Article

Circumscription and Phylogeny of the Campanulales

January 1992
Annals of the Missouri Botanical Garden 79(2):388

January 1992
79(2):388

Authors:

University of Wisconsin - Oshkosh

Systematic relationships of the Campanulales were reexamined through the integration of data accumulated from morphological, anatomical, embryological, palynological, cytological, ultrastructural, chemical, molecular, and paleontological studies. Interpretation of these data suggests that the most natural circumscription of the order may be achieved by retaining Asteraceae and Calyceraceae, adding Menyanthaceae, removing Donatiaceae and Stylidiaceae to Ericales, and including Brunonia in Goodeniaceae rather than its own family. Phylogenies based on chloroplast DNA restriction fragment analysis and rbcL sequencing, supported by embryological and chemical data, suggest a basal dichotomy into two clades. The first (Menyanthaceae, Goodeniaceae, Calyceraceae, and Asteraceae) has multinucleate tapetal cells, lacks endosperm haustoria, and produces deterrent chemicals (either seco-iridoids or sesquiterpene lactones, but not both) via the mevalonate pathway. The second (Campanulaceae, Cyphiaceae, Lobeliaceae, Sphenocleaceae, and Pentaphragmataceae) has binucleate tapetal cells and terminal endosperm haustoria, but cannot synthesize deterrent chemicals via the mevalonate pathway. Numerous characteristic morphological features (e.g., epigyny, zygomorphy, secondary pollen presentation, uniovulate ovaries) appear to have had multiple origins within the order, vitiating their use as synapomorphies. The order originated no later than the Oligocene, very near the base of the Asteridae, probably in the Cornales-Saxifragales complex.

Interfamilial Relationships of the Asteraceae: Insights from rbcL Sequence Variation

Article

Full-text available

Jan 1993

Nucleotide sequences of the chloroplast gene rbcL were analyzed to examine relationships among the large, distinctive family Asteraceae and eight putatively closely related families. Phylogenetic analysis of a total of 24 sequences of rbcL identified a lineage consisting of two families, the Goodeniaceae and Calyceraceae, as the sister group to the Asteraceae. In addition, a strongly supported major monophyletic clade consisting of Asteraceae, Goodeniaceae, Calyceraceae, Corokia (Cornaceae sensu Cronquist), Menyanthaceae, Lobeliaceae, and Campanulaceae was found. These results clearly distance from the Asteraceae certain groups previously considered closely related; moreover, the results support alternative hypotheses of affinity that were based upon floral and inflorescence morphology, biogeography, pollen morphology, chemistry, and pollen-presentation mechanisms.

Gynoecial Structure of Vitales and Implications for the Evolution of Placentation in the Rosids

Article

Full-text available

Dec 2014

• Premise of the Research: Evolutionary relationships of the Vitales (the economically important Vitaceae and its sister family Leeaceae) within the rosids have been difficult to resolve. Gynoecial structure, especially the placentation type, of the two families has been variously interpreted. A survey of gynoecial structure is undertaken within these two families, and they are interpreted in light of phylogenetic comparison of placentation types among all angiosperms. • Methodology: Gynoecial structure and architecture in 21 species were studied with light and scanning electron microscopy. Ancestral character reconstruction of gynoecia with axile, parietal, basal, apical, free-central, marginal, or laminar placentae across 640 genera representing all 58 orders of angiosperms was inferred using maximum likelihood to help interpret the evolution of the gynoecium in the Vitales. • Pivotal Results: The syncarpous ovary is bicarpellate in Vitaceae and tricarpellate in Leeaceae; in both the carpels are congenitally fused to form the synascidiate zone. Placentae are located basally on the septum with generally two ovules per carpel. Distally, septa are incomplete, resulting in the ovary being incompletely bilocular in Vitaceae and incompletely trilocular in Leeaceae. Further morphological variations are a result of differential growth of the septa. In some species of Cyphostemma septa are further reduced, and the ovary is clearly unilocular. Ancestral character reconstruction using maximum likelihood across 640 angiosperm taxa infers marginal placentation to be the ancestral condition, while that of the Superrosidae is axile, with basal placentation inferred to be derived within the clade. • Conclusions: The apically incompletely septate gynoecia and basally septate gynoecia of Vitaceae and Leeaceae are also found in 31 other angiosperm families. These results underscore the labile nature of placentation despite its long use as a character of taxonomic significance. The study provides a basis for inferring the directionality of placentation evolution in the context of a wider investigation of rosid relationships.

Patterns of Embryological and Biochemical Evolution in the Asterids

Article

Full-text available

Apr 2001

The Monopetalae or Sympetalae have been resurrected recently as the Asteridae s.l., a large clade that includes almost all sympetalous angiosperms. Phylogenetic analyses of a large, combined data set of sequences of four genes (18S rDNA, rbcL, ndhF, atpB) have improved our understanding of the evolutionary history of this large clade of angiosperms. Most of the taxa fall within one of four major clades, the Cornales, Ericales, euasterids I (comprising Lamiales, Solanales, Boraginaceae, Gentianales, and Garryales), and euasterids II (comprising Asterales, Apiales, Dipsacales, Escalloniales, and Aquifoliales). We used the phylogenetic tree based on our recent phylogenetic analysis of four genes to elucidate the evolutionary patterns of the ovule and the diversification of iridoid biosynthesis in the asterids. Asterid-specific patterns of character distribution can be seen. Unitegmic-tenuinucellate ovules occur almost universally in the asterid clade. Cellular endosperm is ancestral in the asterids; nuclear endosperm emerged several times. Shedding of tricellular pollen does not seem to be a useful character for higher level taxonomy except for uniting Apiales and Dipsacales. Reversals from nuclear endosperm formation and tricellular pollen are not observed. Iridoids occur in many asterid clades and are probably ancestral. The ability to produce iridoids, however, has been lost frequently. Secoiridoids predominate in the euasterids II, whereas carbocyclic iridoids are most diverse in Lamiales. Although this study showed high correlation of unitegmic-tenuinucellate ovules with iridoid synthesis, several questions remain. Studies of the ontogeny and biosynthesis of these characters can now focus especially on those areas of transition determined in this analysis. Communicating Editor: James R. Manhart

Phylogenetic Analysis of Asterids Based on Sequences of Four Genes

Article

Full-text available

Apr 2001

Asterids, comprising about one-third of all angiosperm species, include almost all species that produce iridoids and that have sympetalous corollas, and most species that have unitegmic-tenuinucellate ovules. To elucidate their phylogeny, we used separate and combined data sets of sequences from 18S rDNA, rbcL, ndhF, and atpB, with a maximum of 7168 base pairs for 158 genera, for phylogenetic analysis. The analyses resolved the major lineages of asterids s.l., confirming the feasibility of analyses of large data sets in phylogenetic systematies. Most of the taxa fall within one of four major clades. Cornales, comprising Cornaceae, Nyssaceae, Hydrangeaceae. Loasaceae, and Hydrostachyaceae, are sister to the remaining genera. Ericales consist of taxa traditionally included in Ericalcs. Primulales, Ebenales, and Theales, as well as Polemoniaceae and Balsaminaceae. They are sister to euasterids I and euasterids II. The euasterids I include Lamiales, Boraginaceae, Solanales, and Gentianales. The relationship of Boraginaceae as sister to Lamiales, even though weakly supported, is reported for the first time. The euasterids II comprise Asterales, Apiales, Dipsacales, Escallonia, Eremosyne, and Berzelia, and Aquifoliales. Dipsacales, including Sambucus and Viburnum, are monophyletic and sister to Apiales. The data set of 158 taxa sequenced for four genes gives us insight into both the evolution of the asterids and the molecular evolution of the four genes. Some striking differences in rates and patterns of molecular evolution appear both within and among the genes; for example, ndhF consists of two very different regions with the second region apparently evolving under reduced selection pressure. The four genes differ in their base composition, transition-transversion bias, and in their most common substitutions.

Secondary Pollen Presentation in Angiosperms and Its Biological Significance

Article

Full-text available

Jan 1993

Secondary pollen presentation is the developmental relocation of pollen from the anthers onto another floral organ which then functions as the pollen presenting organ for pollination. Nine different types have been identified in sixteen angiosperm families according to which organ is used for presentation, whether the pollen is exposed or concealed within a structure and how pollen is loaded onto the presenting surface: (1) Enveloping bloom presenters (Araceae); (2) Perianth presenters with exposed pollen presentation (Epacridaceae); (3) Androecial presenters (Santalaceae); (4) Terminal stylar presenters with passive pollen placement and concealed stigmas (Rubiaceae and Proteaceae); (5) Terminal stylar presenters with passive pollen placement and sub-terminal stigmas (Marantaceae and Polygalaceae); (6) Terminal stylar presenters with active pollen placement (Asteraceae, Calyceraceae and Lobeliaceae); (7) Sub-terminal stylar presenters (Campanulaceae, Cannaceae, Fabaceae and Myrtaceae); (8) Exposed stigmatic presenters (Rubiaceae); (9) Indusial stigmatic presenters (Goodeniaceae and Brunoniaceae). Secondary pollen presentation occurs in three monocotyledon and thirteen dicotyledon families. The presentation types appear to have been independently derived indicating that secondary pollen presentation is a character with a selective advantage. In all but the enveloping bloom type of secondary pollen presentation, developmental relocation of pollen requires simultaneous, introrse anther dehiscence and a close association of the presenting organ to the anthers prior to anthesis. The various secondary pollen presentation systems may be modified to promote xenogamy or autogamy and this can even change during anthesis. Most plants which have secondary pollen presentation, display reduced herkogamy within the flower to facilitate pollination. Increased risk of self-pollination due to this may be overcome through dichogamy, herkogamy within inflorescences, dry stigmas, self-incompatibility systems and passive or active control over pollinator behaviour. Enhanced male function of the flowers of secondary pollen presenting plants is also evident through extension of the male phase by the protection, controlled release and precise placement and receipt of pollen. Plants displaying secondary pollen presentation are almost always protandrous.

Molecular Phylogeny and Taxonomic Revision of the Genus Diastatea (Campanulaceae: Lobelioideae)

Article

Full-text available

Mar 2022

Diastatea (Campanulaceae-Lobelioideae) is a genus of small, herbaceous annuals found mostly in Mexico. Differences in the number of recognized species in the historical treatments, and the lack of a phylogeny, suggested that a revision of the genus was necessary. Species boundaries and species level relationships were tested based on a sampling of nearly 50 individuals. Sequences of the chloroplast spacer regions, atpB-rbcL and ndhF-rpl32 , as well as the ITS, were generated and then analyzed using maximum likelihood and Bayesian inference methods. The analyses of the separate datasets produced congruent topologies. Two strongly supported clades of Diastatea are newly recognized as subgenera in the taxonomic treatment. Diastatea Scheidw. subg. Diastatea and Diastatea subg. Angustifolia E.P.Johnson & T.J.Ayers can be distinguished morphologically by leaf shape and pubescence, hypanthium shape, and the presence or absence of a gibbous projection on the ventral side of the corolla. A morphological treatment of the genus corroborated by the molecular analysis revealed one new species, Diastatea aptera E.P.Johnson & T.J.Ayers, one new subspecies, Diastatea tenera subsp. durangensis E.P.Johnson & T.J.Ayers, and elevated an existing subspecies, Diastatea ciliata (McVaugh) E.P.Johnson & T.J.Ayers, to specific rank.

Typification of Linnaean specific names in the genus Campanula (Campanulaceae)

Article

Dec 2019

P. Pablo Ferrer-Gallego

The lectotypification of nine Linnaean names in the genus Campanula (C. decurrens, C. erinoides, C. graminifolia, C. hederacea, C. hybrida, C. linifolia, C. mollis, C. thyrsoides, C. uniflora) is discussed. The designation of the nomenclatural types is based on the consultation of Linnaeus's original material and the literature cited in the respective protologues. The names are lectotypified using specimens from LINN except for C. erinoides, C. graminifolia, and C. hybrida, which are lectotypified by illustrations of Hermann (1687), Barrelier (1714), and Morison (1680), respectively. An epitype is designated for C. graminifolia due to ambiguity in the interpretation of the lectotype.

Pollen and reproductive morphology of Rhigiophyllum and Siphocodo (Campanulaceae): two unique genera of the fynbos vegetation of South Africa

Article

Full-text available

Dec 2010

Pollen grains of Rhigiophyllum squarrosum Hochst., Siphocodon spartioides Turcz. and S. debilis Schltr., are flattened and triangular with pores at the angles. This morphology is radically different from known pollen of the Campanulaceae s.sfr:: the Campanulaceae are treated here as a family separate from the Lobeliaceae, Cyphiaceae, Nemacladaceae, Pentaphragmataceae and Sphenocleaceae (Lammers 1992). As traditionally conceived, the Campanulaceae is very heterogeneous and, in many classifications, these families were treated as subfamilies of a much-enlarged Campanulaceae. The consistently different floral morphology, biochemistry and pollen structure of the Lobeliaceae favours the recognition of this predominantly tropical group as a separate family.The pollen grains of these species are described in comparison with other members of the Campanulaceae. Based on surface characteristics of their pollen grains, we conclude that they represent an early offshoot o f the wahlenbergioid lineage in southern Africa. We suggest that this unique pollen may also be the result of a highly selective regime in the fynbos, associated with specialized pollinators, and base-poor soils, in addition to possible adaptations for ant dispersal and fire. Rhigiophyllum Hochst. and Siphocodon Turcz. are also unique in having free carpel-like structures within the ovary. These shrink to form seed pockets around the seeds and disperse as units when the capsule matures. Data from molecular studies support the contention that these taxa form a sister group to all other wahlenbergioids and that this should be formally recognized in a classification system. We treat Rhigiophyllum and Siphocodon within the Campanulaceae: Wahlenbergioideae, as a separate tribe, the Rhigiophylleae tribus nov., the species of which are distinguishable from other wahlenbergioids by unique angulaperturatc pollen, epipetalous stamens, free carpel-like structures and seed pockets.

‘False resupinatioń in the flower-pairs of Thalia (Marantaceae)

Article

Sep 2015
FLORA

Resupination, i.e., the turn by 180°, arranges zygomorphic flowers in a suitable position for pollination. It is well-known from orchids, but also occurs in further angiosperm families. Within the genus Thalia L. (Marantaceae), T. dealbata has erect and T. geniculata pendulous inflorescences. Nevertheless, in both species the zygomorphic flower-pairs show the right position for pollination. Thus, resupination is expected in Marantaceae. To reconstruct spatial arrangement, flower development was investigated from meristem formation to anthesis. Early morphogenetic stages were analysed under the scanning electron microscope, late stages were documented by time-lapse records. Gravity experiments were added to test for plastic or genetically fixed flower-pair position. Flowers in both Thalia species develop almost identically. They originate from a common primordium and show the unusual floral organ sequence already described for the family. In T. dealbata, the flower-pair gets its final position by bending 90° backwards, while in T. geniculata a lateral rotation of 90° takes place shortly before anthesis. In both species, position is genetically fixed. Results clearly indicate that the flower-pairs in Thalia do not show resupination according to the given definition. Instead of the expected 180° turn in one species, flower-pairs in both species each dislocate for 90°.

Plant reproduction in the alpine landscape : reproductive ecology, genetic diversity and gene flow of the rare monocarpic "Campanula thyrsoides" in the Swiss Alps

Article

Jan 2007

Hafdís Hanna Aegisdóttir

A new species and a new subspecies of Salvia (Lamiaceae) from Jalisco and MIchoacán, Mexico

Article

Full-text available

Dec 2012

A new species and a new subspecies of Salvia, Salvia vazquezii and Salvia vazquezii subsp. tancitaroensis, from the Jalisco-Colima border and Michoacan, Mexico, respectively, are segregated from the S. gravida complex. They are described, illustrated and compared. A key to the three taxa and a translated and expanded circumscription of S. gravida are provided.

Inflorescence and Flower Development of Pentaphragma horsfieldü (Pentaphragmataceae) and its Systematic Implications

Article

Full-text available

Pentaphragma is a peculiar genus of Malesian rain forest herbs, which was formerly often included in Campanulaceae, but is now generally regarded to represent a family of its own. Recent molecular-systematic and morphological-cladistic studies (LAMMERS 1992; COSNER et al. 1994; GUSTAFSSON & BREMER 1996) have confirmed that the family has its place within Asteridae, but its position there is still uncertain. Collection of flowering material in Malaysia enabled the study of the structure and functional aspects of the flower (VOGEL 1998) as well as the inflorescence and flower development. Floral ontogeny seemed interesting as ontogenetic criteria such as 'early' or 'late sympetaly' turned out to be significant taxonomic markers (ERBAR & LEINS 1996). The flowers are in scorpioid cymes (cincinni). The primordia arise in a pseudo-monopodial manner from a 'false apical meristem' like in Boraginaceae.

Campanula rotundifolia: Biological Flora of the British Isles No. 266

Article

Jan 2012
J ECOL

1. This account presents information on all aspects of the biology of Campanula rotundifolia that are relevant to understanding its ecological characteristics and behaviour. The main topics are presented within the standard framework of the Biological Flora of the British Isles: distribution, habitat, communities, responses to biotic factors, responses to environment, structure and physiology, phenology, floral and seed characters, herbivory and disease, history, and conservation. 2. Campanula rotundifolia L. is a slender, rhizomatous, prostrate to erect herb with long-stalked roundish basal leaves, linear stem leaves, and a blue, bell-shaped corolla. It is widespread in the British Isles though absent from parts of Ireland. Globally, C. rotundifolia has a circumpolar distribution extending from the Arctic Circle to northern Mexico and North Africa. It forms a polyploid complex with some characters linked to ploidy level. Populations in the British Isles are predominantly tetraploid and hexaploid, with occasional pentaploids. The distribution of these cytotypes in the British Isles has a strong spatial structure. 3. Campanula rotundifolia has a wide ecological amplitude, tolerating very dry conditions but also occasionally occurring in permanently saturated habitats, and it grows on a very wide range of soil types, from coarse sands through loams to heavy clays and pure peats. However, C. rotundifolia is rarely found on fertile lowland clays where competition from more vigorous species may limit growth. C. rotundifolia is found in a range of grassland, heath, mire, scrub and sand dune communities spanning a wide range of soil pH. 4. Campanula rotundifolia is perennial and spreads by seed and by rhizome. Plants generally overwinter as frost-resistant green rosettes. As the spring season progresses, plants make slow vegetative growth. Erect flowering stems develop from June onwards. C. rotundifolia is largely self-incompatible, and is insect pollinated, regularly producing large quantities of viable seed. 5. Although C. rotundifolia is a widespread and locally very common species there is some evidence from Britain and across Europe that it is in decline. These losses are likely to be due to a number of factors including agricultural intensification, reversion of grassland to scrub and woodland, disturbance, and atmospheric pollution.

Biological Flora of the British Isles: Campanula rotundifolia

Article

May 2012

1. This account presents information on all aspects of the biology of Campanula rotundifolia L. that are relevant to understanding its ecological characteristics and behaviour. The main topics are presented within the standard framework of the Biological Flora of the British Isles : distribution, habitat, communities, responses to biotic factors, responses to environment, structure and physiology, phenology, floral and seed characters, herbivory and disease, history, and conservation. 2. Campanula rotundifolia is a slender, rhizomatous, prostrate to erect herb with long‐stalked roundish basal leaves, linear stem leaves, and a blue, bell‐shaped corolla. It is widespread in the British Isles though absent from parts of Ireland. Globally, C. rotundifolia has a circumpolar distribution extending from the Arctic Circle to northern Mexico and North Africa. It forms a polyploid complex with some characters linked to ploidy level. Populations in the British Isles are predominantly tetraploid and hexaploid, with occasional pentaploids. The distribution of these cytotypes in the British Isles has a strong spatial structure. 3. Campanula rotundifolia has a wide ecological amplitude, tolerating very dry conditions but also occasionally occurring in permanently saturated habitats, and it grows on a very wide range of soil types, from coarse sands through loams to heavy clays and pure peats. However, C. rotundifolia is rarely found on fertile lowland clays where competition from more vigorous species may limit growth. Campanula rotundifolia is found in a range of grassland, heath, mire, scrub and sand dune communities spanning a wide range of soil pH. 4. Campanula rotundifolia is perennial and spreads by seed and by rhizome. Plants generally overwinter as frost‐resistant green rosettes. As the spring season progresses, plants make slow vegetative growth. Erect flowering stems develop from June onwards. Campanula rotundifolia is largely self‐incompatible, and is insect pollinated, regularly producing large quantities of viable seed. 5. Although C. rotundifolia is a widespread and locally very common species there is some evidence from Britain and across Europe that it is in decline. These losses are likely to be due to a number of factors including agricultural intensification, reversion of grassland to scrub and woodland, disturbance, and atmospheric pollution.

Evolution of the platycodonoid group (Campanulaceae s. str.) with particular references to biogeography and character evolution

Article

Apr 2014
J INTEGR PLANT BIOL

The Qinghai-Tibet Plateau (QTP) is an important centre of origin and diversification for many northern temperate plants. The hypothesis ‘out-of-QTP’ suggests that the majority of northern temperate plants have originated and dispersed from the QTP and adjacent regions. An interesting question is whether the biogeographic history of the platycodonoids, a group mainly distributed in the QTP and adjacent regions, coincides with the hypothesis ‘out-of-QTP’? Furthermore, how have the diagnostic characters of the platycodonoids evolved? In the present study, all 10 genera of the platycodonoids were sampled for molecular phylogeny and dating analyses, and ancestral states of distribution and characters were reconstructed. The results do not support the platycodonoids as an‘out-of-QTP’ group, but instead they might have descended from Tethyan ancestors. The dispersal and diversification of the platycodonoids in Asia might have been driven by the uplift of the QTP. The present study highlights the importance of the Tethyan Tertiary flora for the origin of the Sino-Himalayan flora and the influence of the uplift of QTP on diversification of northern temperate plants. In addition, character reconstruction reveals that the inferior ovary, capsule, long colpate pollen, and chromosome number 2n = 14 are probably ancestral states.

A Reconsideration of Three-Item Analysis, the Use of Implied Weights in Cladistics, and a Practical Application in Gentianaceae

Thesis

Full-text available

Jan 1997

Jan De Laet

[Introduction, pp. xiii-xv] The central theme of this thesis is cladistics. This approach to phylogenetic analysis has its roots in Willi Hennig’s theoretical work of the fifties (see chapter 1), and after a modest take-off in the sixties and a period of exponential growth in the seventies and the eighties, cladistics has now become a basic tool in systematic research. Its merits are that it has stimulated the development of a conceptual framework that enables us to think and talk in a clear way about phylogenetic relationships, and that it provides a set of powerful methods to analyze systematic data in order to discover the underlying phylogenetic relationships. In the first chapter, a basic survey of the main concepts and terms of cladistics is given. It is presented in Dutch because to date no general introductions to cladistics exist in Dutch. At present, there is a set of generally accepted methods in cladistics. However, this does not mean that the theoretical work has come to an end. To the contrary, old ideas are constantly being refined and new ideas keep popping up. Two of those are treated in the second and the third chapter. The first one is three-item analysis, a method that was introduced some years ago as a novel approach to parsimony analysis in both biogeography (Nelson & Ladiges 1991a, b) and systematics (Nelson & Platnick 1991). The name three-item analysis refers to the fact that each statement about relationships between more than three items (areas in biogeography, homologous features in systematics) is decomposed into a series of basic statements, each of which involves only three items. Such a basic statement simply says which two of the three items are thought to be related more closely to each other than either is related to the third. Following its introduction, three-item analysis has been severely criticized because of three basic defects: (1) it is flawed because it presupposes that character evolution is irreversible; (2) it is flawed because basic statements that are not logically independent are treated as if they are; (3) it is flawed because some of the three-item statements that are considered as independent support for a given tree may be mutually exclusive on that tree. In the second chapter it is shown that these criticisms only relate to the particular way that the approach was implemented by Nelson & Platnick (1991), and an alternative implementation that solves each of the three basic problems is derived. However, the resulting method is not an improvement over standard parsimony analysis: it is identical to the standard approach but for one small constraint, which is a highly unnatural restriction on the maximum amount of homoplasy that may be concentrated in a single character state. As this restriction follows directly from the decomposition of character state distributions into basic statements, it is concluded that any approach that is based on such decompositions will be defective. The second one is about character weighting. Some years ago, Goloboff (1993a) proposed a non-iterative homoplasy-based weighting method in which the weight or fit of a character on a cladogram is defined as a hyperbolic decreasing function of its homoplasy. The best trees are those that have the highest total fit over all characters of a data set. Goloboff considered his approach to be in direct agreement with cladistic ideas, but most parsimonious trees are those trees that imply the lowest amount of weighted homoplasy (Farris 1983), and these are not necessarily the trees that imply that the characters have the highest total fit, as is shown in chapter three. Several implications of this observation are discussed, and an alternative way of weighting characters is proposed. A computer program in which this approach is available is discussed in appendix A, and the approach is illustrated by using an indecisive data set (see chapter six) and the morphological Gentianaceae data set that is presented in chapter five. Several cladistic analyses based on various types of data indicate that the Gentianaceae, a cosmoplolitan family of medium size, is one of the principal families of a monophyletic order Gentianales. Recent developments concerning the order Gentianales are reviewed against a historical background in chapter four. While a consensus is emerging about the monophyly of the Gentianales, much work remains to be done concerning the interfamilial and intrafamilial relationships within the order. The most recent worldwide monograph of the Gentianaceae is over a century old (Gilg 1895). The 21 genera that are selected for the current analysis represent all Gilg’s tribes and subtribes except Leiphaimeae, Rusbyantheae and Voyrieae. Standard parsimony analyses and analyses using Goloboff’s approach of maximum fit give congruent results as far as the global relationships are concerned. The best supported clade contains Eustoma (Tachiinae) and all included Gentianinae, Erythraeinae and Chironiinae. The basal parts of the cladograms, involving the woody tropical representatives and Exacum, are poorly resolved. This thesis is concluded with a short chapter on indecisive data sets. Goloboff (1991a, b) defined the cladistic decisiveness of a data set as the degree to which all possible resolved trees for the data set differ in length. He proposed a measure of the decisiveness of data sets, the DD statistic, and discussed some properties of indecisive data sets, a special type of data set for which every possible cladogram has the same length. His discussion of indecisive data sets was restricted to characters that have no missing entries. In this chapter I will first show how indecisive data sets can be constructed when missing entries are present. Without missing entries, there is essentially only a single indecisive data set for a given number of taxa, but by allowing missing entries a wide variety of different indecisive data sets with a wide range of ensemble consistency and retention indices can be constructed (easy-to-calculate formulas for the length of an indecisive data set on a dichotomous tree and on an unresolved bush are derived in Appendix C). Such data sets are useful in the construction of hypothetical examples that illustrate the elusive nature of data decisiveness. It is concluded that simple measures such as Goloboff’s DD statistic are unable to capture the various aspects of the concept.

Phylogeny of Southern African and Australasian Wahlenbergioids (Campanulaceae) based on ITS and tnL-F Sequence Data: Implications for a Reclassification

Data

Full-text available

Jan 2013

A Preliminary Phylogeny of Alloplectus (Gesneriaceae): Implications for the Evolution of Flower Resupination

Article

Full-text available

Jun 2003

Monophyly of the neotropical plant genus Alloplectus (Gesneriaceae) was tested using maximum parsimony and maximum likelihood phylogenetic analyses of molecular sequence data from the nuclear ribosomal (nrDNA) internal transcribed spacer region (ITS). As currently circumscribed, Alloplectus is polyphyletic and includes taxa in three different clades. The clade that contains the type species is described as Alloplectus sensu stricto and is characterized by the presence of resupinate flowers. The Alloplectus s.s. clade is weakly supported as the sister-group to Columnea. A separate clade of non- resupinate ''Alloplectus'' species nests within a paraphyletic Drymonia. A third taxon, Alloplectus cristatus, endemic to the Lesser Antilles and northwestern South America, is also resupinate and unresolved in a basal polytomy, removed from the other species of Alloplectus. The fourth taxon, Alloplectus peruvianus, which was originally described as Columnea peruviana ,i s strongly supported as nesting within Columnea. Resupination of flowers is an important feature that has not been previously reported and should be recognized as a morphological synapomorphy for Alloplectus s.s. Within the tribe Episcieae, flower resupination is a convergent feature that is independently derived in the Alloplectus s.s. clade, Alloplectus cristatus, and a clade comprising some Nematanthus species.

The Pollen Box in Cyphiaceae (Campanulales)

Article

Full-text available

Sep 2003

In the flower of the monogeneric family Cyphiaceae, distributed in tropical and southern Africa, a pollen box is formed by the five connivent anthers and the headlike style tip as bottom. Pollen with abundant pollenkitt is released into this box just before anthesis. The flowers either have a salverform shape or are bilabiate with three petals forming the upper lip; transitions between these flower types can also be observed. On the flower biology, we only can speculate the following: ( 1) Flowers with the pollen box represent the ( perhaps longlasting) male phase of anthesis. ( 2) In the female phase, the anthers are spreading and the mouth of the stylar channel opens presumably by disintegration of closing hairs. A mucilaginous secretion coming out of the mouth may receive the pollen from pollinators. ( 3) Pollinators of the salverform flowers are presumably butterflies, whereas pollinators of bilabiate flowers are bees. ( 4) Pollen is presumably applied at the proboscis of the butterflies that comes in contact with the pollen mass between the anthers. If pollinators are bees, pollen may be loaded on the ventral side of the insect ( sternotribic pollen loading). The insects may press the pollen box down - with flower orientation more or less horizontal - so that pollen comes out of the pollen box between the upper two anthers. The less elaborate pollen presentation in Cyphiaceae is in agreement with a basal position of the family within the Campanulales as indicated by recent molecular studies.

The unusual occurrence of tricolpate pollen within Mutisieae (Asteraceae)

Article

Full-text available

Jun 2005
GRANA

The monotypic genus Hecastocleis and species of Ainsliaea DC. (Asteraceae, Mutisieae) have tricolpate pollen, which is unusual in Asteraceae. The pollen morphology of H. shockleyi and species of Ainsliaea were studied with light microscope and scanning electron microscope. Pollen of H. shockleyi and some species of Ainsliaea have tricolpate aperture but differ in size and exine features. The pollen of Ainsliaea is bigger than that of Hecastocleis , and has more conspicuous microspines, a Mutisia - or an Ainsliaea - exine type, and is slightly thickened at the poles. The pollen of H. shockleyi is scabrate-microechinate with small puncta, and has the Mutisia -exine type, which is regularly thickened over the complete grain. The occurrence of tricolpate pollen supports previous studies that Hecastocleis and Ainsliaea stand apart from other genera of Mutisieae, but the occurrence of Mutisia -exine type does not suggest complete independence of the tribe.

Phylogeny of Southern African and Australasian Wahlenbergioids (Campanulaceae) based on ITS and trnL-F Sequence Data: Implications for a Reclassification

Article

Full-text available

Jun 2013
SYST BOT

The Campanulaceae: Wahlenbergioideae currently comprises 15 genera, one of which, Wahlenbergia, is widespread over the southern continents. Southern Africa is the region with maximum wahlenbergioid diversity with 12 genera and approximately 252 species. A second center is Australasia with 38 Wahlenbergia species. This study used a broad sample of wahlenbergioid diversity from South Africa, Australia, and New Zealand to reconstruct a phylogeny based on chloroplast trnL-F and nuclear ITS sequences. Data were analyzed separately and in combination using parsimony and Bayesian methods. The results suggest that for the wahlenbergioids to be monophyletic Wahlenbergia hederacea has to be excluded and that none of the South African, Australian or New Zealand lineages are strictly monophyletic. There are five species assemblages that are in some disagreement with current classification in the family. Wahlenbergia, Prismatocarpus and Roella are shown to be non-monophyletic and implications for a reclassification are presented. Careful consideration of morphological characters is suggested before the adjustment of generic circumscriptions can be accomplished.

Survival in the alpine landscape Genetic, demographic and reproductive strategies of the rare monocarpic perennial Campanula thyrsoides in the Swiss Alps

Article

Jan 2006

Patrick Kuss

Floral Morphological Studies in the South African Cyphia stenopetala Diels (Cyphiaceae)

Article

Full-text available

Mar 2005

Cyphia stenopetala is a member of the monogeneric South and tropical African family Cyphiaceae, which is characterized by a pollen-presenting box formed by the five (empty) anthers as walls and the stylar tip as the bottom. Cyphia stenopetala uses the pollen box for self-fertilization. Pollen is shed into the pollen box, where it germinates, and the pollen tubes form a dense felt. We assume that the pollen box in C. stenopetala is an area for an increased pollen tube competition presumably compensating for possible inbreeding depression. Flowers of outbreeding Cyphia species are distinctly proterandrous, but in the selfing C. stenopetala, flowers are proterogynous. A mucilaginous liquid secreted from the enlarged stylar channel at the top of the style may cause pollen germination. Self-fertilization in C. stenopetala appears to be a recent evolutionary event because all flowers investigated produce nectar, and different populations can differ in time and degree of pollen germination.

Stipules in angiosperms

Article

Jan 2024

Stipules are generally regarded as the outgrowths of the leaf base in angiosperms. Other interpretations see stipules as independent organs comparable to leaves. Stipules have been recognized as an important trait for plant taxonomy and identification, and there has been great progress in the understanding of their morphology, development, origin, function, and gene regulation over time. Therefore, this review will briefly summarize past research and aims to clarify the occurrence, location, and morphology of stipules in the families recognized by APG IV and reconstruct their ancestral states. Additionally, the developmental morphology of different types of stipule is presented through scanning electron microcopy observations and a survey of the existing literature. The difference between stipules and ligules is discussed in relation to the occurrence of postgenitally or congenitally fused ‘continuous’ stipules. A distinction is made between ‘true’ stipules and ‘pseudostipules’. The origin of stipules at the base of the leaf is explained from different perspectives. About one-third of the families are reported to have stipules, mostly concentrated in the Rosid clade with the highest level of diversity. On the basis of the ancestral state reconstructions, stipules may be absent in the ancestors of angiosperms, but are present in the ancestor of Rosids, with a pair appearing on both sides of the petiole base. The transition between paired and annular stipules is discussed, the latter arising postgenitally or congenitally. Several hypotheses are discussed to explain the abundance of stipules in the Rosid clade and their limited presence in the Asterid clade.

Variation in Chloroplast Genome Size: Biological Phenomena and Technological Artifacts

Article

Full-text available

Jan 2023

The development of bioinformatic solutions is guided by biological knowledge of the subject. In some cases, we use unambiguous biological models, while in others we rely on assumptions. A commonly used assumption for genomes is that related species have similar genome sequences. This is even more obvious in the case of chloroplast genomes due to their slow evolution. We investigated whether the lengths of complete chloroplast sequences are closely related to the taxonomic proximity of the species. The study was performed using all available RefSeq sequences from the asterid and rosid clades. In general, chloroplast length distributions are narrow at both the family and genus levels. In addition, clear biological explanations have already been reported for families and genera that exhibit particularly wide distributions. The main factors responsible for the length variations are parasitic life forms, IR loss, IR expansions and contractions, and polyphyly. However, the presence of outliers in the distribution at the genus level is a strong indication of possible inaccuracies in sequence assembly.

CHROMOSOME NUMBERS OF CAMPANULACEAE. III. REVIEW AND INTEGRATION OF DATA FOR SUBFAMILY LOBELIOIDEAE

Article

Jun 1993

Thomas G. Lammers

Chromosome numbers are now known for 153 species in 21 genera of Lobelioideae (Campanulaceae); this represents almost 13% of the species and 70% of the genera in the subfamily. Numbers reported are n = 6, 7, 8, 9, 10, 11, 12, 13, 14, 19, 21, 35, 70. The subfamily as a whole has x = 7; the best documented exception is Downingia and its allies with x = 11. Only four genera show interspecific variation in chromosome number: Downingia (n = 6, 8, 9, 10, 11, 12); Lobelia (n = 6, 7, 9, 12, 13, 14, 19, 21); Pralia (n = 6, 7, 13, 14, 21, 35, 70); and Solenopsis (n = 11, 14). Intraspecific variation occurs in 13 species, with as many as four different cytotypes in one species. The herbaceous members of the subfamily as a group are quite variable, showing the entire range of chromosome numbers, including numerous dysploids, but are predominantly diploid. The woody species, by contrast, are much less variable; nearly all of the species are tetraploid, with only a few diploids and hexaploids and no dysploid numbers known. These data support the hypothesis that woodiness is apomorphic within the subfamily. A general trend of higher chromosome numbers at higher latitudes and higher elevations is evident within the subfamily. The chromosome number of Apetahia raiateensis (n = 14) is reported here for the first time, on the basis of a count made about 30 years ago by Peter Raven.

Chromosome Numbers Of The East African Giant Senecios And Giant Lobelias And Their Evolutionary Significance

Article

Jul 1993

The gametophytic chromosome number for the giant senecios (Asteraceae, Senecioneae, Dendrosenecio) is n = 50, and for the giant lobelias (Lobeliaceae, Lobelia subgenus Tupa section Rhynchopetalum) it is n = 14. Previous sporophytic counts are generally verified, but earlier reports for the giant senecios of 2n = 20 and ca. 80, the bases for claims of intraspecific polyploidy, are unsubstantiated. The 14 new counts for the giant senecios and the ten new counts for the giant lobelias are the first gametophytic records for these plants and include the first reports for six and four taxa, respectively, for the two groups. Only five of the 11 species of giant senecio and three of the 21 species of giant lobelia from eastern Africa remain uncounted. Although both groups are polyploid, the former presumably decaploid and the latter more certainly tetraploid, their adaptive radiations involved no further change in chromosome number. The cytological uniformity within each group, while providing circumstantial evidence of monophyly and simplifying interpretations of cladistic analyses, provides neither positive nor negative support for a possible role of polyploidy in evolving the giant-rosette growth-form.

Another perspective on cytoevolution in Lobelioideae (Campanulaceae)

Article

Oct 1996

The Lobelioideae is a cosmopolitan group whose cytoevolution is discussed on a model of primitively high diploid chromosome numbers, in which x = 14 is relatively plesiomorphic and x = 21 may be even more plesiomorphic. This model is suggested from the high frequency of lobelioid genera with x = 14, the probably plesiomorphic condition of x = 17 in the sister group Campanuloideae (Campanulaceae), and the primitive x = 15 in Stylidiaceae (Campanulales). It contrasts with that for a primitive x = 7 and paleopolyploidy to higher chromosome numbers. In our analysis, the genus Lobelia shows three broad cytoevolutionary groups, which probably have phylogenetic and infrageneric taxonomic significance: (1) woody diploids with x = 21 in Chile and woody diploids with x = 14 in Africa, Asia, and Hawaii; (2) herbaceous diploids with several series of dysploid chromosome numbers n = 19, 13, 12, 11, 10, 9, 8, 7, 6, mainly in Africa and Australia; (3) widespread and speciose herbaceous taxa based on a very derived n = 7, with recent frequent euploid rises (neopolyploidy) at or below the species level in subgenus Lobelia and allied or segregate genera. Other woody and herbaceous lobeliad genera have comparable cytoevolutionary patterns. New chromosome counts for Australian Lobelia, Pratia, and Isotoma illustrate the last two cytoevolutionary groups.

Morphology and phylogenetic interrelationships of the Asteraceae, Calyceraceae, Campanulaceae, Goodeniaceae, and related families (Asterales )

Article

Feb 1995

In search for the sister group of the Asteraceae, morphological evidence was assembled for investigating the relationship between the Asteraceae and those families most frequently considered to be their closest relatives, in particular the Calyceraceae, Campanulaceae (along with the frequently included Lobeliaceae, Cyphiaceae, Cyphocarpaceae, and Nemacladaceae), and Goodeniaceae (and the sometimes included Brunoniaceae). Several other families that have been associated with this group of families, the “Asterales-Campanulales-complex,” were also considered: Pentaphragmataceae, Sphenocleaceae, Stylidiaceae, Donatiaceae, Menyanthaceae, and Argophyllaceae. In order to delineate the complex more precisely, another eight putatively related families were also included in the analysis. Cladistic parsimony analysis of 46 morphological and chemical characters for the 23 families was undertaken. Stability of the branches was estimated by the number of extra steps necessary to lose the group, as well as by the number of reweighted extra steps (using rescaled consistency indices) necessary to lose the group (a new approach). The results indicate that there is a monophyletic group of 14 families comprising those of the Asterales-Campanulales-complex as well as Pentaphragmataceae, Sphenocleaceae, Stylidiaceae, Donatiaceae, and Menyanthaceae; this group is recognized as the order Asterales. Within the order, the Asteraceae, Calyceraceae, Brunoniaceae, and Goodeniaceae form one comparatively well-supported clade and the five families of the Campanulaceae sensu lato form another well-supported clade.

Portioned pollen release and the syndromes of secondary pollen presentation in the Campanulales-Asterales-complex11)Dedicated in friendship to Prof. Dr. Andreas Sievers on the occasion of his 65th birthday (June 21, 1996))

Article

Full-text available

Nov 1995
FLORA

The release of pollen in portions, i. e. limiting the pollen removed by individual pollinators during a single visit, is a widespread phenomenon. The biological significance of this phenomenon is that it helps to optimize the pollination. Parameters like number of ovules, number of application sites to the pollinator or spectrum of pollinators, which may be correlated with the size of the pollen portion, are illustrated as a net of possible correlations (Fig. 1). Apart from pollenkitt, viscin or cellulosic threads and the non-simultaneous opening of the anthers, the secondary pollen presentation is a special mechanism of pollen portioning. Secondary pollen presentation occurs in several dicotyledonous and in a few monocotyledonous families. Its occurrence is a distinctive feature in the Campanulales-Asterales-complex. The different mechanisms in the Campanulales-Asterales (Asteraceae, Lobeliaceae, Campanulaceae, Goodeniaceae, Brunoniaceae, Calyceraceae) are briefly described. The differentiations of the "basic syndrome" (radial symmetry of the flower bud, complete androecial whorl, introrse anthers, the latter connivent at the time of dehiscence, proterandry, late and sequential growth of filaments and style) are presented in two diagrams (Figs. 30 + 31).

Sphenocleaceae

Chapter

Apr 2016

Thomas G. Lammers

Erect annual glabrous herbs, up to 1.5 m tall. Roots fibrous. Stems fistulose, fleshy or spongy, freely branching. Leaves alternate, simple, exstipulate, petiolate; lamina pinnately veined, margin entire. Inflorescence a dense terminal pedunculate acropetal conico-cylindric spike. Flowers small, tetracyclic, perfect, radially symmetrical, subtended by an inconspicuous spathulate bract and a pair of linear bracteoles. Calyx synsepalous, adnate to the ovary, forming a hypanthium; lobes 5, imbricate, persistent. Corolla sympetalous, radially symmetric, white or pale yellow, urceolate-campanulate, caducous; lobes 5, imbricate. Stamens 5, alternating with the corolla lobes; filaments distinct, adnate to the middle of the corolla tube, slightly dilated at base; anthers tetrasporangiate, dithecal, basifixed, distinct. Nectary 0. Gynoecium syncarpous, 2-locular; ovary half-inferior; ovules numerous; placentae axile, spongy; style simple, short or almost wanting; stigma capitate. Fruit a circumscissile capsule, enclosed by the accrescent calyx lobes. Seeds small, oblong, yellowish brown, shining, numerous, exarillate, testa striate; embryo straight; endosperm very scanty or lacking.

A global perspective on Campanulaceae: Biogeographic, genomic, and floral evolution

Article

Full-text available

Feb 2016

Premise of the study: The Campanulaceae are a diverse clade of flowering plants encompassing more than 2300 species in myriad habitats from tropical rainforests to arctic tundra. A robust, multigene phylogeny, including all major lineages, is presented to provide a broad, evolutionary perspective of this cosmopolitan clade. Methods: We used a phylogenetic framework, in combination with divergence dating, ancestral range estimation, chromosome modeling, and morphological character reconstruction analyses to infer phylogenetic placement and timing of major biogeographic, genomic, and morphological changes in the history of the group and provide insights into the diversification of this clade across six continents. Key results: Ancestral range estimation supports an out-of-Africa diversification following the Cretaceous-Tertiary extinction event. Chromosomal modeling, with corroboration from the distribution of synonymous substitutions among gene duplicates, provides evidence for as many as 20 genome-wide duplication events before large radiations. Morphological reconstructions support the hypothesis that switches in floral symmetry and anther dehiscence were important in the evolution of secondary pollen presentation mechanisms. Conclusions: This study provides a broad, phylogenetic perspective on the evolution of the Campanulaceae clade. The remarkable habitat diversity and cosmopolitan distribution of this lineage appears to be the result of a complex history of genome duplications and numerous long-distance dispersal events. We failed to find evidence for an ancestral polyploidy event for this clade, and our analyses indicate an ancestral base number of nine for the group. This study will serve as a framework for future studies in diverse areas of research in Campanulaceae.

Molecular phylogeny of the platycodonoid group (Campanulaceae s.Str.) with special reference to the circumscription of Codonopsis

Article

Jun 2013

Phylogenetic analyses of 41 taxa representing all genera except Ostrowskia of the platycodonoid group were done using sequences of five DNA fragments from the chloroplast and nuclear genomes. The results show that the platycodonoid group is a monophyletic group, and that four major clades can be recognized: (1) the Platycodon clade with colporate pollen, including Platycodon, Canarina, Cyclocodon, and Echinocodon; (2) the Cyananthus clade with colpate pollen and a superior ovary, including only Cyananthus; (3) the long-colpus clade with long-colpus pollen, an inferior ovary, and a disc or glands, including Codonopsis subg. Codonopsis, Campanumoea, and Leptocodon; and (4) the short-colpus clade with short-colpus pollen, an inferior ovary, and without a disc or glands, including Codonopsis subg. Pseudocodonopsis, subg. Obconicicapsula, and C. purpurea of Codonopsis subg. Codonopsis. These four major clades are strongly supported and well correlated with pollen type and morphology. The results support the separation of Cyclocodon from Codonopsis as a genus. The results also indicate that Campanumoea and Leptocodon should be merged with Codonopsis, whereas all taxa of the short-colpus clade should be separated from Codonopsis.

Compositae

Chapter

Jan 2007

Remarkable nectaries: Structure, ecology, organophyletic perspectives IV. Miscellaneous cases

Article

Jul 1998
FLORA

Stefan Vogel

In the present, final part of a series of contributions on nectaries, further, miscellaneous case histories are presented because of their histological or ecological interest. In the special section, extrafloral and floral nectaries of members of six families are treated. A general survey concludes the paper. Extranuptial glands attracting ant guards are associated with the flowers of Becium and Ocimum (Lamiaceae); they represent novel cases in a family hitherto believed to lack this kind of organs. The glands develop in an unusual way by transformation of abscission scars of floral bracts. The five floral but extranuptial glands on the calyx of Siphocampylus betulifolius (Lobeliaceae), with a similar function, are unusual in sharing the same histological structure and drainage via stomatal sap-holes with the closely adjoining nuptial nectary, a disk. Three case histories refer to zoophilous unisexual flowers with only one of the functional sexual morphs rewarding with nectar. In Pachysandra and Sarcococca (monoecious Buxaceae) the male flowers are nectariferous. The nectarless pistillate flowers are inadvertently pollinated by bees while these exploit the staminate flowers. Two other instances, although heterogeneous in detail, exhibit the reverse situation in that only the pistillate flowers yield nectar: the male capitula of the dioecious composite Heterothalamus alienus are girdled by female-sterile ray florets with copious nectar; bees taking it become dusted with pollen when they move over the dry staminate disk florets. They may subsequently pollinate the female heads, which entirely consist of nectariferous pistillate ray florets. In the hummingbird-pollinated Begonia ferruginea it is also the pistillate flowers that contain a presumably nectareous fluid, while the dry staminate ones fallaciously attract the visitors by mimicking the pistillate flowers. Although basically epigynous, the protandrous revolver flowers of the little-known genus Pentaphragma (Pentaphragmataceae) have maintained the subbasal position of their five separate nectaries on the ovary wall, the nectar being accessible to bees via five receptacular channels. In the concluding chapter, possible evolutionary pathways leading to floral nectaries, particularly floral disks, are discussed. Hydathodes of vegetative provenance rather than extrafloral nectaries are likely precursors of most floral nectaries. Developmental modules comprising a single stomatal sap-hole or groups of them, associated with peristomatal gland cells, may have aggregated in flowers to form disks. Separate nectariferous scales as found in flowers of the Cucurbitaceae, Crassulaceae and the Geraniales also reveal similarities with the respective leaf hydathodes. The organophylesis of the single scale seated in the axial spur of Pelargonium is detailed as an example. A clustered distribution of stomatal sap-holes on disks can be partly interpreted as revealing their origin from meristic hydathodal modules. Examples of the diversity of these patterns are given, which are little explored and may provide useful taxonomic characters.

Viburnales: Cytological Features and a New Circumscription

Article

Feb 2000

Benko‐Iseppon, A. M. & Morawetz, W.: Viburnales: cytological features and a new circumscription. – Taxon 49: 5‐16. 2000. – ISSN 0040‐0262. Cytological studies carried out in 98 species (22 genera out of eight families: Caprifoliaceae, Sambucaceae, Viburnaceae, Adoxaceae, Valerianaceae, Dipsacaceae, Morinaceae , and Calyceraceae) of the Dipsacales s.l. and the eventually related families Cornaceae , and Hydrangeaceae point to a very close relationship between the Viburnaceae, Sambucaceae and Adoxaceae , corroborating the inclusion of these families into an order different from the Dipsacales , namely the Viburnales. Many cytological features shared by these families differ strongly from the Dipsacales s.str., especially (1) chromosome size and morphology, (2) the presence of cold induced chromosome regions (CIRs), (3) interphase nuclear structure, and (4) chromosome condensing behaviour at prophase. Cornaceae and Hydrangeaceae present similar interphase nuclei, but differ from the Viburnales by other karyomorphological characters. The results are discussed with respect to previous morphological, embryological, and molecular findings.

Flavonoid Chemistry of Barnadesioideae (Asteraceae)

Article

Jan 1995

The flavonoid profiles of some species of the following genera of the Barnadesioideae (Asteraceae) consist of mono- and diglycosides of kaempferol and quercetin: Arnaldoa, Barnadesia, Chuquiraga, Dasyphyllum, Doniophyton, Fulcaldea, and Schlechtendalia. These South American genera previously comprised part of the subtribe Barnadesiinae of Mutisieae. Very simple flavonoid profiles were also noted for the Chilean monotypic Gypothamnium and the Hawaiian endemic Hesperomannia arborescens of subtribe Mutisiinae. The simple flavonoid profiles of the barnadesioid genera are taken as further evidence for the primitive status of this subfamily as suggested by the absence of a 22kb cpDNA inversion in this group of genera. Preliminary studies of the flavonoids of the Calyceraceae have shown that the pigment profile of species of Acicarpha and Gamocarpha are very similar to the profile seen in the barnadesioid genera. A brief discussion of relationships between Barnadesioideae and Acicarpha as a possible representative of the prototypical Calyceraceae, indicated by floral development, diploid chromosome number, geography, and flavonoid chemistry, is included.

Phylogeny and Generic Interrelationships of the Stylidiaceae (Asterales), with a Possible Extreme Case of Floral Paedomorphosis

Article

Full-text available

Jul 1998

Cladistic analyses of Stylidiaceae (Asterales), using Donatiaceae as outgroup and with both morphological and molecular characters, produced two equally parsimonious cladograms. The analyses used morphological characters for 26 species and molecular characters from the chloroplast DNA genes rbcL and ndhF for 12 species. The cladograms indicate that Levenhookia and Stylidium are sister groups and that Oreostylidium is nested within Stylidium. The latter result is remarkable because Stylidium has several significant flower specialisations that Oreostylidium lacks and because Oreostylidium is endemic to New Zealand, where Stylidium is missing. The simple flowers of Oreostylidium may have evolved by reduction and paedomorphosis of the zygomorphic and sensitive flowers of a Stylidium-like ancestor, a change caused by adaptation to a new environment lacking a suitable pollinator. In connection with a switch to unspecialised pollinators or self-fertilisation, the flowers of Oreostylidium apparently became fertile at a morphologically immature or reduced stage.

The Impact of Molecular Methods on the Systematics of Angiosperms

Article

Aug 1997
Bot Acta

C. Wagenitz

:A comparison is made between the systematics of selected orders and families based on morphology and other “classical” characters on the one hand, and the results of molecular methods on the other hand. It can be shown that taxa defined by a broad array of characters from morphology, anatomy, embryology and phytochemistry usually are confirmed by molecular results. On the other hand a family like the Saxifragaceae s.l. delimited solely on the basis of floral morphology has been shown to be grossly polyphyletic. Some quite surprising results of the molecular analyses usually agree with some embryological or phytochemical characters, and sometimes even with characters of vegetative morphology and anatomy. As a special case “unequal ancient splits” are discussed, where one clade contains few genera and species usually retaining many primitive characters, and the other shows great diversity and contains the more advanced members.

Nemacladoideae, a New Subfamily of Campanulaceae

Article

Mar 1998

Thomas G. Lammers

Data from morphology and sequencing of the chloroplast gene rbcL show that Cyphioideae (Campanulaceae) as traditionally circumscribed are not monophyletic. To remedy the situation, its genera are divided among three subfamilies, one of which, Nemacladoideae, is new.

A new taxonomic system of the Campanulaceae s.s.: System of Campanulaceae s.s.

Article

Oct 2014

The Campanulaceae s.s. have been proved to be monophyletic, but the subdivision of the family is still controversial among authors. To investigate the intrafamilial structure of the Campanulaceae s.s., four chloroplast DNA fragments, atpB, matK, rbcL, and petD with petB-petD spacer region, were chosen for molecular phylogenetic analysis, and 90 taxa representing 36 genera of this family were sampled. The result shows that the Campanulaceae s.s. consist of three strongly supported monophyletic clades. This result is highly correlated with the data from palynology and external morphology. Therefore, we propose to establish a new three tribal classification system of Campanulaceae s.s. The Cyanantheae is characterized by colpate or colporate pollen with elongate apertures and a loculicidal capsule which is dehiscent by apical values, or a berry. The other two tribes have porate pollen with poroid apertures, but the Campanuleae possesses a poricidal capsule which is dehiscent by lateral pores or valves, or a dry indehiscent fruit, while the Wahlenbergieae possesses a loculicidal capsule which is dehiscent by apical valves, pores or opercula. In the Cyanantheae, we recognize six subtribes and 10 genera. In addition, keys to tribes of Campanulaceae s.s., and to subtribes and genera of the Cyanantheae are here presented respectively.

A New Rare Species of Lobelia (Campanulaceae, Lobelioideae) from Mexican Lowland Rainforest

Article

Full-text available

Jan 2008

Se describe como nueva especie para la ciencia a Lobelia lithophila Senterre & Castillo-Campos. Esta es una especie conocida solamente de la localidad tipo y probablemente corresponde a un taxa microendémico para la selva alta perennifolia de la zona de Uxpanapa al sur de Veracruz, México. La especie aquí descrita esta más relacionada con L. orientalis Rzedowski & Calderón, otro microendémico en este caso de Querétaro, las cuales se destacan por ser especies herbáceas con hojas arrosetadas en el suelo con ovario totalmente súpero y que también son de hábitats muy restringidos. Es fácil distinguir a la especie nueva por sus hojas grandes pero de pecíolos pequeños, y por su pubescencia menos desarrollada. Asimismo, es una especie que se localiza en un hábitat especial, en sustratos rocosos calizos de bajas altitudes, en el sotobosque de la selva alta perennifolia. Esta especie pertenece a la subsección Leiospermae E. Wimmer, sección Lobelia del subgénero Lobelia L.

Sphenocleaceae

Book

Full-text available

Jan 2007

Classification and geography of the flowering plants

Article

Jul 1992

Robert F. Thorne

Thome, Robert F. (Rancho Santa Ana Botanic Garden, Claremont, CA 91711). Classification and geography of the flowering plants. Bot. Rev.58(3): 225–348. 1992.—This treatment of the flowering plants is the latest revision of my classification of the Class Angiospermae and replaces my 1983 and more recent 1992 synopses. An update is necessary because so much new information has been published in the last decade pertinent to the classification of the flowering plants. About 870 such recent books, monographs, and other botanical papers are cited in the Introduction, listed primarily by the botanical discipline that they represent. Also considerable changes in my classification have been necessitated by my narrowed family- and ordinal-gap concepts, acceptance of the ending “-anae” for superorders in place of the traditional but inappropriate “-iflorae,” and acceptance of more prior or more widely used names for the categories above the family. A new phyletic “shrub” replaces earlier versions, and attempts to indicate visually relative sizes and relationships among the superorders, orders, and suborders. One table includes a statistical summary of floweringplant taxa: ca. 233,900 species of 12,650 genera, 437 families, and 708 subfamilies and undivided families in 28 superorders, 71 orders, and 71 suborders of Angiospermae. Three other tables summarize the known indigenous distribution of the families and subfamilies of angiosperms about the world. The synopsis lists the flowering plant taxa from the class down to the subfamily (and in Asteraceae down to the tribe) with indication of the degree of confidence I place in the circumscription and placement of each category above the subfamily, the best available estimates of the number of genera and species for each category, and the known indigenous distribution of each subfamily and family. Table V lists alphabetically the geographical abbreviations used in the synopsis. The extensive bibliography of recent literature should be helpful to those persons interested in the classification of the flowering plants.

Pentaphragmataceae

Article

Jan 2007

Thomas G. Lammers

Another perspective on Cytoevolution in Lobelioideae (Campanulaceae)

Article

Oct 1996

The Lobelioideae is a cosmopolitan group whose cytoevolution is discussed on a model of primitively high diploid chromosome numbers, in which x = 14 is relatively plesiomorphic and x = 21 may be even more plesiomorphic. This model is suggested from the high frequency of lobelioid genera with x = 14, the probably plesiomorphic condition of x = 17 in the sister group Campanuloideae (Campanulaceae). and the primitive x = 15 in Stylidiaceae (Campanulales). It contrasts with that for a primitive x = 7 and paleopolyploidy to higher chromosome numbers. In our analysis, the genus Lobelia shows three broad cytoevolutionary groups, which probably have phylogenetic and infrageneric taxonomic significance: (1) woody diploids with s = 21 in Chile and woody diploids with x = 14 in Africa, Asia, and Hawaii: (2) herbaceous diploids with several series of dysploid chromosome numbers n = 19, 13, 12, 11, 10, 9, 8, 7, 6, mainly in Africa and Australia; (3) widespread and speciose herbaceous taxa based on a very derived n = 7, with recent frequent euploid rises (neopolyploidy) at or below the species level in subgenus Lobelia and allied or segregate genera. Other woody and herbaceous lobeliad genera have comparable cytoevolutionary patterns. New chromosome counts Cor Australian Lobelia, Pratia, and Isotoma illustrate the last two cytoevolutionary groups.

Chromosome Numbers of the East African Giant Senecios and Giant Lobelias and their Evolutionary Significance

Article

Jul 1993

Molecular phylogeny of the Campanula pyramidalis species complex (Campanulaceae) inferred from chloroplast and nuclear non-coding sequences and its taxonomic implications

Article

Full-text available

Jun 2013

The Campanula pyramidalis complex is a group of closely related taxa with a distribution across the Balkans, from the Gulf of Trieste in the north to the Peloponnese Peninsula in the south, with small disjunct parts of the range in the south Apennines. Although 21 taxa were described within this complex, only three, C. pyramidalis, C. versicolor, and C. secundiflora, have been generally accepted in recent synoptical taxonomic treatments. Our molecular phylogenetic analyses based on sequences of three non-coding chloroplast regions (psbA-trnH, psbZ-trnfM, trnG-trnS) as well as of nuclear ribosomal internal transcribed spacers (nrITS), lend strong support to the recognition of several lineages which only partially correspond to generally accepted taxonomic concepts. Molecular data presented in this study showed that C. pyramidalis is a polyphyletic assemblage that segregates into three distinct lineages, one of which is described here as a new species, C. austroadriatica sp. nov. The lectotype of C. pyramidalis, redefined in a strict sense, is designated. Neither C. versicolor nor C. secundiflora were found to be strictly monophyletic, but their monophyly could not be rejected. Morphological and biogeographical implications are discussed.

Chromosome Numbers of Campanulaceae. III. Review and Integration of Data for Subfamily Lobelioideae

Article

Jun 1993

Thomas G. Lammers

Chromosome numbers are now known for 153 species in 21 genera of Lobelioideae (Campanulaceae); this represents almost 13% of the species and 70% of the genera in the subfamily. Numbers reported are n = 6, 7, 8, 9, 10, 11, 12, 13, 14, 19, 21, 35, 70. The subfamily as a whole has x = 7; the best documented exception is Downingia and its allies with x = 11. Only four genera show interspecific variation in chromosome number: Downingia (n = 6, 8, 9, 10, 11, 12); Lobelia (n = 6, 7, 9, 12, 13, 14, 19, 2 1); Pratia (n = 6, 7, 13, 14, 21, 35, 70); and Solenopsis (n = 11, 14). Intraspecific variation occurs in 13 species, with as many as four different cytotypes in one species. The herbaceous members of the subfamily as a group are quite variable, showing the entire range of chromosome numbers, including numerous dysploids, but are predominantly diploid. The woody species, by contrast, are much less variable; nearly all of the species are tetraploid, with only a few diploids and hexaploids and no dysploid numbers known. These data support the hypothesis that woodiness is apomorphic within the subfamily. A general trend of higher chromosome numbers at higher latitudes and higher elevations is evident within the subfamily. The chromosome number of Apetahia raiateensis (n = 14) is reported here for the first time, on the basis of a count made about 30 years ago by Peter Raven.

The classification and geography of the flowering plants: Dicotyledons of the class Angiospermae

Article

Oct 2000

Robert F. Thorne

This latest revision of my classification and geography of the Dicotyledons replaces my 1992 (Bot. Rev. [Lancaster] 58(3): 225–348) review and is necessitated by the plethora of new information that has become available about the classification of the Angiospermae, especially in the currently popular approaches of cladistic, particulate, and molecular taxonomy. This review attempts to bring up-to-date our knowledge of the dicotyledons, with emphasis on new information published in the last decade. Nearly 600 such recent books, monographs, and other botanical articles are cited in the introduction, listed primarily by the botanical discipline they represent, and in the explanation of the classification. More than 2,000 additional works are listed in the “Literature Cited” section. The numerous changes in the classification created by this new information are listed by subclass and superorder, with pertinent references. A new phylogenetic “shrub” replaces earlier versions and attempts to indicate visually relative sizes and relationships among the superorders, orders, and suborders, with all of these divided into 10 subclasses. One table includes a statistical summary of all known and generally accepted flowering-plant taxa: approximately 257,400 species in 13,678 genera, 389 subfamilies in 490 families, and 756 subfamilies and undivided families in 10 subclasses, 31 superorders, 73 orders, and 64 suborders of Angiospermae. Figures for the dicotyledons are 199,500 species in 10,900 genera, 307 subfamilies in 376 families, and 586 subfamilies and undivided families in 7 subclasses, 22 superorders, 49 orders, and 48 suborders. Three other tables summarize the known indigenous distribution of the families and subfamilies of dicotyledons around the world (the monocotyledons are treated elsewhere). The synopsis lists the dicotyledonous taxa from the subclass down to the subfamily (and in Asteraceae down to the tribe), with indications of the degree of confidence I place in the circumscription and placement of each category above the subfamily, the best available estimates of the number of genera and species for each category, and the known indigenous distribution of each subfamily and family. Table V lists the geographical abbreviations used in the synopsis. The extensive bibliography of pertinent literature on which I have based my decisions should be helpful to persons interested in the classification of the dicotyledons.

Seed Morphology in North American Campanulaceae

Article

Full-text available

Jan 1986

Campanulaceae (excluding Lobeliaceae) in North America comprise four genera and 35 species of annual, biennial, and perennial herbs. Generic and specific circumscriptions have been treated variously, and studies of seeds have been few and limited. In this study, seeds of all but one of the native North American species and of selected Eurasian putative relatives were examined with the light and scanning electron microscope. Characteristics of the seeds and their surface cells are described and compared. The seed-coat morphology was found to be relatively uniform, but there are recognizable generic patterns and a number of distinctive individual species. Seeds of Campanula americana, C. divaricata, and the other eastern species of Campanula are each distinctive and do not show the relative uniformity seen within Githopsis, Triodanis, Heterocodon, and the western species of Campanula. Seeds of the recently rediscovered C. robinsiae and of the wide-ranging C. aparinoides are highly distinctive and would appear to set each of these species apart within the genus. However in Triodanis, T. texana stands apart. Seeds of Campanula reverchonii resemble those of Triodanis coloradoensis more than those of other campanulas. Brief speculation on the adaptive significance of seed-coat sculpturing and ornamentation is given.

A Taxonomic Analysis and Revised Classification of Centrospermae

Article

Full-text available

Jul 1984

Clustering and ordination studies demonstrated a phenetically coherent assemblage of 11 core families of Centrospermae, widely separated from Plumbaginaceae and Polygonaceae. Cladistic analyses based on 71 out-group-polarized characters provided strong evidence for monophyly of centrosperms but only weak support for close sister-group status with Plumbaginaceae and Polygonaceae. Both phenetic and cladistic results indicated divergence between Caryophyllaceae, problematically allied with Molluginaceae, and the betalain-producing centrosperms. Three distinct clades emerged among the betalain taxa: a primitive group of Nyctaginaceae + Phytolaccaceae; an intermediate group of Amaranthaceae + Chenopodiaceae; and an advanced group of succulents-Aizoaceae + Cactaceae, Portulacaceae, and Basellaceae + Didiereaceae. We present a revised classification of Centrospermae, emphasizing cladistic relationships of the 11 core families and including several aberrant or little-known genera subordinated and sequenced according to conventions described by Wiley (1981). The new classification is compatible with recent interpretations of floral homologies and with external data from nucleic acid, ultrastructural, and serological comparisons.

A Chloroplast DNA Phylogeny of the Solanaceae: Subfamilial Relationships and Character Evolution

Article

Full-text available

Jan 1992

Phylogenetic relationships among 42 species of Solanaceae representing 12 of the 14 currently recognized tribes were assessed by chloroplast DNA restriction site mapping. Over 1,000 cleavage sites were identified for 10 restriction enzymes and of these, 447 provided information concerning relationships among the included taxa and the outgroup, Ipomoea (Convolvulaceae). The results establish that subfamily Cestroideae is ancestral in the family and is paraphyletic, and that the subfamily Solanoideae is derived from within the Cestroideae and is monophyletic, if it is circumscribed to include Nolana. The tribe Salpiglossideae, characterized by floral zygomorphy and reduction in stamen number, is.probably polyphyletic and hence artificial. An analysis of character evolution in the family suggests that the tribe Nicotianeae retains the most primitive morphological characters of any tribe in the family and helps to explain the disjoint position of members of the tribe in two distinct lineages in the Cestroideae. The chromosome base number x = 12 unites the Solanoideae with the Anthocercideae and Nicotiana. The worldwide distribution of the Solanoideae versus the almost exclusively New World distribution of the Cestroideae argues for a predominantly long-distance dispersal, rather than a vicariance explanation of biogeographic distributions in the family. The morphologically distinctive genus Schizanthus is the earliest diverging lineage in the family. Tribal relationships within the Solanoideae remain poorly resolved and await more detailed study.

Iridoid Compounds, Their Occurrence and Systematic Importance in the Angiosperms

Article

Full-text available

Jan 1975

Naturally occurring iridoid compounds are divided into ten groups on biosynthetic grounds, demonstrated or postulated. lridoid-bearing genera of plants are tabulated according to the iridoids found in them. lridoid compounds are found in thirteen orders within the superorders Hamamelidanae, Cornanae, Gentiananae, Loasanae and Lamianae (sensu DAHLGREN). The mutual relationships of the orders are discussed with regard to the groups of iridoids found, together with other characters. Arguments for a monophyletic origin of these orders are presented, and the traditional "Sympetalae" is rejected as a natural group. The results of an investigation for iridoids in 44 species from 36 families are recorded in an appendix. lridoid glucosides have heen detected or identified for the first time in the families Retziaceae, Dipsacaceae, Calyceraceae, Roridulaceae, Stylidiaceae, Sarraceniaceae and Goodeniaceae. Comments on the systematic position of the last four families are presented.

Index to plant chromosome numbers 1967-1971

Article

Jan 1973

R.J. Moore

Chromosome numbers and cytoevolution in the Goodeniaceae

Article

Jan 1963
J NEW MAT ELECTR SYS

W.J. Peacock

Index to plant chromosome numbers 1996-1997

Article

Jan 2000

Chiropterophilie in der neotropischen Flora. Neue Mitteilungen I

Article

Jan 1968
FLORA

S. Vogel

Systematics of the genus Cyananthus Wall. ex Royle

Article

Jan 1991
ACTA PHYTOTAXON SIN

Studies in the flora of Thailand. 14. Cytological studies in vascular plants of Thailand

Article

Jan 1963

K. Larsen

Principles and Concepts in the Classification of Higher Taxa

Article

Jan 1977

Vernon Heywood

Despite the very great interest shown by botanists during the last 100 years in the “phylogenetic classification” of the Flowering Plants, much more attention seems to have been paid to the phylogenetic components of such schemes (despite the lack of adequate evidence) than to their taxonomic/classificatory components (for which much evidence is available). There is in fact an inbuilt conflict involved in constructing a scheme that attempts to express both the dynamic/historical phylogenetic component and a static horizontal present day classification, which makes most of the conventional phylogenetic tree-like diagrams conceptually suspect. In the Flowering Plants, only the classes and families are widely accepted whereas there are numerous plausible ordinal classifications. The kind of questions we have to ask in the case of higher categories and taxa such as the subclass and order are: are the criteria to be primarily phylogenetic, or taxonomic in the sense of aiming at better circumscriptions of the taxa, greater predictive value, stability, etc.? Only when we have answered these and similar questions can we decide whether we should be aiming at cladogenetic schemes or anagenetic grades or an explicit combination of both.

Index to plant chromosome numbers for 1965

Article

Jan 1967

Chromosome Numbers of Some Grasses from the Southwestern United States and Mexico

Article

May 1980

Stephan L. Hatch

Chloroplast DNA Variation and Plant Phylogeny

Article

Jan 1988

Several features, foremost its conservative mode of evolution, make chloroplast DNA an extremely valuable molecule for phylogenetic studies. Its conservatism is also its only serious drawback, as this can limit the amount of useful DNA variation at the intraspecific level. Comparative restriction site mapping is currently the preferred molecular method for examining interspecific relationships. At this level, one typically encounters less than 5% homoplasy. Furthermore, the predominantly uniparental inheritance of chloroplast DNA provides unique insights into the origin of hybrid and polyploid complexes, as illustrated by examples from the genus Brassica. In many families of angiosperms, such as the Asteraceae and Orchidaceae, restriction site mapping can also be used to determine intergeneric relationships. The greater expense of DNA sequencing makes it most appropriate only at those higher taxonomic levels-above the family level-where restriction site mapping fails. Within angiosperms, the rbcL gene appears to be the chloroplast gene of choice for phylogenetic studies. Twenty-five rbcL sequences have already been accumulated, and several laboratories are making a coordinated effort to sequence this gene widely among angiosperms and gymnosperms. The more conservatively evolving ribosomal RNA genes hold the greatest promise for resolving the deepest branchings of plant evolution and, indeed, have already settled the ultimate question of chloroplast evolution, namely, its endosymbiotic origin. A third approach to extracting phylogenetic information from chloroplast DNA is by analyzing the distribution of major structural rearrangements, such as inversions and the loss or gain of genes and introns. Although such rearrangements are rare relative to point mutations, their great rarity and freedom from homoplasy also make them extremely powerful characters. Examples to be discussed include an inversion defining the most ancient branching in the Asteraceae, rearrangements that mark several major divisions within the Fabaceae, and events that identify the green algal ancestors of land plants.

On the Mechanisms of Secondary Pollen Presentation in the Campanulales-Asterales-Complex 1

Article

Feb 1990
Bot Acta

Families of the Campanulales-Asterales-complex are characterized by special mechanisms for secondary pollen presentation: pump mechanism, brushing mechanism, deposition mechanism, cup mechanism and combination of cup mechanism with brushing mechanism. These different mechanisms are based on three successive events: elongation of the filaments, opening of the anthers, elongation of the style. The diversity of the mechanisms arises through different auxiliary structures such as an indusium, hairs or an anther tube, and the relative rate and time of filament and style growth. An advantage of secondary pollen presentation lies in the prolongation of the male phase of anthesis through portioned pollen release. A probable phylogeny of the different mechanisms of secondary pollen presentation is proposed (Fig. 7).

The Bases of Angiosperm Phylogeny: Embryology

Article

Jan 1975

Barbara F. Palser

Many of the embryological characteristics that may be useful, when employed judiciously and in conjunction with other characters, in arriving at taxonomic conclusions are listed. Several characteristics that show a non-random distribution between families of the Magnoliatae and those of the Liliatae are discussed. Features which are more predominant in the monocotyledons than dicotyledons are: monocotyledonous type of development of the anther wall; amoeboid tapetum; successive cytokinesis of the microspore mother cells to form isobilateral tetrads; helobial endosperm development; and a single cotyledon in the mature embryo. In contrast, those characteristics that are more prevalent among dicot families than monocot families include: basic and dicotyledonous patterns of anther wall formation; simultaneous cytokinesis of microspore mother cells with the formation of tetrahedral tetrads; hemitropous, amphitropous, or circinotropous ovules; ovules with a single integument; an endothelium; Oenothera, Penaea, Peperomia, Plumbago, or Plumbagella types of megagametophyte development; cellular endosperm; and two cotyledons in the mature embryo. In spite of these differences, those characteristics that are most common--occurring in at least 70%, and usually more, of all angiosperms --are evenly distributed between the two classes and afford a strong embryological unity to the angiosperms. These widespread characters include: four microsporangia per anther; differentiated endothecium; two-celled pollen grains; bitegmic anatropous ovules; Polygonum type of megagametophyte development; and nuclear endosperm. Within the two classes distribution of the various characters is not always proportional among the several subclasses and superorders; for example, a unitegmic, tenuinucellate ovule with an endothelium, cellular endosperm with haustoria and Solanad embryogeny predominates in the Ericanae and Asteridae; or helobial endosperm is a distinctive feature of the Alismidae. On the whole, embryological characteristics are remarkably constant at the family level. In those families where variation does occur, genera are usually constant, although a few notable examples of intrageneric, and even intraspecific, variation do exist, as for example, in patterns of megagametophyte development. In addition to the taxonomic usefulness of the grosser aspects of embryology--the major categories of structure or development--some evidence is presented suggesting that variation in details within a single category, such as size, shape, and cellular characteristics of the developing and mature megagametophyte, may be helpful in determining relationships within families, genera, or species.

The Bases of Angiosperm Phylogeny: Vegetative Morphology

Article

Jan 1975

Coherent patterns of morphology of apparent value in determining taxonomic and phylogenetic relationships are present in dicotyledonous leaves. Features of greatest value in assessing these affinities include leaf organization; marginal features, including morphology of the tooth; major vein configuration; characters of the intercostal venation; and gland placement. Of these, recognition of tooth morphology appears to be an overlooked tool of major systematic importance. Variation in these features is most coherent when analyzed in terms of the Takhtajan and Cronquist systems of dicot classification. Essential to our procedure was a recognition of the "basic" leaf features of each taxon. These were regarded as the most generalized type from which all of the more specialized types in a taxon could have been derived and they were derived from an analysis of the comparative morphology of modern leaves with limited input from the fossil record. The resulting scheme indicates strong correlation of leaf features with six of the seven Takhtajan subclasses, in addition to paralleling and clarifying both systems at the ordinal and familial levels. Conspicuous exceptions are the breakdown of the Asteridae into a possible rosid and a possible dilleniid group, reassignment of the Celastrales and Myrtales to the Dilleniidae, and of the Juglandales to the Rosidae. Affinities of numerous problem taxa, such as the Didymelaceae and Medusagynaceae, are resolved, as are some of the points of disagreement between the Takhtajan and Cronquist arrangements. This analysis also provides the first systematic summary of dicot leaf architectural features and the outlines of a regular systematic method for leaf determination.

Plant Chemosystematics

Article

Jan 1986

The distribution of flavonoids in the Angiosperms

Article

Jan 1979

Polyacetylenes as systematic markers in Dicotyledons

Article

Jul 1982
BIOCHEM SYST ECOL

In the sequence of superorders Rutiflorae-Santaliflorae-Araliiflorae-Asteriflorae an increase in the mean oxidation state for each series of Cn-polyacetylenes and an extension in the range of the carbon atoms of these polyacetylenes in the direction of smaller numbers are observed. These trends of polyacetylene evolution also seem to be operative at lower hierarchic levels in the family Asteraceae.

Alkaloid evolution and angiosperm systematics

Article

Mar 1980
BIOCHEM SYST ECOL

The correlation of biosynthetic steps leading to the primary precursors of the shikimate pathway with the distribution of derived alkaloids on dahlgren's system of classification of angiosperm orders suggests that evolution paralleled gradual blocking of these steps. Phenylalanine-derived alkaloids, with the centre of radiation situated in the magnoliales, are of widespread occurrence in angiosperms, an indication of the antiquity of the character. Anthranilic acid-derived alkaloids, with the centre of radiotion in the rutales, are less widespread. Orders in which such alkaloids co-occur with the former biogenetic group are considered to be of more recent origin. Finally, mevalonate-derived iridoid alkaloids, with the centre of radiation in the gentianales, are even less widespread. Orders in which such alkaloids co-occur with the former biogenetic groups should thus be of still more recent origin. These concepts are summarized bg a phylogenetic tree, which illustrates the divergence of three major groups of angiosperm superorders.

Phylogenetic Implications of Chloroplast DNA Restriction Site Variation in the Mutisieae (Asteraceae)

Article

May 1988

Phylogenetic relationships among 13 species in the tribe Mutisieae and a single species from each of three other tribes in the Asteraceae were assessed by chloroplast DNA restriction site mapping. Initially, 211 restriction site mutations were detected among 16 species using 10 restriction enzymes. Examination of 12 of these species using nine more enzymes revealed 179 additional restriction site mutations. Phylogenetic analyses of restriction site mutations were performed using both Dollo and Wagner parsimony, and the resulting monophyletic groups were statistically tested by the bootstrap method. The phylogenetic trees confirm an ancient evolutionary split in the Asteraceae that was previously suggested by the distribution of a chloroplast DNA inversion. The subtribe Barnadesiinae of the tribe Mutisieae is shown to be the ancestral group within the Asteraceae. The molecular phylogenies also confirm the paraphyly of the Mutisieae and provide statistical support for the monophyly of three of its four currently recognized subtribes (Bamadesiinae, Mutisiinae, and Nassauviinae). The fourth subtribe, Gochnatiinae, is shown to be paraphyletic. Within the subtribes, several closely related generic pairs are identified. Chloroplast DNA sequence divergence among genera of the Asteraceae ranges between 0.7 and 5.4%, which is relatively low in comparison to other angiosperm groups. This suggests that the Asteraceae is either a relatively young family or that its chloroplast DNA has evolved at a slower rate than in other families.

Ornithophily Among the Hawaiian Lobelioideae (Campanulaceae): Evidence from Floral Nectar Sugar Compositions

Article

Nov 1986

Data on the sugar compositions of floral nectars were used to test the hypothesis that the Hawaiian Lobelioideae (Campanulaceae) were pollinated by nectarivorous passerine birds (Fringillidae subfam. Drepanidinae, Meliphagidae) prior to widespread extinctions among the native avifauna. Nectars from 24 individuals (representing different taxa [three genera, ten species, and three interspecific hybrids], different conspecific populations, and different individuals within a population) were analyzed via high-performance liquid chromatography (HPLC). Mean percentages (?one S.D.) and ranges (parenthetically) for each sugar were: fructose, 43.5 ? 2.8% (37.20/o-48.3%); glucose, 54.5 ? 3.1% (47.20/oe62.8%); and sucrose, 2.0 ? 1.6% (0.00/o-5.3%). These nectars are thus hexose-dominant (mean sucrose-hexose ratio of 0.019 ? 0.017), the type preferred by passerine birds.

Floral Morphology, Organogenesis and Interpretation of the Inferior Ovary in Downingia Bacigalupii

Article

Nov 1967

Donald R. Kaplan

Chromosome Numbers and Genetic Systems in the Trigger Plants of Western Australia ( Stylidium ; Stylidiaceae)

Article

Jan 1979

SH James

Chromosome numbers for 116 taxa of the genus Stylidium in south Western Australia are reported. The primitive number is n = 15, but haploid numbers ranging from 5 to 16 occur, with polyploidy on 13, 14 and 15. This dysploid variation is associated with a widespread occurrence of recessive lethal factors which eliminate most of the products of self-pollination at or soon after fertilization while permitting a majority of the cross-pollination products to survive.

Fossil pollen records of extant angiosperms

Article

Jan 1981

Jan Muller

The fossil record for angiosperm pollen types which are comparable to recent taxa is evaluated, following a similar survey published in 1970. Special attention is paid to the dating of the sediments. Evidence for 139 families is considered to be reliable, for others the records are cited as provisional, pending the accumulation of more evidence. Some published records are shown to be erroneous. In the early Cretaceous only types occur indicating the presence of plants ancestral to Magnoliidae and Liliatae. In the Turonian increased differentiation is evident, culminating in the Maestrichtian when evidence for the presence of most major taxa of angiosperms is available. In the Tertiary diversification at lower taxonomic level continues and identification with living taxa becomes more certain. Herbaceous groups tend to appear later in the record than woody ones.

Flora of Tierra del Fuego

Article

Dec 1984
BIOSCIENCE

Alkaloid Bearing Plants and Their Contained Alkaloids

Article

Jan 1961
Lloydia

PoUen Morphology and Plant Taxonomy

Article

Mar 1953
SOIL SCI

G. ERDTMAN

Some tropical African Lobeliaceae. Chromosome numbers, new taxa and comments on taxonomy and nomenclature

Article

Mar 2008
NORD J BOT

Mats Thulin

Chromosome numbers of 21 taxa in Lobelia L. and Cyphia Berg, are reported, the majority being new records. 2n = 18 in Cyphia is a new generic report. The new species Lobelia paludigena Thulin (Zaire), L. kundelungensis Thulin (Zaire), L. cheranganiensis Thulin (Kenya) and L. corniculata Thulin (Swaziland and Natal) are described. The distributions of these species, as well as of L. thermalis Thunb., are mapped. L. kalobaënsis E. Wimm. ex Thulin is validated. The new combinations L. trullifolia Hemsl. ssp. delicatula (Compton) Thulin and L. flaccida (Presl) A. DC. ssp. gnmvikii (T. C. E. Fries) Thulin are made. Typifications are provided for the following names: L. stuhlmannii Schweinf. ex Stuhlm., L. gregoriana Bak. f., L. fervens Thunb., L. anceps L. f. and L. depressa L. f.

Iridoids as systematic markers in dicotyledon

Article

Dec 1982
BIOCHEM SYST ECOL

Iridoid evolution is postulated to involve a gradual increase in the state of oxidation. From this, it appears that the iridoid-containing dicotyledons bifurcated at a primitive stage near the Ericales into the Cornales, with an extension to a Dipsacales branch and a Gentianiflorae branch, and into the Lamiiflorae.

Ovular morphology and the major classification of the dicotyledons

Article

Feb 1974
BOT J LINN SOC

W. R. Philipson

The distribution within the dicotyledons of the number of integuments and the thickness of the nucellus is reviewed. It is suggested that these characters indicate relationship among major dicotyledonous groups. A unitegmic group with crassinucellate or tenuinucellate ovules is considered ancestral to the main sympetalous complex (Asteridae) and, combined with it, forms one of the major phyletic lines within the dicotyledons. A second group, with bitegmic-tenuinucellate ovules, leads to the Primulales-Ebenales complex.

The tapetum: Its form, function, and possible phylogeny in Embryophyta

Article

Sep 1985

It appears that the tapetum is universally present in land plants, even though it is sometimes difficult to recognize, because it serves mostly as a tissue for meiocyte/spore nutrition. In addition to this main function, the tapetum has other functions, namely the production of the locular fluid, the production and release of callase, the conveying of P.A.S. positive material towards the loculus, the formation of exine precursors, viscin threads and orbicules (= Ubisch bodies), the production of sporophytic proteins and enzymes, and of pollenkitt/tryphine. Not all these functions are present in all land plants:Embryophyta. Two main tapetal types are usually distinguished in theSpermatophyta: the secretory or parietal type and the amoeboid or periplasmodial type; in lower groups, however, other types may be recognized, with greater or lesser differences. A hypothetical phylogenesis of the tapetum is proposed on the basis of its morphological appearance and of the nutritional relations with meiocytes/spores. The evolutionary trends of the tapeta tend towards a more and more intimate and increasingly greater contact with the spores/pollen grains. Three evolutionary trends can be recognized: 1) an intrusion of the tapetal cells between the spores, 2) a loss of tapetal cell walls, and 3) increasing nutrition through direct contact in narrow anthers.

Phylogenetisch-systematische Betrachtungen

Article

Sep 1949

Rudolf Leonhardt

A contribution to the life-history ofSphenoclea Zeylanica gaertn.

Article

Distribution of L-(+)-bornesitol in the Gentianaceae and Menyanthaceae

Article

Dec 1976
PHYTOCHEMISTRY

N. Schilling

l-(+)-Bornesitol was detected in 23 of 33 genera of Gentianaceae investigated. The only subtribe without l-(+)-bornesitol (3 species tested) was Exacinae. None of the five genera of Menyanthaceae examined were found to contain l-(+)-bornesitol.

Triggerplants. Patterson Broken-sha

Jan 1958

R Erickson

ERICKSON, R. 1958. Triggerplants. Patterson Broken-sha, Perth. FERREIRA, Z. S. &

Explosive Chromosome Evolution in Downingia yina Secondary me-tabolites and the higher classification of angiosperms

Jan 1972
5-34

R I Foster
Davis
D Frohne
U Jensen
J Gershenzon
T J Mabry

FOSTER, R. I. 1972. Explosive Chromosome Evolution in Downingia yina. Unpublished Ph.D. Dissertation. University of California, Davis. FROHNE, D. & U. JENSEN. 1973. Systematik des Pflanz-enreichs unter Besonderer Berficksichtigung Che-mischer Merkmale und Pflanzlicher Drogen. G. Fi-scher, Stuttgart. GERSHENZON, J. & T. J. MABRY. 1983. Secondary me-tabolites and the higher classification of angiosperms. Nordic J. Bot. 3: 5-34.

Chemotaxonomy of Flowering Plants Index to plant chromo-some numbers 1975-1978. Monogr. Syst. Bot. Mis-souri Bot. Gard. 5. -1984. Index to plant chromosome numbers 1979-1981 Index to plant chromosome numbers 1982-1983

Jan 1974

R D Gibbs

GIBBS, R. D. 1974. Chemotaxonomy of Flowering Plants, Volume 2. McGill-Queen's Univ. Press, Montreal. GOLDBLATT, P. (editor). 1981. Index to plant chromo-some numbers 1975-1978. Monogr. Syst. Bot. Mis-souri Bot. Gard. 5. -1984. Index to plant chromosome numbers 1979-1981. Monogr. Syst. Bot. Missouri Bot. Gard. 8. -1985. Index to plant chromosome numbers 1982-1983. Monogr. Syst. Bcqt. Missouri Bot. Gard. 13. 1988. Index to plant chromosome numbers 1984-1985. Monogr. Syst. Bot. Missouri Bot. Gard.

Affinities of the Cichoriaceae

Jan 1904
59-63

E L Greene

GREENE, E. L. 1904. Affinities of the Cichoriaceae. Leafl. Bot. Observ. Crit. 1: 59-63.

Comparative phytochemistry of al-kaloids. Pp. 211-230 in Compar-ative Phytochemistry Birkhauser, Basel. -1977. The chemistry of the Compositae The Biology and Chemistry of the Compositae

Jan 1964
283-33547

R Hegnauer
Birkhauser
Basel

HEGNAUER, R. 1964. Chemotaxonomie der Pflanzen, Volume 3. Birkhauser, Basel.. 1966a. Comparative phytochemistry of al-kaloids. Pp. 211-230 in T. Swain (editor), Compar-ative Phytochemistry. Academic Press, London.. 1966b. Chemotaxonomie der Pflanzen, Vol-ume 4. Birkhauser, Basel. -1973. Chemotaxonomie der Pflanzen, Volume 6. Birkhauser, Basel. -1977. The chemistry of the Compositae. Pp. 283-335 in V. H. Heywood, J. B. Harborne & B. L. Turner (editors), The Biology and Chemistry of the Compositae. Academic Press, London. HERZ, W. 1977. Sesquiterpene lactones in the Com-positae. Pp. 337-357 in V. H. Heywood, J. B. This content downloaded from 128.230.234.162 on Mon, 29 Jul 2013 07:15:47 AM All use subject to JSTOR Terms and Conditions Missouri Botanical Garden Harborne & B. L. Turner (editors), The Biology and Chemistry of the Compositae. Academic Press, Lon-don.

1889. Calyceraceae. Pp. 84-86 in A. Engler, Die Natfirlichen Pflanzenfamilien

F Hock

HOCK, F. 1889. Calyceraceae. Pp. 84-86 in A. Engler, Die Natfirlichen Pflanzenfamilien, Teil 4, Abteilung

Em-bryology of Pentaphragma horsfieldii (Miq.) Airy Shaw with a discussion on the systematic position of the genus

Jan 1965
93-102

R N Kapil
M R Vijayaraghavan

KAPIL, R. N. & M. R. VIJAYARAGHAVAN. 1965. Em-bryology of Pentaphragma horsfieldii (Miq.) Airy Shaw with a discussion on the systematic position of the genus. Phytomorphology 15: 93-102.

Campanulaceae, Lobeliaceae Flowering Plants of the World

Jan 1912
254-256

M Kovanda

KOVANDA, M. 1978. Campanulaceae, Lobeliaceae. Pp. 254-256 in V. H. Heywood (editor), Flowering Plants of the World. Mayflower, New York. KRAUSE, K. 1912a. Goodeniaceae. Pp. 1-207 in A.

Abteilung 277 Wilhelm Engelmann, Leipzig. 1912b. Brunoniaceae. Pp. 1-6 in Some aspects of the classification and evolution of higher taxa

Jan 1977
21-31

Das Engler
Pflanzenreich
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Engler, Das Pflanzenreich, Teil IV, Abteilung 277. Wilhelm Engelmann, Leipzig. 1912b. Brunoniaceae. Pp. 1-6 in A. Engler (editor), Das Pflanzenreich, Teil IV, Abteilung 277a. Wilhelm Engelmann, Leipzig. KUBITZKI, K. 1977. Some aspects of the classification and evolution of higher taxa. P1. Syst. Evol., Suppl. 1: 21-31.

Cytology of Cyananthus linifolius Wall Chromosome numbers of Ha-waiian Lobelioideae (Campanulaceae), and the sys-tematic utility of cytological data in the subfamily Chromosome numbers and their sys-tematic implications in the Hawaiian Lobelioideae (Campanulaceae)

Jan 1975
1130-1134

V Kumar
K P S Chauhan
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KUMAR, V. & K. P. S. CHAUHAN. 1975. Cytology of Cyananthus linifolius Wall. Cat. Proc. Indian Sci. Congr. Assoc. 62(3), Group B: 130. LAMMERS, T. G. 1987. Chromosome numbers of Ha-waiian Lobelioideae (Campanulaceae), and the sys-tematic utility of cytological data in the subfamily. Ohio J. Sci. 87: 7. [Abstract.]. 1988. Chromosome numbers and their sys-tematic implications in the Hawaiian Lobelioideae (Campanulaceae). Amer. J. Bot. 75: 1130-1134.

Campanulaceae (Lobelioideae)

Jan 1943
1-134

R Mcvaugh

MCVAUGH, R. 1943. Campanulaceae (Lobelioideae). N. Amer. Fl. 32A: 1-134.

Caffeic acid as a taxonomic mark-er in dicotyledons

Jan 1960
107-141

B Moeliono
P Tuyn

MOELIONO, B. & P. TUYN. 1960. Campanulaceae. Pp. 107-141 in C. G. G. J. van Steenis (editor), Flora Malesiana, Series 1, Volume 6, Part 1. Noordhoff-Kolff, Djakarta. M0LGAARD, P. 1985. Caffeic acid as a taxonomic mark-er in dicotyledons. Nordic J. Bot. 5: 203-213.

A propos du Sphenoclea zeylanica (Sphenocleaceae) Adansonia (series 2

Jan 1980
147-164

T Monod

MONOD, T. 1980. A propos du Sphenoclea zeylanica (Sphenocleaceae). Adansonia (series 2) 20: 147-164.

A revision of the genus Nemacladus (Campanulaceae)

Jan 1924
233-248

P A Munz

MUNZ, P. A. 1924. A revision of the genus Nemacladus (Campanulaceae). Amer. J. Bot. 11: 233-248 + pls. IX-X.

Circumscription and Phylogeny of the Campanulales

Abstract

No full-text available

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