Phylogenetics and evolution of the Macaronesian members of the genus Aichryson (Crassulaceae) inferred from nuclear and chloroplast sequence data

Aichryson is a genus of annual or perennial herbs, comprising approximately 13 species. The genus is nearly endemic to Macaronesia, with the center of diversity on the Canary Islands. Previous studies indicate that Aichryson is monophyletic and sister to a clade comprising two other genera, Monanthes and Aeonium. However, phylogenetic relationships within the genus have yet to be investigated. Phylogenetic relationships were estimated for Aichryson using DNA sequence data from both the nuclear and chloroplast genome. Parsimony analyses were conducted using separate nuclear and chloroplast data sets, as well as a combined data set. These combined analyses provide a well-resolved topology that is used to investigate patterns of evolution within the genus. For example, although typically herbaceous in habit, two members display a woody growth form. Our analyses indicate that woodiness is a derived feature in Aichryson that has arisen once. Furthermore, the data presented suggest that the Aichryson pachycaulon group, a group of five subspecies, are in fact not monophyletic and that a reexamination of classification is necessary. The present study indicates that the biogeographical patterns exhibited by Aichryson appear to be rare, occurring in only one other known genus.The authors would like to thank Susana Fontinha of the Madeiran Botanical Garden for plant material of Madeiran Aichryson; Drs. D. Crawford and J. Archibald, as well as two anonymous reviewers for helpful comments; Eastern Illinois Undergraduate Honors Research Fund, and University of Kansas Plant Biology Research Award. Additional financial support for this work was provided by the Department of Ecology and Evolutionary Biology and the Museum of Natural History and Biodiversity Research Center, University of Kansas and a Kansas NSF EPSCoR First Award (NSF 32171) to MEM.     

Infrasectional systematics of the genus Sideritis L. section Sideritis (Lamiaceae)

RIVERA NUÑEZ, D., OBON DE CASTRO, C., TOMAS-LORENTE, F., FERRERES, F. & TOMAS-BARBERAN, F. A., 1990. Infrasectional systematics of the genus Sideritis L. section Sideritis (Lamiaceae). A new taxonomic division of the section Sideritis is proposed on the basis of morphological, cytological and chemical characters. The following subsections art-recognized: Grandiflora, Ovata, Camarae, Linearifolia, Gymnocarpae, Stachydioides, Lacaitae, Hirsuta, Chamaedryfolia, Arborescens, Flavovirens, Leucantha, Angustifolia, Serrata and Scordioides . Possible evolutionary pathways are discussed.     

A numerical analysis of Sideritis L. (Lamiaceae) from North Africa

REJDALI, MOH, 1992. A numerical analysis of Sideritis L. (Lamiaceae) from North Africa. A numerical analysis is shown to be useful means of discriminating taxa at all levels of the hierarchy in the genus Sideritis. This analysis supports both the separation of the genus into the two sections recognized by Bentham, the three groups of Briquet's treatment and the four groupings as suggested by various lines of evidence presented by Rejdali (1988).     

Diversification and vicariance of desert plants: Evidence inferred from chloroplast DNA sequence variation of Lagochilus ilicifolius (Lamiaceae)

To document arid Northern China as a diversification center for desert plants, and to better understand the mechanisms of desert taxa diversification, we used five cpDNA spacers (trnL-trnF, rps16, psbA-trnH, psbK-psbI and trnS-trnG) to investigate Lagochilus ilicifolius in all distribution areas, including Northern China, Mongolia and Russia. Phylogenetic analyses showed that L. ilicifolius comprises two distinctive lineages, one distributed in China, and another in Mongolia–Russia. Our data confirmed that arid Northern China, is a distinctive area with many endemic genera. Biogeographic inferences, based on a Bayesian uncorrelated lognormal model together with molecular dating, suggested that the main diversification within the species occurred in the Pleistocene (ca. 1.38–0.3 Ma), resulting from the transition of the climate of Eurasia to a dry-cold pattern as well as the desertification caused the vicariance of desert plants.     

Preliminary phylogeny of Valerianaceae (Dipsacales) inferred from nuclear and chloroplast DNA sequence data

Valerianaceae is a relatively small (ca. 350 species), but morphologically diverse angiosperm clade. Sequence data from the entire ndhF gene, the trnL-F intergenic spacer region, the trnL intron, the matK region, the rbcL-atpB intergenic spacer region and internal transcribed spacer (ITS) region of nuclear ribosomal DNA were collected for 21 taxa within Dipsacaceae and Valerianaceae (1 and 20, respectively). These data were included in several phylogenetic analyses with previously published sequences from Dipsacales. Results from these analyses (maximum parsimony, maximum likelihood, and Bayesian analysis) are in strong agreement with many of the conclusions from previous studies, most importantly: (1) Valerianaceae is sister to Dipsacaceae; (2) Triplostegia is more closely related to species of Dipsacaceae than to Valerianaceae; and (3) Valeriana appears not to be monophyletic, with Valeriana celtica falling outside the remainder of the species of Valeriana sampled here (with very strong support). With the exception of V. celtica, these data support two major clades within Valeriana; one that is exclusively New World and another that is distributed in both the Old and New World. Although the species of Valerianaceae and its sister group Dipsacaceae plus Triplostegia, are widely distributed in the Northern Hemisphere, and the data imply that Valerianaceae diversified initially in Asia (the Himalayan Patrinia and Nardostachys falling at the base of the clade), the center of modern species diversity for the group is in the Andes of South America with as many as 175 species restricted to that region. Although the exclusively South American taxa form a clade in the chloroplast and combined ITS and chloroplast analyses, support values tend to be low. Future studies will need to include additional data, in the form of both characters and taxa, before any strong conclusions about the character evolution, diversification, and biogeography of the South American valerians can be made.     

Phylogenetics and diversification of Cotyledon (Crassulaceae) inferred from nuclear and chloroplast DNA sequence data

Crassulaceae includes approximately 35 genera and 1500 species of leaf and stem succulent flowering plants. The family is nearly cosmopolitan in distribution, but is particularly diverse in southern Africa, where five genera comprising approximately 325 species are found. One of these genera, Cotyledon, includes 10 species that are largely confined to South Africa, where they are commonly found on rocky hillsides, coastal flats, and cliff faces. Species of Cotyledon are characterized by five-parted, pendulous, sympetalous flowers, but the genus is highly diverse in growth form, flower color and size, and leaf morphology. One particularly variable species, C. orbiculata, has been divided into five varieties based on leaf morphology and biogeography; however, the monophyly of this species as well as the relationships among the varieties have not previously been investigated. Parsimony analyses of a combined data set of DNA sequences from chloroplast and nuclear genome provided the first estimate of phylogeny for Cotyledon, and resulted in two minimum-length trees and a fully resolved phylogeny for the genus. Results indicate that C. orbiculata is not monophyletic and suggest the need for additional studies and a revised classification within the genus.     

中国唇形科草糙苏属分类学研究杂记(英文)

最近的研究结果表明,国产糙苏属(Phlomis)的所有种类(除了栽培种橙花糙苏P.fruticosa以外)以及沙穗属(Eremostachys)、独一味属(Lamiphlomis)、钩萼草属(Notochaete)均应并入草糙苏属(Phlomoides)。新界定的草糙苏属因其草本习性等特征而与糙苏属易于区别。尽管已有不同作者对国产草糙苏属大部分物种名称进行了新组合,但仍遗漏了11个物种名称。为方便使用,本文对这11个名称进行了新组合,同时提供了新界定的草糙苏属国产物种名录及检索表,共50种9变种。     

Leucas pachmarhiensis sp. nov. (Lamioideae; Lamiaceae) from central India

A new species, Leucas pachmarhiensis (Lamiaceae), is described and illustrated from the Pachmarhi Biosphere Reserve, Hoshangabad district, Madhya Pradesh, India with conservation status assessed as ‘Critically Endangered’. This novelity is endemic to central India and belongs to Leucas sect. Plagiostoma. It differs from other species of this section by its height up to 120 cm, and inflorescence positioned at multiple nodes. Detailed notes on its conservation status and ecology are provided.     

Leaf micromorphology and taxonomy of North African species of Sideritis L. (Lamiaceae)

REJDALI M., 1991. Leaf micromorphology and taxonomy of North African species of Sideritis L. (Lamiaceae). This study reports on the structure of trichomes, epidermal cells, stomata and venation patterns. The trichomes are either glandular or eglandular. The latter are unicellular or multicellular, soft or rigid pointed hairs. The glandular hairs are short or long stalked, the latter are generally flattened and may be sessile. The epidermal cells are variously shaped with sinuous or straight walls usually randomly orientated. Stomata are either anomocytic or diacytic and evenly distributed throughout the epidermis without any definite pattern of orientation. Venation is of the semicraspedodermous type.     

Genome differentiation between two closely related species of Sideritis L. (Lamiaceae)

We have analysed the karyotype structure, DNA amounts and seed protein profiles of three isolated populations of two species: S. saetabensis and S. tragoriganum. These species are closely related from a karyological and morphological point of view. The differences in karyotypes and seed protein profiles between species and among populations could be the result of the differentiation that they are undergoing at present due to geographical isolation.     

Phylogenetic relationships of Salvia (Lamiaceae) in China: Evidence from DNA sequence datasets

With 84 native species, China is a center of distribution of the genus Salvia (Lamiaceae). These species are mainly distributed in Yunnan and Sichuan provinces (southwestern China), notably the Hengduan Mountain region. Traditionally, the Chinese Salvia has been classified into four subgenera, Salvia, Sclarea, Jungia, and Allagospadonopsis. We tested this classification using molecular phylogenetic analysis of 43 species of Salvia from China, six from Japan, and four introduced species. The nuclear ribosomal internal transcribed spacer region and three chloroplast regions (rbcL, matK, and trnH-psbA) were analyzed by maximum parsimony, maximum likelihood, and Bayesian methods. Our results showed that the Chinese (except Salvia deserta) and Japanese Salvia species formed a well-supported clade; S. deserta from Xinjiang grouped with Salvia officinalis of Europe. In addition, all introduced Salvia species in China were relatively distantly related to the native Chinese Salvia. Our results differed from the subgeneric and section classifications in Flora Reipublicae Popularis Sinicae. We suggested that sections Eusphace and Pleiphace should be united in a new subgenus and that sect. Notiosphace should be removed from subg. Sclarea and form a new subgenus. Our data could not distinguish a boundary between subg. Allagospadonopsis and sect. Drymosphace (subg. Sclarea); the latter should be reduced into the former. Further clarification of the phylogenetic relationships within Salvia and between Salvia and related genera will require broader taxonomic sampling and more molecular markers.     

Nomenclatural notes on the genus Paraphlomis (Lamiaceae: Lamioideae) from China

Lectotypes of Paraphlomis albiflora (Hemsl.) Hand.‐Mazz. and Paraphlomis gracilis (Hemsl.) Kudô are designated. The name Paraphlomis javanica (Blume) Prain var. angustifolia C. Y. Wu & H. W. Li ex C. L. Xiang, E. D. Liu & H. Peng is validated by a type designation.     

Seed morphology and taxonomy of the North African species of Sideritis L. (Lamiaceae)

REJDALI, M., 1990. Seed morphology and taxonomy of the North African species of Sideritis L. (Lamiaceae). Seed morphology data are shown to support data from general morphology and palynology; they can be used for taxonomy at the sectional and specific levels. At times clear differences are apparent at the infraspecific level. Seed sculpturing was found to be of great value for separating taxa at all levels of the hierarchy.     

Pericarp structure of Leucas and related genera (Lamiaceae subfam. Lamioideae)

The pericarp has been studied in 72 taxa belonging to the genera Leucas, Leonotis, Acrotome, Otostegia and Isoleucas. Otostegia, Isoleucas and many Leucas (the Asiatic sections Astrodon, Leucas, Plagiostoma , and the African Spiculifolia, Squarrosicyme, Stachydiformis and Virgata ) have been found to agree with most other members of the subfamily Lamioideae by having high bone cells which contain crystals. Acrotome, Leonotis and most African Leucas (sections Hemistoma, Lasiocotys, Lonostoma, Neuflizeana, Ogadenia and Physoleucas ) deviate by having thin bone cells which often lack crystals, and due to their similarity, these African taxa are suggested to be closely related. The Asiatic Leucas (Ortholeucas), Plagiostoma and Astrodon also seem to form a group. Leucas milanjiana and perhaps also Leucas hyssopifolia are considered to be extraneous to Plagiostoma and to be more closely related to the section Leucas and the African Lasiocorys, Ogadenia and Physoleucas respectively.     

The evolution of growth-forms in the Macaronesian genus Aeonium (Crassulaceae) inferred from chloroplast DNA RFLPs and morphology

Phylogenetic relationships among species of Aeonium were studied using morphological characters and chloroplast DNA RFLPs. Cladistic analysis of weighted morphological data indicates that the small, herbaceous and least woody species are basal in the genus. Chloroplast DNA data gave similar results, supporting the separation of the herbaceous or small, woody species from the large, hapaxanth rosettes, rosette trees, and branched subshrubs with yellow, white or red flowers as well as the only (herbaceous) species with axillary inflorescences. The relationships among the species descending from a polytomy that comprises all species of the genus as well as a polytomy which comprises 18 of the 26 species studied, are only very incompletely resolved, except for two monophyletic clades that contain the branched subshrubs with yellow flowers ( A . sect. Aeonium ) and the branched subshrubs and rosette trees with white or red flowers ( A . sect. Leuconium ), respectively. Cladistic analysis of the combined morphological and chloroplast DNA data improved resolution considerably. Four monophyletic clades are distinguished, each of which, except for three species, comprises only one of the five main growth-form types. Although Aeonium is generally regarded as an outstanding example of adaptive radiation, this mode of speciation seems to have been of minor significance in the evolution of the genus, because each growth-form apparently evolved only once. Instead, island speciation in the absence of major ecological shifts, is probably more important in the evolution of the genus.     

Systematic position of Asteropyrum (Ranunculaceae) inferred from chloroplast and nuclear sequences

Sequences of chloroplast gene rbcL and partial nuclear 26S rDNA were used to evaluate phylogenetic relationships of Asteropyrum. Four primary clades were recognized in Ranunculaceae, corresponding to subfamilies Hydrastidoideae, Coptidoideae, Thalictroideae, and Ranunculoideae. Our results place Asteropyrum in Ranunculoideae, sister to the tribe Actaeeae, which includes Beesia, Cimicifuga, and Eranthis. This is supported by chromosome characters, including chromosome size and basic number, and the stainability of prophase chromosomes and interphase nuclei. Our results do not support previous placements of Asteropyrum in either Coptidoideae or Thalictroideae. Considering its uniqueness in a few characters (e.g. simple peltate leaves, accumulating benzylisoquinoline alkaloids, vessel elements with only typical scalariform perforation plates), we recognize Asteropyrum as a monotypic tribe of Ranunculoideae, Asteropyreae W. T. Wang et C. Y. Chang.     

Phylogeny and evolution of Dyckia (Bromeliaceae) inferred from chloroplast and nuclear sequences

The genus Dyckia (Bromeliaceae) comprises more than 150 terrestrial or epilithic species with a strongly xeromorphic habit. Most of its members belong to the azonal rock vegetation of Neotropical savannas and forests of Brazil and adjacent countries. Dyckia is relatively species-rich compared with its closest relatives Encholirium (27 species) and Deuterocohnia (17 species). Here, we present the first molecular phylogenetic analysis of Dyckia using DNA sequence data from six plastid loci (matK gene, rps16 intron, petD intron, rpl32-trnL, rps16-trnK and trnD-trnT) and a portion of the nuclear gene phyC. A total of 124 accessions were included, corresponding to 79 taxa from six genera. Phylogenetic trees were generated using parsimony, likelihood and Bayesian methods. DNA sequence variation among Dyckia species turned out to be extremely low, and phylogenies were poorly resolved. The monophyly of Dyckia is supported, whereas evidence is provided that Encholirium is paraphyletic. Based on a dated plastid DNA tree, Dyckia experienced a recent radiation starting around 2.9 million years ago. Four major clades could be identified that roughly correspond to the geographic origin of the samples. A parsimony network based on plastid DNA haplotypes shows a star-like pattern, indicating recent range expansions. Our data are compatible with a scenario where Dyckia and Encholirium diverged in northeastern Brazil, whereas one lineage of Dyckia dispersed to southern Brazil from where a rapid colonization of suitable habitats was initiated. We discuss our results in relation to species delimitation in Dyckia.