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Based on new studies on style morphology of basal subfamilies and tribes of Asteraceae (Barnadesioideae, Famatinanthoideae, Mutisioideae, Stifftioideae, Wunderlichioideae, Gochnatioideae, Hecastocleidoideae), we establish 15 style types in view of shape, bifurcation and distribution of stylar hairs and stigmatic tissue. From the stylar characters and observing different anthetic stages of the proterandrous flowers, we inferred the mechanisms of secondary pollen presentation. The high diversity of stylar shapes in the basal groups of Asteraceae is remarkable. In the most basal subfamily Barnadesioideae, the stylar surface is smooth or, at most, papillate and secondary pollen presentation is either a deposition/primitive brushing or a pump mechanism. Considering Calyceraceae (deposition mechanism) as sister group, we conclude that presumably a deposition/primitive brushing mechanism is the beginning of all secondary pollen presentations in Asteraceae. Whereas in the higher Asteraceae both the brushing and pump mechanisms function perfectly, in some lower Asteraceae, the mechanisms work less efficiently regarding pollen release. The subfamily Mutisioideae is well-defined by the occurrence of true stylar hairs and the tribe Mutisieae by its stigmatic tissue on the margins of the stylar arms, confluent at their tips. The early step from a continuous stigma covering the whole inner surface of the stylar branches (as in most basal members) to a stigma arrangement in two separate lines (as in most Asteroideae) is realized in two ways: either by adhesion of the median ventral tissues of the stylar branches (e.g. Onoseris , Wunderlichia ) or by receptivity loss of the median tissues (e.g. Mutisia ). Our study supports further DNA-based clades by style morphology (e.g. Famatinanthoideae).

Barnadesioideae (94 species) is the sister subfamily to the rest of the Asteraceae (23,000 species). Pollen grains in this subfamily are structurally and sculpturally distinctive and diverse. Although pollen morphology has contributed to the taxonomy of the subfamily, there is a gap of knowledge concerning the evolution of the exine structure. This study aims at exploring the systematic and phylogenetic significance of optimizing selected pollen characters of Barnadesioideae on the latest molecular phylogenetic tree. Transmission electron microscope (TEM) observations on pollen of selected species, some of them never explored so far, show that the exine probably evolved from a thin pattern (ca. 1–3 μm), with a well-developed foot layer and solid and free columellae, present in sister family Calyceraceae, towards a thicker (> 6–11 μm) and a more complex columellate-granulate bilayered exine in Barnadesioideae (with very delicate columellae). The particular exine structure observed in the monotypic Schlechtendalia luzulaefolia, which combines compact and independent columellae (common in more derived Asteraceae) with a granular internal tectum as the inner ectexine layer (as in Barnadesioideae), reinforces its distant phylogenetic position within Barnadesioideae. More derived lineages within Asteraceae (e.g. Mutisioideae) retained some ancestral exine features although evolved an even thicker exine and a columellate trilayered exine (with robust columellae), rare in the angiosperm pollen grains. Fil: Tellería, María Cristina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Laboratorio de Sistemática y Biología Evolutiva; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Palazzesi, Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Barreda, Viviana Dora. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina

Background and Aims The plant family Asteraceae (Compositae) exhibits remarkable morphological variation in the styles of its members. Lack of studies on the styles of the sister families to Asteraceae, Goodeniaceae and Calyceraceae, obscures our understanding of the origin and evolution of this reproductive feature in these groups. The aim of this work was to perform a comparative study of style morphology and to discuss the relevance of important features in the evolution of Asteraceae and its sister families. Methods The histochemistry, venation and general morphology of the styles of members of Goodeniaceae, Calyceraceae and early branching lineages of Asteraceae were analysed and put in a phylogenetic framework to discuss the relevance of style features in the evolution of these families. Key Results The location of lipophilic substances allowed differentiation of receptive from non-receptive style papillae, and the style venation in Goodeniaceae and Calyceraceae proved to be distinctive. There were several stages of style evolution from Goodeniaceae to Asteraceae involving connation and elongation of veins, development of bilobation from an initially cup-shaped style, and a redistribution of the receptive and non-receptive papillae. Conclusions These developments resulted in bifurcation in the styles of Asteraceae, with each branch face having a different function, and it is suggested here as a mechanism that promoted outcrossing, which in turn led to the great diversification in the family., Facultad de Ciencias Naturales y Museo, Facultad de Ciencias Agrarias y Forestales

The nomenclatural revision of 51 names previously published, combined or synonymyzed under the genus Nastanthus (Calyceraceae) is presented. Six species are recognized with correct names and new synonyms established. The nomenclature of Nastanthus ventosus, Nastanthus scapigerus and Nastanthus compactus is updated, giving a total of 45 synonyms. Lectotypes for Boopis caespitosa, Boopis scapigera, Boopis spathulata, Nastanthus laciniatus, Boopis araucana, Calycera ventosa, Nastanthus pinnatifidus and Boopis breviflora are here designated. A map with the geographic distributions, complete species descriptions, corrected illustrations, and a new key to accepted species of Nastanthus, are included.Se presenta la revisión nomenclatural de 51 nombres publicados, combinados o sinonimizados bajo el género Nastanthus (Calyceraceae). Se reconocen seis especies, estableciendo los nombres correctos y sinónimos nuevos. Se actualizó la nomenclatura de Nastanthus ventosus, Nastanthus scapigerus y Nastanthus compactus, con un total de 45 sinónimos. Se designaron lectotipos para Boopis caespitosa, Boopis scapigera, Boopis spathulata, Nastanthus laciniatus, Boopis araucana, Calycera ventosa, Nastanthus pinnatifidus y Boopis breviflora. Se incluyeron, además, un mapa de distribución geográfica, descripciones completas, ilustraciones corregidas y una clave nueva para las especies aceptadas de Nastanthus.

Pollen morphology of 13 species from all six genera of Calyceraceae (Acicarpha, Boopis, Calycera, Gamocarpha, Moschopsis, and Nastanthus) and representatives of the Campanulaceae and Goodeniaceae is examined with light (LM), scanning (SEM), and transmission (TEM) electron microscopy. Acicarpha, Calycera, and Nastanthus pollen grains are distinguished by angulaperturate apertures, colpar ledges and surface depressions between colpi known as intercolpar concavities (IC). Pollen of Gamocarpha and Moschopsis is tricolporate rather than angulaperturate and without an IC. Some species of Boopis are similar to the preceding genera (e.g., B. graminea), while others (e.g., B. gracilis) are angulaperturate with ICs. Structural features derived from fractured pollen in SEM and sections in TEM show pollen walls composed of prominent columellae ca. 0.55–1.1 µm high and

Barreda et al. (1) claim a Cretaceous fossil pollen type is an extinct Asteraceae. Concluding this pollen type is “nested within Dasyphyllum (crown representative),” they calibrate a Dasyphyllum + Barnadesia crown node (Dasyphyllum crown absent) and estimate an 85.9-Ma Asteraceae crown age that potentially compresses asterid evolution by tens of millions of years. However, the bootstrap majority consensus topology reported could not be reproduced from the data; instead, the fossil resolved in a trichotomy with Calyceraceae and Asteraceae. Thus, unambiguous assignment of these pollen grains to Asteraceae is premature.

Palynomorphological characteristics of the order Asterales are discussed. Particular attention is paid to the pollen morphology of basal families of this group and to that of problematic taxa that are considered as sister groups to the group under study. Ultrastructurally similar sporoderms of several families, including (1) Asteraceae, Calyceraceae, and Goodeniaceae; (2) Campanulaceae, Phellinaceae, and Menyanthaceae; (3) Rousseaceae, Abrophyllaceae, and Columelliaceae, are described. Pollen grains of Alseuosmiaceae and Stylidiaceae show unique ultrastructural features of the exine.

Rhamnaceae allied to fetid honeys in Argentine. Melissopalynological analysis of six honey samples, commercially rejected for its fetid odour, was performed. These honeys were produced from Espinal and Pampeana Phytogeographical Provinces. The main nectar sources that characterized monofloral honeys belong to the following families: Rhamnaceae (Discaria americana, Scutia buxifolia) and Calyceraceae (Acicarpha tribuloides) along with pollen of S. buxifolia as secondary. According with the results of the present study it seems that the odour has its origin in the Rhamnaceae flowers.

We obtained two most parsimonious cladograms using a data set of 100 characters derived from morphology, anatomy, embryology, chemistry, and karyology, combined with three nucleotide sequence data sets (the chloroplast genes atpB, ndhF, and rbcL) in a phylogenetic analysis of all 12 currently recognized families in the angiosperm order Asterales, represented by 40 genera. Most clades were supported by a jackknife value of at least 50% and a Bremer support of 5 or more. Rousseaceae sensu lato (including Carpodetaceae), together with Pentaphragmataceae and Campanulaceae s.l., is the sister group to the rest of the Asterales. A sister group relationship between Donatia and Stylidiaceae is well supported both morphologically and by molecular data, and we suggest that Donatia should again be treated as a subfamily in Stylidiaceae. The sister group relationship between Calyceraceae and Asteraceae is well supported.

The subfamily Barnadesioideae of the Asteraceae consists of nine genera and approximately 90 species. Both molecular and morphological phylogenies indicate that this subfamily is sister to the rest of the family. We have used scanning electron microscopy (SEM) and transmission electron microscopy (TEM) to study pollen of 41 species from all genera of the Barnadesioideae. Three general pollen types are described in the subfamily: Barnadesia-type (Barnadesia, Huarpea), Chuquiraga-type (Chuquiraga, Doniophyton, Duseniella, Fulcaldea) and Dasyphyllum-type (Dasyphyllum and Schlechtendalia). A fourth type, Amaldoa-type, consisting solely of Arnaldoa, is intermediate between the Chuquiraga- and Dasyphyllum-types. These types parallel and confirm findings from previous studies. Psilolophate grains are found only in the Barnadesia-type. Pollen with a cavity (cavea) between pollen wall units in each of the three interapertural regions is present in Barnadesia (Barnadesia-type), Dasyphyllum (Dasyphyllum-type) and Arnaldoa (Arnaldoa-type). The Chuquiraga-type does not have cavate pollen. Intercolpar concavities occur only in the Dasyphyllum- and Arnaldoa-types. In the latter, intercolpar regions are accompanied by pairs of indentations flanking the colpi. The presence of intercolpar concavities in Dasyphyllum and Schlechtendalia, often cited as a synapomorphy for the Barnadesioideae and Calyceraceae, has apparently evolved independently within the sub-family. Chuquiraga pollen exhibits the least derived palynological features in the subfamily. Palynological characters, when placed in the context of current phylogenies for the Barnadesioideae, suggest additional phylogenetic analyses are needed to re-evaluate intergeneric relationships within the subfamily.

The relationships between Asteraceae, Calyceraceae, Goodeniaceae, and Brunoniaceae are discussed, and some SEM-observations of microcharacters within the two latter families are presented. Based on morphological evidence neither Goodeniaceae s.str. nor Goodeniaceae s.lat. (i.e., incl. Brunonia) appear to be the putative sister taxon of Asteraceae. A cladistic analysis demonstrates that Brunonia australis may constitute a separate, monotypic family or alternatively be deeply nested within Goodeniaceae s.lat Brunonia shares a number of potential synapomorphies with Asteraceae. Of special interest are the peculiar hairs and feathery pappus bristles which very much resemble those found within Asteraceae subfam. Barnadesioideae. However, studies with SEM disclose that these structures rather constitute autapomorphies for the taxa in question. In conclusion, Calyceraceae remain to be the best sister candidate of the sunflower family.

Palynological data obtained by LM, SEM and TEM support a division of the Goodeniaceae into three groups. The Lechenaultia group has rhomboidal tetrads with pores, interconnected columellae, and lacks a foot layer. The Anthotium‐Dampiera group has tricolporate, rugulate grains with a thin infratectum and rounded ora. The Scaevola‐Goodenia group, finally, comprises the remaining genera in the family and is characterized by tricolporate, microspinulose grains with distally branched columellae and lalongate ora. Within the latter group, Brunonia stands somewhat apart, having ridges and depressions along the colpi. The Scaevola‐Goodenia group shows some similarity to members of the Asteraceae, and particularly, to the Calyceraceae. The other two groups are morphologically isolated, both having a number of rare or unique palynological features.

Embryological studies of Campanulaceae and related families were reviewed to add to discussion of appropriate circumscription of Campanulaceae and Campanulales. Embryological features are very uniform within Campanulaceaesens. lat. (Campanuloideae and Lobelioideae), and provide no evidence for the separation of Lobelioideae as a distinct family. Campanulaceaesens. lat. share with several other families traditionally included in Campanulales (or Asterales) a few derived embryological features (i.e., unitegmic and tenuinucellate ovules, an endothelium,ab initio Cellular type endosperm, and an endosperm haustorium). However, an extensive comparison with families in an expanded Asteridae shows that those derived embryological features are symplesiomorphies, rather than synapomorphies indicative of close relationships among them. Combined with other morphological (inferior ovary), chemical (presence of inulin), and molecular (rbcL gene nucleotide sequence) characters, embryological features rather support a broad circumscription of Campanulales (or Asterales) including Pentaphragmataceae, Stylidiaceae, Campanulaceaesens. lat., Donatiaceae, Menyanthaceae, Calyceraceae, Goodeniaceae, and Asteraceae. Within Campanulalessens. lat. (or Asteralessens. lat.), Menyanthaceae, Calyceraceae, Goodeniaceae, and Asteraceae are obviously closely related to one another. To understand more exact relatonships within the order, embryological studies are needed onBrunonia, Cyphia, Cyphocarpus, Nemacladus, Corokia, Abrophyllum, Argophyllaceae, and Alseousmiaceae.

Summary Pesacreta, T. C. & Stuessy, T. F.: Autofluorescent walls of connective bases in anthers of Barnadesioideae (Asteraceae), and systematic implications. - Taxon 45: 473-485. 1996. - ISSN 0040-0262. Stamens from 20 species of the subfamily Barnadesioideae were examined with confocal scanning laser microscopy to determine some of the characteristics of the connective base epidermis. All species had autofluorescent cell walls. Wall autofluorescence was more intense in the abaxial connective epidermis than in the filament epidermis. In most species all autofluorescent cell walls were evenly thickened, but in several species of Barnadesia some either had spiral thickenings or were irregularly thickened, perhaps an adaptation for increased strength in longer filaments and anthers as part of a hummingbird pollination syndrome, a derived condition within the subfamily. In slightly less than half the species the connective base exhibited a more intense autofluorescence than any other region of the connective. The adaxial epidermis was autofluorescent in eight species. The occurrence of autofluorescent cells in both Calyceraceae and Barnadesioideae suggests this as a possible synapomorphy, but in the former the cells are tangentially elongate and irregular, rather than longitudinally elongate with rectangular outline. Taxonomic investigations in Asteraceae over the past two centuries have taught us that, in general, reproductive features contain more useful taxonomic information than vegetative ones. Features of stamens and carpels have been used extensively at different levels of classification of the family, particularly in the delimitation of tribes and subtribes. Style branch morphology and anther features have been especially employed (e.g. by Bentham, 1873; Small, 1919; Cronquist, 1955, 1977; Bremer, 1994). Characters of anthers that have been used taxonomically include length, colour, shape and size of thecal cells, tails, apical appendages, and aspects of filaments. All have been documented as taxonomically efficacious in certain contexts. One particular portion of the anthers, the "article antherifere" (Cassini, 1814), "antheropodium" (or "antheropode"; Beauvard, 1910; Briquet 1917, 1918), "collar" (Robinson & King, 1977), or "connective base" (Pesacreta & al., 1993), which is the area at the base of the connective where it attaches to the filament, has recently been shown to be helpful in suggesting taxonomic affinities, particularly at the generic level (e.g., King & Robinson, 1987). Current work in the laboratory of the senior author is surveying detailed variation in aspects of the connective bases within the family and assessing its taxonomic potential. Because these cells have thick walls and are autofluorescent, confocal scanning laser microscopy can be used to reveal their characteristics in conjunction with regular light microscopic techniques. Autofluorescence reveals the presence of lignin, and therefore gives data different from simple observations of wall thickness with conventional transmitted-light microscopy.

TherbcL gene of 25 taxa was sequenced and analyzed cladistically in order to define more precisely the orderAsterales s.l. and to reconstruct the phylogeny ofGoodeniaceae. The cladistic analyses show that theAsterales comprise the familiesAbrophyllaceae, Alseuosmiaceae, Argophyllaceae, Asteraceae, Calyceraceae, Campanulaceae s.l.,Donatiaceae, Goodeniaceae (includingBrunoniaceae),Menyanthaceae, Pentaphragmataceae, andStylidiaceae. Abrophyllaceae, Alseuosmiaceae, Brunoniaceae, andDonatiaceae have previously not been studied in this respect. Within theGoodeniaceae, four groups supported by therbcL data can be distinguished: the genusLechenaultia, theAnthotium-Dampiera-group, the genusBrunonia, and a group formed by the remaining genera, theScaevola-Goodenia-group.

Flavonoid profiles were determined for 11 species representing five genera of Calyceraceae: Acicarpha, Boöpis, Calycera, Gamocarpha, and Nastanthus. Kaempferol, quercetin, 6-methoxykaempferol, and 6-methoxyquercetin (patuletin) were unequivocally identified. Kaempferol and quercetin occurred as 3-O-mono- and 3-O-diglycosides, whereas the latter two flavonols were observed only as aglycones. Several unidentified phenolic compounds were also noted. The simplest profile in the family consisted solely of the kaempferol and quercetin glycosides in Acicarpha, Gamocarpha, and one specimen of Calycera leucanthema. Morphological evidence suggests that Acicarpha represents prototypical Calyceraceae and that a close relationship exists between primitive Calyceraceae and some basal members of Asteraceae (subfamily Barnadesioideae). This last possibility is reinforced by the similarity of flavonoid profiles of Acicarpha and members of Barnadesioideae. Key words: Calyceraceae, Asteraceae, Compositae, Barnadesioideae, flavonoids.

Summary 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). In addition, a few remarks are given on pollen presentation in Rubiaceae, Cyphiaceae, Sphenocieaceae, Pentaphragmataceae, Stylidiaceae , of which the latter four are usually assigned to Campanulales-Asterales .

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 ...

 Premise of the study: Phylogenies based on molecular data are revealing that generalizations about complex morphological structures often obscure variation and developmental patterns important for understanding the evolution of forms, as is the case for infl orescence morphology within the well-supported MGCA clade (Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae). While the basal families share a basic thyrsic/thyrsoid structure of their infl orescences, Asteraceae possesses a capitulum that is widely interpreted as a racemose, condensed infl orescence. Elucidating the poorly known infl orescence structure of Calyceraceae, sister to Asteraceae, should help clarify how the Asteraceae capitulum evolved from thyrsic/thyrsoid infl orescences.  Methods: The early development and structure of the infl orescence of eight species (fi ve genera) of Calyceraceae were studied by SEM, and patterns of evolutionary change were interpreted via phylogenetic character mapping.  Key results: The basic infl orescence structure of Calyceraceae is a cephalioid (a very condensed botryoid/thyrsoid). Optimization of infl orescence characters on a DNA sequence-derived tree suggests that the Asteraceae capitulum derives from a simple cephalioid through two morphological changes: loss of the terminal fl ower and suppression of the cymose branching pattern in the peripheral branches.  Conclusions: Widely understood as a condensed raceme, the Asteraceae capitulum is the evolutionary result of a very reduced, condensed thyrsoid. Starting from that point, evolution worked separately only on the racemose developmental control/pattern within Asteraceae and mainly on the cymose developmental control/pattern within Calyceraceae, producing head-like infl orescences in both groups but with very different diversifi cation potential. We also discuss possible remnants of the ancestral cephalioid structure in some Asteraceae.

We describe and illustrate two new species of Boopis from Argentina: Boopis pterocalyx (Mendoza province) and Boopis necronensis (Catamarca and La Rioja provinces). Both species are easily distinguished from the remaining species of Boopis (sensu lato, including Nastanthus) by the foliose, imbricate sepals, which become foliose and orbicular in the fruit of B. pterocalyx, or navicular, obtuse-mucronate and corky in the fruit of B. necronensis. A geographical distribution map, complete descriptions, and illustrations are also included. Fil: Zavala Gallo, Lucio Martín. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Botánica Darwinion. Academia Nacional de Ciencias Exactas, Físicas y Naturales. Instituto de Botánica Darwinion; Argentina Fil: Denham, Silvia Suyai. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Botánica Darwinion. Academia Nacional de Ciencias Exactas, Físicas y Naturales. Instituto de Botánica Darwinion; Argentina Fil: Pozner, Raúl Ernesto. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Botánica Darwinion. Academia Nacional de Ciencias Exactas, Físicas y Naturales. Instituto de Botánica Darwinion; Argentina

The South American family Calyceraceae is analysed and some new observations on pollen and petal epidermis-patterns are presented. The new characters give some indication of a relationship to Compositae. The choice of Calyceraceae as the sister-group of Compositae is discussed, and it is further suggested that Compositae, Calyceraceae, and Lobeliaceae constitute a clade. Two new specific combinations in the Calyceraceae are formally established: Gamocarpha alpina and Calycera andina.

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.

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

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 (...)

Stems from more than 555 species representing 440 genera of approximately 210 dicotyledonous families were surveyed for the presence of fructose oligosaccharides. The tissues from 372 species representing 312 genera and 187 families did not contain such oligosaccharides. The sugars occurred sporadically or in small amounts in members of the following families: Crossosomataceae, Lauraceae, Malpighiaceae, Cornaceae, Myrisinaceae, Melianthaceae, Greyiaceae, and Simaroubaceae. The fructosyl sucrose trisaccharide, isokestose, aswell as its fructose tetrasaccharide homolog (but not higher inulin oligosaccharides) were present in members of the families Hippocastanaceae and Limnathaceae. Stems from members of the families Clethraceae and Monotropaceae, and some members of the Polemoniaceae, contained two series of fructose oligosaccharides: one based on isokestose (the inulin series) and the other one based on kestose (the levan series). The inulin or isokestose-based series of fructose oligosaccharides, alone, was present in members of the following families: Menyanthaceae, Boraginaceae, Campanulaceae, Goodeniaceae, Stylidiaceae, Brunoniaceae, Calyceraceae and Compositae. The primary metabolites occur ubiquitously and consistently in tissues of plants of these families. We hypothesize that this conservative character represents several phylogenetic alliances among them.

Ten taxa of the families Valerianaceae, Dipsacaceae, and Calyceraceae are recorded as naturalised in New Zealand. One additional record is unsubstantiated.