Revision of Bomarea subgenus Sphaerine (Alstroemeriaceae)

The subgenus Sphaerine of Bomarea (Alstroemeriaceae) recently contains 33 validly published names. Extended field studies in Peru and extensive investigation of herbarium material revealed the urgent necessity for a revision. As a result, the number of species is reduced to 12, among them two species are newly described. 9 species are members of an other subgenus. A key to determine the species is given in English and Spanish. The typical growth form and its variability, habitat preferences and distribution are discussed separately for each species.     

Panbiogeographical analysis of the genus Bomarea (Alstroemeriaceae)

Aim  A panbiogeographical analysis of the genus Bomarea was undertaken in order to determine generalized tracks and biogeographical nodes, and to evaluate the current distribution of the genus based on the available tectonic information and the biogeographical regionalization of Latin America.
Location  The Neotropical region from northern Mexico to northern Argentina, and the Nearctic and Andean regions.
Method  A total of 2205 records of 101 species were analysed, representing 95% of the species assigned to Bomarea . Localities were represented on maps and their individual tracks were drawn. Based on their comparison, generalized tracks were detected and mapped. Nodes were identified in the areas where different generalized tracks were superimposed.
Results  Five generalized tracks were recovered. One is located in the Eastern Central America and Western Panamanian Isthmus provinces (Caribbean subregion, Neotropical region), which was supported by three species of Central American distribution. The four remaining generalized tracks were located in South America, in the North Andean Paramo, Cauca and Puna biogeographical provinces. These tracks were supported by species of Bomarea with an Andean distribution. Biogeographical nodes were established in the Central Andean region of Colombia, central Ecuador and central Peru.
Main conclusions  The nodes obtained for Bomarea support a hybrid origin for the Andean region, which presents diverse components from both northern and southern South America. Likewise, the track recovered between Colombia and Ecuador includes Andean and Neotropical areas, providing further support for this hypothesis. The nodes obtained are coherent with vicariant elements evident for Bomarea. Species of three clades proposed for Bomarea are distributed in specific generalized tracks.     

Functional leaf anatomy of Bomarea Mirb. (Alstroemeriaceae)

The leaf anatomy of Bomarea is described and related to ecological conditions. The principal architecture of all species is very similar; adaptations are developed in numerous differences, for instance degree of lignification. All species have inverse leaves, the adaxial side being the stomatous side. In most species, the leaves are resupinate, the lower surface being the adaxial. Theories for the cause of resupination are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 73–90.     

Chromosome studies in Andean taxa of Alstroemeria (Alstroemeriaceae)

Meiotic or mitotic chromosomes of seven Alstroemeria taxa, native in Argentina and Chile and with Andean distribution were studied: A. andina ssp. venustula, A. hookeri ssp. cummingiana , A. hookeri ssp. recumbens , A. pallida , A. patagonica , A. pseudospathulata and A. pygmaea . All were diploid with 2 n = 16. Karyotypes of A. andina ssp. venustula and A. pygmaea were analysed, revealing similarity to previously analysed species. Thus, to all existing arguments for not retaining Schickendantzia as a separate genus, we can add another one which merges A. pygmaea with other Alstroemeria species, and does not support its taxonomic uniqueness. In general, the meiotic behaviour was normal, with regular formation of eight bivalents except in A. hookeri ssp. cummingiana , in one plant of which meiotic irregularities at various stages were observed. At the tetrad stage a large percentage of the cells presented micronuclei. The presence of 0–2 supernumerary chromosomes in A. hookeri ssp. recumbens is recorded. The karyotype asymmetry presented by most Alstroemeria species is discussed. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 451–459.     

Two new species of Alstroemeria L. (Alstroemeriaceae) from Brazil

Alstroemeria hygrophila andA. orchidioides are both described as new from the state of Goiás (including Distrito Federal) in Brazil.Alstroemeria hygrophila is a cerrado bog-dwelling species that grows semi-epiphytically on the culms of a sedge. The vegetative stems of this species characteristically zig-zag between the pseudonodes. Like other wetlandAlstroemeria in Brazil (e.g.,A. apertiflora, A. isabellana, andA. sellowiana),A. hygrophila has wiry stems bearing narrow, lanceolate, nonresupinate leaves, and does not form root tubers.Alstroemeria orchidioides is a hysteranthus forest understory species with large vegetative leaves aggregated at the apex of the stems. The leaves on flowering stems are reduced to scarious bract-like scales. Its flowers are a pale, nearly white, greenish yellow, a color not reported in any other Brazilian species. Both species have 2n=16 chromosomes.     

Comparative karyotypic analysis in the Alstroemeria hookeri Lodd. (Alstroemeriaceae) complex sensu Bayer (1987)

Alstroemeria L. (Alstroemeriaceae) is an American genus of monocots with two principal distribution centers in Chile and Brazil. In Chile, it is represented by about 32 species, most of them in central Chile, an area known for its high level of endemism. The "complex" Alstroemeriahookeri is endemic to Chile, where it is distributed from the Coquimbo to the Bío-Bío Region. We analyzed the karyotypes of 36 populations of this complex along its natural distribution. Ten metaphases per population were used for chromosome measurements. All analyzed subspecies presented a well defined asymmetric karyotype. The populations of A. hookeri subsp. hookeri collected in the coastal range of the Bío-Bío Region and the populations from the Central Valley of this Region (Pangal del Laja) presented striking morphological differences in the karyotype, mainly on chromosome 3. The population of A. hookeri subsp. recumbens from Pichicuy showed a polymorphism on chromosome 7, which differed from the other analyzed populations of this subspecies. Phenetic analysis suggested that A. hookeri subsp. cummingiana, which showed a more symmetrical karyotype and did not grow in sandy soil, should be alocated to A. cummingiana rather than considered as part of the hookeri complex.     

Effects of crossing distance and genetic relatedness on pollen performance in Alstroemeria aurea (Alstroemeriaceae)

Prezygotic barriers may represent effective mechanisms to avoid the deleterious effects of inbreeding. This study reports the existence of distance-dependent prezygotic barriers in self-compatible Alstroemeria aurea, a clonal herb native to temperate forests of the southern Andes. We analyzed pollen germination and tube growth as indicators of donor-recipient affinity using crossing distances of 1, 10, and 100 m. We used allozyme electrophoresis to determine the actual genetic relatedness between donor and recipient ramets. Pollen germination was not affected by distance between mates, but the number of pollen tubes reaching the base of the style increased strongly with distance between donor and recipient. This pattern was related to an increase in genetic dissimilarity with distance between mates. In contrast, pollen tube-style interactions did not change with distance when we restricted analysis to individuals at different distances that appeared to be genetically identical. This test implied genetic dissimilarity as the critical factor affecting pollen performance. We propose that the existence of prezygotic barriers might contribute to the high degree of genetic mixing exhibited by some clonal species.     

Selective fruit filling in relation to pollen load size in Alstroemeria aurea (Alstroemeriaceae)

 We conducted an experiment in a natural population of Alstroemeria aurea, a clonal perennial, to determine (1) if reproduction was resource limited, and (2) if fruits would be selectively filled based on differences in pollination intensity when pollen loads were adequate for full seed set. Under these conditions, differences in pollination intensity are unlikely to affect seed number, but could affect seed quality, providing an interesting test of the gametophytic competition hypothesis. To test for resource limitation, percent fruit maturation, number of seeds per fruit and average seed weight were compared to paired controls for ramets in which all but one fruit was removed. To test the effect of pollination intensity on selective resource allocation, three types of pollination treatments were performed: (1) all flowers of the single inflorescence received a low pollen load, (2) all flowers received a high pollen load, (3) alternate flowers of the inflorescence received either a high or a low pollen load. We determined the percentage of fruit that reached maturity, counted the number of seeds and ovules and calculated the average seed weight for all capsules in each treatment. Resources appeared to limit reproduction in this population since seed number and weight were significantly higher than in controls when competing capsules were removed. At the whole ramet level, a four fold difference in pollen loads had no significant effect on any of the parameters measured. However, when pollination intensity varied within an inflorescence, the number of seeds per fruit increased by about 10% in flowers that received the higher pollen load. We observed the same trend in each of 2 years, but the increase was significant in only 1 year. The differences, although not great, were only slightly smaller than when all competing fruits were removed, and were consistent with selective resource allocation based on pollination intensity independent of seed set. Received: 28 September 1997 / Accepted: 30 April 1998     

Ribosomal DNA distribution and a genus-wide phylogeny reveal patterns of chromosomal evolution in Alstroemeria (Alstroemeriaceae)

? Premise of the study: Understanding the flexibility of monocot genomes requires a phylogenetic framework, which so far is available for few of the ca. 2800 genera. Here we use a molecular tree for the South American genus Alstroemeria to place karyological information, including fluorescent in situ hybridization (FISH) signals, in an explicit evolutionary context. ? Methods: From a phylogeny based on plastid, nuclear, and mitochondrial sequences for most species of Alstroemeria, we selected early-branching (Chilean) and derived (Brazilian) species for which we obtained 18S-25S and 5S rDNA FISH signals; we also analyzed chromosome numbers, 1C-values, and telomere FISH signals (in two species). ? Key results: Chromosome counts for Alstroemeria cf. rupestris and A. pulchella confirm 2n = 16 as typical of the genus, which now has chromosomes counted for 29 of its 78 species. The rDNA sites are polymorphic both among and within species, and interstitial telomeric sites in Alstroemeria cf. rupestris suggest chromosome fusion. ? Conclusions: In spite of a constant chromosome number, closely related species of Alstroemeria differ drastically in their rDNA, indicating rapid increase, decrease, or translocations of these genes. Previously proposed Brazilian and Chilean karyotype groups are not natural, and the n = 8 chromosomes in Alstroemeria compared to n = 9 in its sister genus Bomarea may result from a Robertsonian fusion.     

Comparative karyotype analysis of populations in the Alstroemeria presliana Herbert (Alstroemeriaceae) complex in Chile

Alstroemeria L., one of the most diverse genera of the Chilean flora and of high floricultural value, is represented by 35 species, most of them distributed between 28–38° S in the Mediterranean zone of Central Chile. There are 24 complex-forming taxa, of which 18 have conservation problems (8 are considered “endangered” and 10 as “vulnerable”). One of these complexes is Alstroemeria presliana Herb. with two subspecies: subsp. presliana and subsp. australis Bayer. Alstroemeria presliana grows in Chile and Argentina: subsp. presliana is distributed from Reserva Nacional Siete Tazas (35°27′ S, Region of Maule) to Antuco, (37°25′ S, Region of Bío-Bío), and is also found in Neuquén, Argentina; subsp. australis is endemic to the Cordillera of Nahuelbuta. A comparative karyotype study was carried out among six populations of A. presliana subsp. presliana and five populations of A. presliana subsp. australis. The eleven populations presented an asymmetric karyotype, with 2n = 2× = 16 chromosomes but with different karyotype formulae. A. presliana subsp. presliana shows the haploid formula 2m + 2m-sat + 1sm-sat + 1st-sat + 1t + 1 t-sat, and A. preslianasubsp. australis presents a formula 1m + 2m-sat + 1sm + 2t + 2t-sat chromosomes. The architecture of the karyotype between the subspecies is very different. The scatter plot among CV_CL vs. M_CA shows different groupings between populations of the two subspecies. According to the results obtained it is possible to consider raising Alstroemeria presliana subsp. australis at species level.     

Foliar anatomy and micromorphology of southern South American Alstroemeriaceae: Alstroemerieae,and its systematic implications in Alstroemeria

Alstroemerieae is an exclusively Central and South American tribe belonging to Alstroemeriaceae, which comprises two large genera, Alstroemeria and Bomarea. Alstroemeria has two areas of distribution, mediterranean Chile and central southeastern Brazil. Most Bomarea species grow in forests and hedges in moist areas, however, some species are adapted to dry Andean valleys and high altitudes. Previous leaf anatomical data were obtained from a limited group of species. To assess the value of the anatomical characters for the systematics and their importance as adaptations to different environments, we compared representative species from different geographical areas and habitats. Data regarding leaf anatomy and micromorphology were obtained from light microscopy and scanning electron microscopy and were combined with macromorphology for 27 Alstroemerieae species. In accordance with earlier studies, our results show variation in relation to several leaf morpho‐anatomical characters. Based on these we define seven types. We furthermore analyzed the morpho‐anatomical characters in a phylogenetic context. Morpho‐anatomical characters are highly homoplastic within the family. Leaf anatomy may support monophyly of Baker's informal grouping of Alstroemeria Brazilian species with rigid leaves, however, a more thorough study of Brazilian Alstroemeria species are needed to confirm this.     

Influence of local floral density and sex ratio on pollen receipt and seed output: empirical and experimental results in dichogamous Alstroemeria aurea (Alstroemeriaceae)

Local density and sexual composition are two aspects of floral neighborhoods thought to influence pollination and seed output of recipient plants. I characterized the floral neighborhood of 436 flowering ramets of Alstroemeria aurea, a southern Andean perennial, distributed among three sites. On each ramet, I measured total pollen receipt and seed output. The long-lived, bumblebee-pollinated flowers of A. aurea are synchronously protandrous with a given ramet being either all male or all female and thus incapable of self or geitonogamous pollination at the ramet level. Even though each ramet changes sex over time, A. aurea forms floral neighborhoods that remain stable with respect to density and sex ratio during the span of a focal ramet female phase. Contrary to expectation, under field conditions neither local density nor sexual identity explained significant amounts of variation in pollen receipt. Density of neighboring flowering ramets marginally affected pollen receipt in two of the three populations but in opposite directions. Despite the absence of strong effects of neighborhood sexual composition on pollen receipt, the sexual identity of neighbors affected seed output which suggests effects on the quality of pollination due to changes in patterns of pollen flow. I also compared pollen loads on the stigmas of artificially isolated ramets (6 m) with those on experimental focal ramets surrounded by six close neighbors (20 cm) that were either all male or all female. Here, pollen receipt by focal ramets in all-male neighborhoods was 1.3 times greater than in isolated ramets, and 3.8 times greater than in ramets in all-female neighborhoods. In these artificial neighborhoods, stigmatic pollen deposition increased significantly over time. In nature, rates of bumblebee visits were higher in female-biased (early-flowering) than in male-biased (late-flowering) co-occurring floral patches. Thus, spatio-temporal shifts in visitation frequencies associated with the sexual composition of floral neighborhoods might compensate for spatial variability in pollen availability within populations and explain the discrepancies between empirical and experimental results. Received: 11 June 1996 / Accepted: 24 March 1997     

Generic Delimitation and Phylogeny of the Subtribe Centaureinae (Asteraceae): A Combined Nuclear and Chloroplast DNA Analysis

Taxonomic complexity has hindered the partitioning of the artificial genus Centaurea, even though it has long been recognized as a polyphyletic assemblage. On the basis of morphology, pollen type, karyology and DNA sequence analysis, previous workers have defined five informal groups in the genus (Acrocentron,Centaureasensu stricto , Cyanus, Jacea and Psephellus). However, the precise delimitation of these groups and their relationships remain largely unknown. Moreover, although some informal groups have been established among the rest of the subtribe (Amberboa, Carthamus and Stemmacantha), many genera cannot be classified in any group. Newer molecular approaches are essential for resolving these problems. Sequences of the internal transcribed spacers (ITS) of nuclear ribosomal DNA and the chloroplast gene mat K were analysed for a comprehensive sample of the whole subtribe, with the aim of clarifying the delimitation and the phylogeny of the groups of the Centaureinae. Results largely confirm the suggested informal entities as natural groups, with some interesting changes of placement of some genera, especially in the Acrocentron and the Stemmacantha groups. Our results confirm that the sections of Centaurea with Dealbata pollen type should be classified as a different genus, Psephellus. In addition to morphology and suggested pollen type evolution, the ITS and mat K phylogenies are also supported by karyological evidence. Our results confirm that the natural delimitation of Centaurea that minimizes nomenclatural changes is possible only if a new type of the genus is designated.     

Satellite DNA probes of Alstroemeria longistaminea (Alstroemeriaceae) paint the heterochromatin and the B chromosome,reveal a G-like banding pattern,and point to a strong structural karyotype conservation

Alstroemeria species present a well-conserved and asymmetric karyotype. The genus is divided into a Chilean clade, rich in heterochromatin, and a Brazilian clade, poor in heterochromatin. We investigated the distribution of the main repetitive sequences in the chromosomes of the Brazilian species A. longistaminea (2n = 16 + 0-6B) aiming to evaluate the role played by these sequences on the structural organization of the karyotype. In situ hybridization of the three most abundant retrotransposons, corresponding to ~ 45% of the genome, was uniformly distributed. Three satellite DNA sequences, representing near half of the whole satellite fraction (1.93% of the genome), were mainly concentrated on the heterochromatin and one of them painted the whole B chromosome. Noteworthy, some satellites were located on euchromatin, either dispersed or concentrated in clusters along the chromosomes, revealing a G-band-like pattern. The two satellites that presented more C-band- and G-band-like labeling were also hybridized in situ in two other Alstroemeria species. They revealed astonishing similar patterns of distribution, indicating an unusually structural karyotype conservation among Brazilian species.

     

Molecular Phylogeny of Tribe Theeae (Theaceae s.s.) and Its Implications for Generic Delimitation

Tribe Theeae, which includes some economically important and widely grown plants, such as beverage tea and a number of woody ornamentals, is the largest member of the Theaceae family. Using five genomic regions (chloroplast: atpI-H, matK, psbA5''R-ALS-11F, rbcL; nuclear: LEAFY) and 30 species representing four of the five genera in this tribe (Apterosperma, Camellia, Polyspora, and Pyrenaria s.l.), we investigated the phylogeny of Theeae and assessed the delimitation of genera in the tribe. Our results showed that Polyspora was monophyletic and the sister of the three other genera of Theeae investigated, Camellia was paraphyletic and Pyrenaria was polyphyletic. The inconsistent phylogenetic placement of some species of Theeae between the nuclear and chloroplast trees suggested widespread hybridization between Camellia and Pyrenaria, Polyspora and Parapyrenaria. These results indicate that hybridization, rather than morphological homoplasy, has confused the current classification of Theeae. In addition, the phylogenetic placement and possible allies of Laplacea are also discussed.     

Delimitation and classification ofBromus fasciculatus (Poaceae)

The syntypes ofBromus madritensis var.delilei Boiss. comprise two different elements: specimens ofB. haussknechtii Boiss. and ofB. fasciculatus C. Presl s.l. By its lectotypification and on the basis of new morphological characters, the validation ofB. fasciculatus subsp.delilei (Boiss.)H. Scholz 1971 was substantiated. This subspecies represents the eastern marginal segregate of the south-Mediterranean species.B. fasciculatus var.alexandrinus Thell. is a minor variant of the typical subspecies.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

泡果荠属和阴山荠属（十字花科）的分属界限及两属的分类校订

泡果荠属Hilliella和阴山荠属Yinshania是十字花科中国特有的两小群植物,但是,它们的分属界定却一直颇有争议。本文结合前人的研究结果重新对这两群植物做了研究,从而进一步确认这两群植物应该界定为两个属,即泡果荠属和阴山荠属。文中比较了两属间重要的性状特征,并指出它们的主要区别在于:泡果荠属果实无假隔膜,种子较大,表面具小瘤状突起,染色体倍性为六倍体（2n=6x=42）;而阴山荠属果实有假隔膜,种子较小,表面具网纹,染色体倍性为二倍体（2n=2x=12）。两属其他方面如叶形态、叶表皮结构、被毛类型     

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.