids are fairly fertile. So far only three hybrids between species of the two sections have been known. Three more have been found by the author in Chile, all being highly sterile and showing the growth characteristics resulting from sterility as occur in hybrids between species of sect.Rumex.

---

Resumen Rumex secciónAxillares fue descrito por primera vez por el autor en 1937. Se conoce en América del Norte con 20 especies endémicas. De América del Sur se conocían sólo 5 especies, todas ellas endemicas; este número asciende a 15 en la presente revision. Fuera de las Américas se encuentran pequeños grupos de especies endémicas en Sudáfrica, en las Islas de Hawaii y en Australia; especies aisladas se encuentran en Africa Central, en las lejanas Islas Atlánticas de Tristan de Cunha y Gough Island y en Asia Oriental. La sección no tiene representantes en Europa, con excepción de tres especies americanas introducidas, poco frecuentes. La secciónAxillares se caracteriza bien, tanto morfológicamente por su forma de creciemiento—sin rosetas foliares, período floral prácticamente ilimitado por la formación de inflorescencias axilares parciales, flores a menudo monoicas—como por su estructura genética que se manifiesta en un comportamiento diferente de hibridación. Híbridos son frecuentes en las especies de la secciónRumex (syn.Simplices) y reconocibles por su esterilidad que se expresa en cierta particularidad del habito. Al contrario, no hay seguridad de híbridos espontáneos en la secciónAxillares. La mayor parte de los híbridos artificiales son totalmente fértiles. Hasta ahora se conocían solo tres híbridos entre las especies de ambas secciones. Otros tres han sido encontrados por el autor en Chile. Todos completamente estériles y con las características de hábito típicas resultantes la esterilidad, como ocurre en los híbridos de la secciónRumex.

     

A reappraisal ofPelargonium sect.Ligularia (Geraniaceae)

A study of 33 species ofPelargonium sect.Ligularia reveals four basic chromosome numbers, x = 8, 9, 10, and 11, and variation in chromosome size. From evidence of karyology and hybridization attempts, proposals are made to divide the section into smaller groups and to transfer some species to other sections.     

Synopsis ofVerbesina sect.Ochractinia (Asteraceae)

A taxonomic revision ofVerbesina sect.Ochractinia is presented. 40 species plus 7 additional infraspecific taxa are recognized. A table of the distribution of the species, key to the species and synonymy are included. Relationships to other sections ofVerbesina and species relationships within sect.Ochractinia are discussed.     

Sero-systematics ofCytisus sect.Trianthocytisus (Fabaceae)

The serological reaction of seed proteins provides evidence for a partly new systematic arrangement ofCytisus sect.Trianthocytisus and ofCytisus s.l. Proposed modifications agree with recent advances in morphological taxonomy. Sect.Trianthocytisus includes only two species,C. villosus andC. aeolicus. Its position is central within the genus, and this fact agrees with the proposed retypification ofCytisus (type species:C. villosus).C. emeriflorus, formerly included in the same section, constitutes the monospecific sect.Emeroides, which is intermediate towards the genusLembotropis. This is serologically isolated and includes onlyL. nigricans. It is confirmed thatC. sessilifolius should be removed from the genusCytisus as a monospecific genus:Cytisophyllum Lang which is closely allied toHesperolaburnum and toPodocytisus, the most primitive genera ofGenisteae.     

Gibberellin-like activity in suspensors of Tropaeolum majus L. and Cytisus laburnum L.

Gibberellins in the embryo-suspensor system have been considered so far only in Phaseolus coccineus. We present in this report the localization of gibberellin-like substances in the suspensors of Tropaeolum majus L. and Cytisus laburnum L. The total gibberellin activity (expressed as gibberellic-acid equivalent in the -amylase bioassay) in 2000 suspensors (106 mg fresh weight; FW) of C. laburnum and in 600 suspensors (236 mg FW) of T. majus were 50.9 g g^-1 FW and 8.9 g g^-1 FW respectively.Abbreviation GA gibberellin     

Chromoplasts of Tropaeolum majus L.: Lipid synthesis in whole organelles and subfractions

Isolation of tubulous chromoplasts from Tropaeolum majus L. petals was achieved in pure form. Their main substructures-lipid bodies, tubules, and envelope membranes-have been enriched. Whole chromoplasts as well as substructures have been tested for their activities in lipid synthesis. The following activities were found: fatty acid synthesis from acetate, glycosyl transfer reactions from UDP-galactose and UDP-glucose to galactolipids and sterols, acyltransferase reactions from palmitoyl-CoA, and a very active acyl-CoA hydrolase (EC 3.1.2.2.). Fatty acid synthesis was restricted to whole chromoplasts. Glycosyl- and acyltransferases were essentially confined to envelope membranes, whereas acyl-CoA hydrolase was found in all fractions. The chemical composition of chromoplast subfractions was determined. The lipid bodies consisted mainly of galactolipids and carotenoid esters in a 1:1 ratio, together with small amounts of protein.     

Natural hybrids in Chamaecrista sect. Absus subsect. Baseophyllum (Leguminosae-Caesalpinioideae): genetic and morphological evidence

Summary Chamaecrista sect. Absus subsect. Baseophyllum (Leguminosae–Caesalpinioideae) includes eight species with geographic distribution concentrated in the Cadeia do Espinha?o mountain range of the States of Bahia and Minas Gerais, in eastern Brazil. During the taxonomic revision of this group, individuals were encountered with intermediate morphology between three sympatric species. Allozyme and multivariate morphometric analyses were carried out to investigate the occurrence of hybridization between these species in four populations and 14 suspected hybrid individuals between C. confertiformis and C. depauperata and between C. blanchetii and C. confertiformis. The results of both analyses support the occurrence of natural hybridization between them, and that all the individuals with intermediate morphology are probably F1 hybrids. The hybrids are formally described as C. × blanchetiformis and C. ×patyensis.     

Phenetic studies in Nuphar Sm. (Nymphaeaceae):variation in sect. Nuphar

Previous taxonomic assessments of the hydrophytic genus Nuphar Sm. (Nymphaeaceae) have inadequately considered the range of morphological variation existing in the genus. A comprehensive statistical analysis of morphological features among the recognized taxa of Nuphar is presented here, first using 10 fruit characters to delimit the separation of two infrageneric groups of taxa, and secondly using 31 characters to quantify the morphological variation found within sect. Nuphar. Results from combined phenetic analyses provided empirical support for recognition of two infrageneric groups of species in the genus (largely by fruit, stigmatic disk margin, neck, and stigmatic disk size) and for separation of taxa within sect. Nuphar (by leaf size, and fruit disk and neck size). Statistical analyses demonstrate that extensive variation in ranges of character states exists among sect. Nuphar taxa. However, a suite of qualitative and quantitative characters can be used to recognize five species and three subspecies within this predominantly Old World assemblage.     

Isozyme evidence of relationships withinSphagnum sect.Acutifolia (Sphagnaceae,Bryophyta)

Ten enzyme systems coding for 13 putative loci were studied in 12 European species ofSphagnum sect.Acutifolia (peat mosses). On average seven plants from each of 73 collections made throughout N Europe were analysed. Despite complex morphological variation, all species expressed unique allele combinations. Principal components and cluster analyses based on allele frequencies showed that sect.Acutifolia can be divided into two major groups; one comprisingSphagnum fimbriatum, S. girgensohnii andS. molle; the second comprisingS. capillifolium, S. angermanicum, S. fuscum, S. quinquefarium, S. rubellum, S. subfulvum, S. subnitens andS. warnstorfii. These groups are linked byS. russowii, which is shown to be an allopolyploid, with fixed heterozygosity at 6 to 8 of the loci. The presumed progenitors ofS. russowii areS. girgensohnii andS. rubellum. Allelic data also indicate thatS. teres from sect.Squarrosa is related toS. girgensohnii andS. fimbriatum. The bisexual species tend to be less variable than the unisexual species.     

Breeding systems and hybridization inPetrorhagia sect.Kohlrauschia (Caryophyllaceae)

In the diploid speciesP. prolifera, the evolution of autogamy has resulted in the presence of distinct selfing and outcrossing races. The change in breeding system toward autogamy is associated with a reduction in the size of floral features and a lower pollen production. The outbreeding populations have only been found in Greece, while the selfing populations have colonized much of Europe. The two races appear to be reproductively isolated from one another. —Petrorhagia sect.Kohlrauschia contains four species.P. glumacea is outbreeding and shows unilateral incompatibility with the small-flowered race ofP. prolifera and interfertility with the large-flowered race; althoughP. glumacea is sympatric with the large-flowered race, they do not appear to hybridize in the wild. Amongst the allopatric species, internal breeding barriers are also found. These may take the form of hybrid sterility or seed incompatibility.P. velutina is autogamous and reproductively isolated from all the other species of the section regardless of whether they are sympatric or allopatric. It is suggested that the evolution of autogamy has been an important factor in the spread ofP. velutina, the tetraploidP. nanteuilii, and the small-flowered race ofP. prolifera in Europe.     

Ribosomal DNA spacer-length variation inSecale spp. (Poaceae)

Variation in ribosomal DNA spacer length was analysed in 23 populations of 12Secale spp. by restriction enzyme analysis. Digestion of rDNA with Taq I endonuclease enzyme yields spacer fragments that include the subrepeat array and the non-repetitive region downstream of the array. Extensive spacer length variation existed in most species with Taq I fragment lengths ranging from 0.9–3.1 kb. These length variants have been attributed to the differences in number of 134 bp spacer subrepeats within rDNA arrays.S. silvestre was the only species to exhibit a unique spacer length variant of 0.9 kb and this was shown to result from the presence of an extra Taq I site in the spacer. rDNA spacer length frequencies were determined for the species. These frequencies were used to derive phenetic relationships between the species by numerical taxonomic methods. In plots constructed fromGower's distance matrices,S. silvestre appeared well separated from the major cluster consisting of the other species. On the basis of morphological and cytogenetic criteria,S. silvestre is considered the most ancient species. The rDNA data is consistent with this interpretation as it shows a clear differentiation ofS. silvestre from all the other species based on length and nucleotide sequence composition of the spacer region.     

Investigations on the content of steroidal alkaloids and sapogenins withinSolanum sect.Solanum ( = sect.Morella) (Solanaceae)

Plant material of 55 strains from 32 species ofSolanum sect.Solanum ( = sect.Morella) has been investigated for the content of steroidal alkaloids and sapogenins. The leaf-extracts of all species contain diosgenin resp. tigogenin or both together in considerable amount. On the contrary the characteristic steroidal alkaloid solasodine was absent in these samples. For 29 species the presence of steroidal sapogenins has been proved for the first time. In unripe fruits, however, solasodine could be detected in most of the species examined. New occurences of solasodine are reported for seven further species. These chemical results may contribute to a better comprehension of the difficult genusSolanum.—Ontogenetic studies on different organs ofS. atriplicifolium show the relationship between accumulation of steroidal compounds and plant maturity.     

Chromosomal evolution inEpilobium sect.Epilobium (Onagraceae)

WithinEpilobium sect.Epilobium, a cytological analysis of 121 experimental hybrids, involving 40 species, indicates the presence of a widespread BB chromosome arrangement in Eurasia, Africa, and Australasia, as well as in North and South America less commonly. The AA chromosome arrangement, which differs from BB by one reciprocal translocation, occurs in North America, South America, and in at least three European species. The CC arrangement, which differs from AA by two reciprocal translocations, characterizes theAlpinae, a circumboreal group. Distinctive or only partly worked out chromosome arrangements occur in the EuropeanE. duriaei andE. nutans and in the North AmericanE. luteum, E. obcordatum, E. oregonense, andE. rigidum. With earlier results, the chromosome arrangements of some 65 of the estimated 185 species of the section have been established fully or partly.     

DNA evidence for multiple origins of intergeneric allopolyploids in annualMicroseris (Asteraceae)

Seventy populations of North American annualMicroseris, Stebbinsoseris, andUropappus species were examined for chloroplast and nuclear ribosomal DNA restriction site variability to determine the origin of the allotetraploid speciesS. heterocarpa andS. decipiens. Previously identified chloroplast DNA restriction site variants were used in concert with restriction site variation forNco I in the nuclear-encoded ribosomal DNA repeat. The presence of two, mutually exclusive restriction site gains were observed in diploid populations ofM. douglasii; these same variants were also found in populations of allotetraploidS. heterocarpa, indicating mutiple origins of this species from different maternal diploid populations ofM. douglasii. Variation in the rDNA repeat between the diploid annual species and the putative paternal genome ofU. lindleyi was found to be additive inS. heterocarpa. A similar relationship was observed for the origin ofS. decipiens; cpDNA restriction site variants found inM. bigelovii andM. douglasii were present inS. decipiens. The rDNANco I variants also were additive in this purported allotetraploid. These results confirm the reticulate evolutionary pattern inStebbinsoseris and provide another example of multiple origins of intergeneric allopolyploids.     

Ribosomal RNA genes inQuercus spp. (Fagaceae)

The taxonomy of the genusQuercus is still unclear. In order to elucidate the taxonomy of Mediterranean oaks we have analyzed ribosomal RNA genes ofQuercus cerris, Q. coccifera, Q. trojana, Q. ilex, Q. suber, andQ. macrolepis by means of Southern blot hybridization. Oak nuclear DNA was extracted from root tips of 300 acorns and from catkins of single plants. EcoRI and BamHI restriction endonucleases were used. DNA electrophoresis and rRNA/DNA hybridization were performed usingVicia faba rRNA 18 S and 25 S as probes. The rRNA genes of all the species studied have an identical restriction mapping in the 18 S and 25 S regions, while differences in length are present in the intergenic regions.Q. cerris possesses at least four types of genes of 12.1, 11.5, 8.5, and 8.3 kb;Q. coccifera at least three types of 12.4, 10.4, and 10.1 kb;Q. trojana possesses the same rRNA genes asQ. cerris plus another gene type 12.0 kb long, with EcoRI and BamHI restriction sites in the intergenic spacer;Q. ilex at least three types of 12.4, 10.85, and 9.5 kb;Q. suber at least five types of 11.5, 11.0, 8.6, 8.5, and 8.3 kb;Q. macrolepis, finally, at least seven types of 11.5, 11.0, 10.2, 8.6, 8.5, 8.3, and 8.15 kb.Q. coccifera andQ. ilex rDNA appears quite different respect to other species examined, while high similarity seems to exist betweenQ. cerris, Q. trojana, Q. suber, andQ. macrolepis. These results are in agreement with the taxonomic model proposed bySchwarz for the genusQuercus.     

Delimitation ofPelargonium sect.Glaucophyllum (Geraniaceae)

Pelargonium otaviense Knuth andP. spinosum Willd. are excluded from sect.Glaucophyllum, whileP. grandiflorum (Andr.)Willd.,P. patulum Jacq. andP. tabulare (Burm. f.)L'Hérit. of sect.Eumorpha are included. Sect.Glaucophyllum is characterized by green to glaucous vegetative organs and zygomorphic white to pink corolla with five narrow petals. All the species have an identical pollen and chromosome morphology, the same basic chromosome number (x = 11) and similar flavonoid patterns. A close relationship between sect.Glaucophyllum and sect.Pelargonium is indicated by the occurrence of natural hybrids and concordant characters. Isorhamnetin and luteolin have been detected in the genus for the first time.     

Species relationships inFestuca (sect.Ovinae,Poaceae)

Morphological and seed protein studies of selected species ofFestuca were performed to elucidate the relationships between species of sect.Ovinae and their affinity with other fescues.Festuca rubra andF. heterophylla (extravaginale group) had higher phenetic affinity with taxa of sect.Scariosae, Montanae andBovinae than with members of their own section. 5 species of the intravaginale group (F. ovina, F. capillata, F. valesiaca, F. sulcata, F. trachyphylla) were fairly uniform and well defined by protein and morphological data. This supports current systematic thinking and does not agree withHackel's (1882) classification of this group.     

Karyological notes onCentaurea sect.Acrocentron (Asteraceae)

The karyology ofCentaurea sect.Acrocentron is surveyed. 19 chromosome counts on 8 species are reported; those onC. acaulis, C. crocata, C. galianoi, C. pubescens, andC. malinvaldiana are new. The basic chromosome numbers of the section are x = 11 and x = 10. Karyological arguments have been used to show that evolution was from x = 11 to x = 10. This is supported by biogeographical data. Two main centres of diversification of sect.Acrocentron were studied from that point of view: the East and the Southwest Mediterranean region.