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The Calyceraceae (47 spp.) is a small family of plants that is sister to the Asteraceae (∼ 25,000 spp.), one of the largest families of angiosperms. Most members of Calyceraceae are endemic to the Andes and Patagonia, representing an excellent model within which to study diversification patterns in these regions. The single phylogenetic study of Calyceraceae conducted to date revealed that the boundaries of most genera and several species of this family require further analyses, especially the “Nastanthus–Gamocarpha” clade. In this study, we reconstructed the phylogeny of the “Nastanthus–Gamocarpha” clade using multispecies coalescent models under BPP and StarBeast2 programs, sampling 63 individuals from 13 of the 14 species recognized to date. We then used this phylogenetic framework to delimit species using BFD and the A11 method implemented in BPP. Species limits suggested through a coalescent approach were then re-evaluated in the light of morphology, geography, and phenology. Coalescent-based methods indicated that most putative lineages could be recognized as distinct species. Morphological, geographical, ecological, and phenological data further supported species delimitation. Necessary taxonomic changes are proposed. Namely, the paraphyletic Nastanthus is synonymized under Gamocarpha, while five species of Boopis are transferred into Gamocarpha. We used an integrative taxonomic approach to recognize 13 species and one subspecies within the newly circumscribed genus Gamocarpha. Fil: Denham, Silvia Suyai. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo; Argentina. 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: Brignone, Nicolás Fernando. 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: Johnson, Leigh. University Brigham Young; Estados Unidos 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

Biodiversity results from multiple evolutionary mechanisms, including genetic variation and natural selection. Whole-genome duplications (WGDs), or polyploidizations, provide opportunities for large-scale genetic modifications. Many evolutionarily successful lineages, including angiosperms and vertebrates, are ancient polyploids, suggesting that WGDs are a driving force in evolution. However, this hypothesis is challenged by the observed lower speciation and higher extinction rates of recently formed polyploids than diploids. Asteraceae includes about 10% of angiosperm species, is thus undoubtedly one of the most successful lineages and paleopolyploidization was suggested early in this family using a small number of datasets. Here, we used genes from 64 new transcriptome datasets and others to reconstruct a robust Asteraceae phylogeny, covering 73 species from 18 tribes in six subfamilies. We estimated their divergence times and further identified multiple potential ancient WGDs within several tribes and shared by the Heliantheae alliance, core Asteraceae (Asteroideae-Mutisioideae), and also with the sister family Calyceraceae. For two of the WGD events, there were subsequent great increases in biodiversity; the older one proceeded the divergence of at least 10 subfamilies within 10 My, with great variation in morphology and physiology, whereas the other was followed by extremely high species richness in the Heliantheae alliance clade. Our results provide different evidence for several WGDs in Asteraceae and reveal distinct association among WGD events, dramatic changes in environment and species radiations, providing a possible scenario for polyploids to overcome the disadvantages of WGDs and to evolve into lineages with high biodiversity.

Spatial and temporal differences in ecological opportunity can result in disparity of net species diversification rates and consequently uneven distribution of taxon richness across the tree of life. The largest eudicotyledonous plant family Asteraceae has a global distribution and at least 460 times more species than its South American endemic sister family Calyceraceae. In this study, diversification rate dynamics across Asteraceae are examined in light of the several hypothesized causes for the family's evolutionary success that could be responsible for rate change. The innovations of racemose capitulum and pappus, and a whole genome duplication event occurred near the origin of the family, yet we found the basal lineages of Asteraceae that evolved in South America share background diversification rates with Calyceraceae and their Australasian sister Goodeniaceae. Instead we found diversification rates increased gradually from the origin of Asteraceae approximately 69.5Ma in the late Cretaceous through the Early Eocene Climatic Optimum at least. In contrast to earlier studies, significant rate shifts were not strongly correlated with intercontinental dispersals or polyploidization. The difference is due primarily to sampling more backbone nodes, as well as calibrations placed internally in Asteraceae that resulted in earlier divergence times than those found in most previous relaxed clock studies. Two clades identified as having transformed rate processes are the Vernonioid Clade and a clade within the Heliantheae alliance characterized by phytomelanic fruit (PF Clade) that represents an American radiation. In Africa, subfamilies Carduoideae, Pertyoideae, Gymnarrhenoideae, Cichorioideae, Corymbioideae, and Asteroideae diverged in a relatively short span of only 6.5millionyears during the Middle Eocene.

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

Calyceraceae is a small family with six traditionally recognized genera and 47 species from southern South America. Most species grow along the Andes (of both Argentina and Chile) and in arid regions of the Patagonian steppe. This family belongs to the well-supported MGCA clade within Asterales, which includes Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae. Calyceraceae is monophyletic and sister to Asteraceae, one of the five largest families of angiosperms. Although Calyceraceae is clearly distinct as a family, its genera are not, and taxonomic revisionary effort has confirmed the lack of sharp boundaries among genera. We performed a phylogenetic analysis of Calyceraceae with a broad taxon sampling (41 of 47 species), and with sequence data from multiple regions from the nuclear (ITS) and plastid genomes (ycg6-psbM, psbM-trnD,trnS-trnG, trnH-psbA, trnD-trnT) using maximum parsimony and Bayesian approaches. We aimed at identifying monophylectic groups, their putative morphological synapomorphies and their geographical distribution; we also estimated divergence times and examined chromosomes numbers in an evolutionary context. We obtained well-resolved and strongly supported phylogenies that show Calyceraceae to be divided into two major clades with geographically structured subclades within each. Our results indicate that an early split within Calyceraceae occurred about 27.4 Ma, probably related to differential changes in chromosome numbers, which allowed the two lineages to evolve in sympatry. We found that major natural subgroups diverged 15?12 Ma, following the Early-Miocene South Andes construction stage. Finally, the diversification of the extant species is probably associated to Andean orogeny and climate changes in the last 5?4 Myr. We recovered Acicarpha as monophyletic, while the remaining traditionally recognized genera of Calyceraceae are para- or polyphyletic. Most species of Moschopis are included in the Glutinose group, but M. monocephala is more closely related to some Calycera species. Calycera is divided into two clades: the Calycera group and the Pilose group. All species of Nastanthus are placed in a well-supported main group with species of Gamocarpha and Boopis. Gamocarpha could be monophyletic after exclusion of G. dentata and G. angustifolia, but is nested within Nastanthus and Boopis species. Boopis is clearly polyphyletic with its species distributed in all main groups. 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. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo; Argentina 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: Johnson, LeighA.. Brigham Young University; Estados Unidos 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

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

Premise of the studyLike many other flowering plants, members of the Compositae (Asteraceae) have a polyploid ancestry. Previous analyses found evidence for an ancient duplication or possibly triplication in the early evolutionary history of the family. We sought to better place this paleopolyploidy in the phylogeny and assess its nature.MethodsWe sequenced new transcriptomes forBarnadesia, the lineage sister to all other Compositae, and four representatives of closely related families. Using a recently developed algorithm, MAPS, we analyzed nuclear gene family phylogenies for evidence of paleopolyploidy.Key resultsWe found that the previously recognized Compositae paleopolyploidy is also in the ancestry of the Calyceraceae. Our phylogenomic analyses uncovered evidence for a successive second round of genome duplication among all sampled Compositae exceptBarnadesia.ConclusionsOur analyses of new samples with new tools provide a revised view of paleopolyploidy in the Compositae. Together with results from a high densityLactucalinkage map, our results suggest that the Compositae and Calyceraceae have a common paleotetraploid ancestor and most Compositae are descendants of a paleohexaploid. Although paleohexaploids have been previously identified, this is the first example where the paleotetraploid and paleohexaploid lineages have survived over tens of millions of years. The complex polyploidy in the ancestry of the Compositae and Calyceraceae represents a unique opportunity to study the long-term evolutionary fates and consequences of different ploidal levels.

Much of our knowledge of the past distribution and radiation of Asteraceae and allied families depends on the fossil pollen record. In recent years, new discoveries are coming to light from southern Africa, Australia, New Zealand, and southern South America (Patagonia). Unequivocally assigned morphotaxa from accurately dated sediments have permitted for the first time a comprehensive review of the past distribution of the most important core of the sunflower alliance of families (Menyanthaceae, Goodeniaceae, Calyceraceae and Asteraceae). The main goal of this contribution is to explore the major evolutionary radiation of the basal lineages of Asteraceae (Mutisioideae and Barnadesioideae) and allied relatives (Menyanthaceae, Goodeniaceae and Calyceraceae) on the basis of the worldwide fossil pollen records. Several taxa, which today are restricted to isolated geographic regions, were widespread in the Southern Hemisphere during Paleogene times. Menyanthaceae, Goodeniaceae and Mutisioideae (Asteraceae), for example, had a wide distribution over Gondwanan landmasses in the Oligocene and are now drastically reduced in their geographic range. Early Neogene records, in contrast, suggest extinction and diversification events that progressively led to the present day configuration. In broad terms, the distribution of Miocene fossils assigned to this clade (Barnadesioideae, Nassauvieae, and Calyceraceae) agrees with that of their present distribution. The major floristic turnovers coincided with the final isolation of Antarctica, leading to cooler, drier, and more seasonal climates and forced the evolution and distribution of these Gondwanan elements. 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 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: Tellería, María Cristina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; Argentina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Laboratorio de Sistemática y Biología Evolutiva; Argentina Fil: Katinas, Liliana. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; Argentina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División de Plantas Vasculares; Argentina Fil: Crisci, Jorge Victor. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; Argentina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Laboratorio de Sistemática y Biología Evolutiva; Argentina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División de Plantas Vasculares; Argentina

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

Chromosome numbers were determined from microsporocytes (meiotic counts) and root-tip cells (mitotic counts) in 201 collections of 51 genera and 119 specific and infraspecific taxa belonging to nine tribes of Asteraceae and Acicarpha spathulata R. Br. of Calyceraceae from Australia, Bolivia, Brazil, Chile, Mexico, and the United States. These include the first reports and new numbers for the basal members of the Barnadesieae and Mutisieae. First reports are provided for 45 taxa, including five genera, Dasyphyllum HBK (2n = 54), Dithyrostegia A. Gray (2n = 14), Epitriche Turcz. (2n = 10), Revealia R. M. King & H. Rob. (2n = 22), and Stifftia J. C. Mikan (2n = 54). In addition, new chromosome numbers are established at the generic level in the genera Barnadesia Mutis (2n = 54), Eremanthus Less. (2n = 34, 30), Macvaughiella R. M. King & H. Rob. (2n = 24), and Trichocline Cass. (2n = 38), and in 12 additional species. Remaining counts augment and agree with previously reported determinations. The base chromosome numbers of Calyceraceae and Asteraceae are discussed in published records and our original counts.

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.

Background The transposable element (TE) content of the genomes of plant species varies from near zero in the genome of Utricularia gibba to more than 80 % in many species. It is not well understood whether this variation in genome composition results from common mechanisms or stochastic variation. The major obstacles to investigating mechanisms of TE evolution have been a lack of comparative genomic data sets and efficient computational methods for measuring differences in TE composition between species. In this study, we describe patterns of TE evolution in 14 species in the flowering plant family Asteraceae and 1 outgroup species in the Calyceraceae to investigate phylogenetic patterns of TE dynamics in this important group of plants. Results Our findings indicate that TE families in the Asteraceae exhibit distinct patterns of non-neutral evolution, and that there has been a directional increase in copy number of Gypsy retrotransposons since the origin of the Asteraceae. Specifically, there is marked increase in Gypsy abundance at the origin of the Asteraceae and at the base of the tribe Heliantheae. This latter shift in genome composition has had a significant impact on the diversity and abundance distribution of TEs in a lineage-specific manner. Conclusions We show that the TE-driven expansion of plant genomes can be facilitated by just a few TE families, and is likely accompanied by the modification and/or replacement of the TE community. Importantly, large shifts in TE composition may be correlated with major of phylogenetic transitions. Electronic supplementary material The online version of this article (doi:10.1186/s12864-015-1830-8) contains supplementary material, which is available to authorized users.

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.

This taxonomic treatment is the first in the genus Calycera based on a detailed study of morphology and a critical analysis of species boundaries. In this revision, nine species and four varieties are recognized, one new combination (C. crassifolia var. spinulosa), nine new synonymies (Anomocarpus axillaris, Boopis integrifolia, Calycera boopidea, C. crenata, C. foliosa, C. intermedia, C. pulvinata f. cauligera, C. squarrosa, and Gymnocaulus viridiflorus), six new lectotypifications (for Anomocarpus, Boopis gracilis, Calycera crenata, C. pulvinata f. cauligera, C. sessiliflora, and C. sympaganthera), and one neotypification (for Calycera involucrata) are established. Updated morphological descriptions (including an emended description of C. herbacea var. sinuata) and geographical distributions are included for each species. Analyses of their life forms and inflorescence, flower, and fruit morphology are presented. 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: 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: 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

ISSN 0046-0262. Previous molecular sequence and restriction site data have confirmed subfamily Barnadesioideae as basal within Asteraceae. The subfamily consists of 88 species in nine genera: Arnaldoa, Barnadesia, Chuquiraga, Dasyphyllum, Doniophyton, Duseniella, Fulcaldea, Huarpea, and Schlechtendalia. Two morphological cladistic analyses among genera or subgeneric groups within this subfamily have been completed previously, with contrasting results. Because of the importance of understanding relationships in this group of early Asteraceae, especially for insights on character evolution within the family, comprehensive morphological cladistic analyses were conducted at the specific level. Initial investigations utilising 31 characters with all 88 species led eventually to curtailed studies with 52 species. Five different outgroup hypotheses were employed: Acicarpha laxa, A. spathulata, Calycera leucanthema, C. spinulosa (all Calyceraceae), and Schlechtendalia luzulaefolia (Barnadesioideae; as functional outgroup). The results confirm Schlechtendalia as basal within Barnadesioideae. Monophyly is also confirmed for all genera, sections Chuquiraga and Acanthophyllae of Chuquiraga, and subgenus Archidasyphyllum of Dasyphyllum. All cladograms show two major groups (although statistical support values are low): (1) Chuquiraga, Doniophyton and Duseniella; and (2) Arnaldoa, Dasyphyllum, Fulcaldea, Huarpea and Barnadesia. Although bilabiate (4 + 1) corollas occur in Schlechtendalia, tubular corollas are believed plesiomorphic for the subfamily because they are pervasive in Calyceraceae as well as in all other basal Barnadesioideae. Pollen with no shallow depressions also are hypothesised as plesiomorphic for Barnadesioideae, even though one depression occurs in Schlechtendalia. A southern South American origin of Barnadesioideae (and the entire family) is re-emphasised. The previously hypothesised ancient lineage of Dasyphyllum subg. Archidasyphyllum, consisting of large trees, appears more derived in the new analyses.

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.

A comparison of cytological characters of the selected species of Calyceraceae (Acicarpha) and Dipsacaceae (Dipsacus, Scabiosa) is given, including chromosome numbers, condensing behavior at prophase and description of interphase nuclei. The results corroborate the isolation of Calyceraceae from the Dipsacales s.str. The cytological data are discussed with respect to previous morphological and molecular results. Karyotype description and banding patterns (C-banding and fiuorochromes) are described here for the first time for the genus Acicarpha.

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.

La familia Calyceraceae es un pequeño grupo ca. 54 especies endémicas Sudamérica austral. La mayoría de ellas se distribuye a lo largo de la cordillera de los Andes y Patagonia a partir del sur de Perú, aunque algunas especies presentan distribución chaqueña, alcanzando la costa atlántica y algunos enclaves de las selvas paranaense y atlántica en Brasil. Las Calyceraceae están estrechamente relacionadas con las Asteraceae, una de las cinco familias más ricas en especies de Angiospermas y una de las familias botánicas con mayor importancia económica. El subclado Calyceraceae + Asteraceae comparte varias sinapomorfías: las flores dispuestas en inflorescencias capituliformes rodeadas por brácteas involucrales, las anteras connadas en un anillo que rodea el estilo, con dehiscencia introrsa y presentación secundaria de polen, el polen espinulado o liso con concavidades intercolpares, ovarios uniloculares con un único óvulo, y cipselas con un cáliz persistente modificado. Recientes análisis filogenéticos han demostrado que el subclado Calyceraceae + Asteraceae queda incluido en el clado MGCA (Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae), un grupo monofilético con alto nivel de soporte. Las Calyceraceae se distinguen de las Asteracae principalmente por su óvulo péndulo (erecto en las Asteraceae), el estigma indiviso, punctiforme o capitado (dividido en dos ramas en las Asteraceae, que a veces no se separan simiulando un estigma indiviso), y la presencia de un tubo estaminal más o menos desarrollado (muy raro en las Asteraceae). Si bien las Calyceraceae se distinguen claramente como familia, no sucede lo mismo con sus géneros. Diversos problemas de interpretación de su morfología, más la variación morfológica de algunas de sus especies con amplia distribución geográfica, han desdibujado los límites entre los géneros (todos ellos basados en taxonomía alfa). Así, la familia Calyceraceae necesita una revisión de su morfología, de la taxonomía de sus especies y una reevaluación de sus géneros dentro de un marco filogenético, que permita proponer una clasificación natural y desarrollar hipótesis de evolución morfológica. De este modo, se propuso realizar una revisión taxonómica detallada de todas las especies de esta familia, más un análisis filogenético exploratorio combinando caracteres morfológicos y moleculares. Para la revisión taxonómica detallada, se efectuó una recopilación y revisión exhaustiva del material bibliográfico relacionado con la diversidad y taxonomía de la familia (catálogos, floras, tratamientos taxonómicos parciales y protologos), se procedió a la búsqueda y reconocimiento de ejemplares tipo, se estudiaron plantas vivas en su hábitat; se examinaron especímenes de varios herbarios, se observaron caracteres vegetativos y reproductivos de los ejemplares de herbario y de plantas vivas en su hábitat de las especies conocidas de la familia Calyceraceae. Como resultados de la revisión taxonómica, se reconocieron 47 especies distribuidas en los 6 géneros tradicionales. Cada especie se delimitó claramente, actualizando las listas sinonímicas, efectuando las lectotipificaciones necesarias y realizando descripciones ampliadas y detalladas para cada una de ellas. Además, se señalaron sus respectivas áreas de distribución geográfica con comentarios acerca de su hábitat y se presentaron discusiones acerca de los caracteres morfológicos entre especies similares y cómo diferenciarlas. Se publicaron dos especies nuevas para ciencia y una nueva combinación; se establecieron 25 nuevas sinonimias y se realizaron 21 lectotipificaciones y una neotipificación. Finalmente se confeccionó una clave artificial para identificar las 47 especies reconocidas de Calyceraceae. Para el análisis filogenético exploratorio, se realizó un el análisis cladístico de las 47 especies de la familia Calyceraceae reconocidas en esta tesis (con algunas excepciones dependiendo de la disponibilidad de datos para unas pocas especies raras). La elección de los grupos externos se basó en la selección de taxones hermanos de Calyceraceae, pertenecientes al clado MGCA. Se identificaron 37 caracteres exomorfológicos variables entre las especies y se definieron sus estados homólogos. Para los datos moleculares se trabajó con material coleccionado en sílica gel o material de herbario de 35 taxones, representando todos los géneros de Calyceraceae. También se obtuvieron secuencias de Genbank. Se amplificaron, mediante la reacción en cadena de la polimerasa (PCR), secuencias nucleotídicas correspondientes a espaciadores intergenes del cloroplasto: trnS–trnG y trnH–psbA. El análisis cladístico se basó en el criterio de máxima parsimonia. Se realizaron análisis con datos morfológicos, moleculares y combinados. Para cada análisis se obtuvieron árboles de consenso estricto y de mayoría (50%). Para evaluar la homoplasia de los caracteres en los árboles se obtuvieron los índices de consistencia y de retención. Para calcular el soporte de las ramas en los árboles óptimos se usó Jackknife Parsimony. Para calcular el soporte de las ramas en los árboles óptimos resultantes del análisis morfológico se usó Bremer support. De acuerdo con los resultados del análisis filogenético, la estructura de la inflorescencia de tipo cefalodio se agrega a las sinpomorfías de ls Calyceraceae. De los seis géneros tradicionalmente aceptados, sólo Acicarpha es monofilético; Boopis, Calycera, Gamocarpha, Moschopis y Nastanthus son géneros polifiléticos. La mayoría de los caracteres morfológicos son altamente homoplásicos. Con excepción de la forma columnar del receptáculo y la presencia de flores estaminadas en el centro de la inflorescencia (sinapomorfías de Acicarpha), los caracteres utilizados en taxonomía clásica para delimitar los géneros de Calyceraceae no permiten delimitar grupos naturales. Sin embargo, surgieron algunos pocos nuevos caracteres morfológicos con valor taxonómico: la presencia de pelos (especialmente en el pedúnculo y el involucro), la ubicación de los nectarios, la presencia de una secreción glutinosa en los tallos y las hojas, y en menor medida la forma de la base de las anteras, la nervadura central de los sépalos. Otro caracteres con valor taxonómico en ls Asteraceae, como el ápice de las anteras, las brácteas del receptáculo, la proporción del hipanto y de la porción lobada de la corola (éstos dos últimos como indicadores de la estructura general de la corola), resultaron altamente homoplásicos y no permitieron delimitar grupos naturales. Se reconocen dos grandes grupos monofiléticos dentro de Calyceraceae, especies con glándulas nectaríferas estaminales (clado A) y especies con glándulas nectaríferas hipantiales (clado B). Dentro del clado A se reconocen dos grupos monofiléticos: el clado 1, integrado por el género Acicarpha junto con su especie hermana Boopis anthemoides; y el grupo de plantas formado por las especies “Glutinosas” del clado 2 junto con su especie hermana Moschopis leyboldi y las dos variedades de Boopis gracilis (en su mayoría de distribución austral). Dentro del clado B se reconocen tres grupos monofiléticos: el grupo de especies “Pubescentes”, que incluye especies de los géneros Boopis y Calycera (clado 3); el clado 4, que incluye a Moschopis monocephala y algunas especies del género Calycera; y el clado 5, que incluye varias especies de los géneros Nastanthus, Boopis y Gamocarpha. Todos los clados del análisis combinado, inclusive los de bajo soporte, presentan coherencia en la distribución geográfica de sus especies. Los resultados del análisis filogenético sugieren que son necesarios cambios profundos en la taxonomía de la familia, para delimitar nuevamente y enmendar los géneros que la integran., Facultad de Ciencias Naturales y Museo

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 ...