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Calcium oxalate (CaOx) is one of the most common bio-mineral in extant plants and is believed to serve a variety of functions such as calcium storage and herbivore defense. However, traces of CaOx crystals have rarely been identified in fossil plants, and they are primarily known from fossil gymnosperms, where empty cavities of former CaOx crystals or ghost crystals have been reported from leaf cuticles of some Late Cretaceous and Cenozoic conifers. Here we investigate fossil angiosperm leaves from the late Oligocene Rott Fossil Lagerstätte and report ghost crystals of various shapes, sizes and topology (distribution patterns), and cavities. These micromorphological structures of fossil leaves are compared to CaOx deposits in leaves of extant plants: globular structures in fossil leaves resemble CaOx druses (crystal aggregates) in fresh leaves in size and distribution; and angular or brick-shaped structures in the vascular system of fossil leaves closely resemble prismatic CaOx crystals in the vascular system of extant leaves in both size and topology. Chemically, CaOx druses have survived fossilization as cavities only, and were replaced by organic matter and ghost minerals containing Ca, Si, Al, S, and Fe. The identification of former CaOx remains in leaf fossils provides novel insights on the fate of plant bio-minerals during fossilization. More importantly, it provides an additional aspect of the ecophysiology of fossil plants thus improving the accuracy of palaeoecological reconstructions and can provide a broader perspective on the evolution of CaOx and their rule in plant ecology across geological timescales. Alternative interpretations of the fossil microstructures are discussed but ruled out.

Biomineralization is a common phenomenon in plants and has been shown to be chemically, functionally and topologically diverse. Silica and calcium carbonate have long been known as structural plant biominerals and calcium phosphate (apatite)–long known from animals–has recently been reported. Strikingly, up to three different biominerals may occur in a single trichome in, e.g., Urticaceae and Loasaceae, and in combination with organic compounds, can form organic/inorganic composite materials. This article presents an extension of previous studies on the distribution of these biominerals in Loasaceae trichomes with a focus on their spatial (three-dimensional) distribution and co-localization with organic substances. Light microscopy and scanning electron microscopy with high-resolution EDX element analyses of sample surfaces and sections illustrate the differential distribution and composition of the different biomineral phases across cell surfaces and cell walls. Raman spectroscopy additionally permits the identification of organic and inorganic compounds side by side. All three biominerals may be found in a nearly pure inorganic phase, e.g., on the plant surfaces and in the barbs of the glochidiate trichomes, or in combination with a larger proportion of organic compounds (cellulose, pectin). The cell lumen may be additionally filled with amorphous mineral deposits. Water-solubility of the mineral fractions differs considerably. Plant trichomes provide an exciting model system for biomineralization and enable the in-vivo study of the formation of complex composite materials with different biomineral and organic compounds involved.

International audience; Multiple family-level subdivisions of Boraginales have been proposed in the past. The relationships of several constituent genera have been enigmatic, including Codon (Codonaceae), Hoplestigma (Hoplestigmataceae), Pholisma (Lennoaceae), Vahlia (Vahliaceae), and Wellstedia (Wellstediaceae), all of which are included in the present study. We present a molecular analysis with four chloroplast loci, including 89 ingroup taxa and a broad outgroup sampling in the asterids. The genus Vahlia is excluded from Boraginales and appears to represent an early branching lineage of Lamiales. The study provides a well supported topology for the relationships within Boraginales, including all of the genera with previously unclear relationships. Within Boraginales, two major clades are recognized, with “herbaceaous” Boraginales I resolved as [Codonaceae,[Wellstediaceae,[Boraginaceae]]] and “woody” Boraginales II resolved as [Hydrophyllaceae I,[Hydrophyllaceae II,[Heliotropiaceae,[Cordiaceae,[Ehretiaceae,Lennoaceae]]]]. A close relationship between Ehretiaceae and Lennoaceae is well supported, but the exact placement of Lennoaceae remains unresolved. The Cordiaceae lineage includes the monotypic genus Coldenia and the aberrant western and central African genus Hoplestigma. Woody Boraginales II are retrieved in two highly supported clades. Hydrophyllaceae are retrieved in two separate clades, but with poor support. There appear to be clear morphological progressions in vegetative, floral, and fruit morphology in both major Boraginales lineages. Thus capsular fruits are found in the first branching lineages of both clades, whereas reduced seed numbers in indehiscent fruits predominate in the more derived phylogenetic positions. Based on these results, we advocate the recognition of eight morphologically well defined clades in the order, namely Boraginaceae s.str., Codonaceae, Cordiaceae (incl. Coldenia and Hoplestigmataceae), Ehretiaceae (incl. Lennoaceae), Heliotropiaceae, Hydrophyllaceae I and Hydrophyllaceae II, and Wellstediaceae.

Loasaceae subfam. Loasoideae are a nearly exclusively American plant group with a center of diversity in Peru. Numerous new taxa have been described over the past decades; one of the most striking discoveries was that of the narrowly endemic

Many hummingbird-pollinated plant species evolved from bee-pollinated ancestors independently in many different habitats in North and South America. The mechanisms leading to these transitions are not completely understood. We conducted pollination and germination experiments and analyzed additional reproductive traits in three sister species pairs of which one species is bee- and the other hummingbird-pollinated. All hummingbird-pollinated species showed higher seed set and germination rates in cross-pollinated than in self-pollinated flowers. In the self-compatible, bee-pollinated sister species this difference did not exist. As expected, seed set and germination rate were higher after cross-pollination in the largely self-incompatible genus

Stachydeae, comprising c. 470 species, are one of the most diverse and taxonomically puzzling groups in Lamioideae. In the present study, the phylogenetic relationships in the Eurystachys clade (a phylogenetic name for all genera attributed to Stachydeae except Melittis) were reconstructed utilizing nuclear ribosomal DNA sequences (nrETS, 5S-NTS) from 148 accessions in 12 genera. Our phylogenetic results recovered Stachys as paraphyletic with numerous traditionally recognized genera nested in it. A broadly defined Eurystachys clade, however, was monophyletic. Unlike previous studies, the present study was able to resolve the group into 12 well-supported clades, named here as (1) Eriostomum, (2) Stachys, (3) Prasium, (4) Setifolia, (5) Distantes, (6) Burgsdorfia, (7) Hesiodia, (8) Empedoclia, (9) Sideritis, (10) Marrubiastrum, (11) Swainsoniana and (12) Olisia. These 12 clades were formally named in a phylogenetic nomenclature for the Eurystachys clade. Several infrageneric units were retrieved as monophyletic, namely Sideritis sections Burgsdorfia, Empedoclia and Hesiodia, Sideritis subgenus Marrubiastrum and Stachys sections Eriostomum (including Stachys section Mucronata) and Setifolia. The findings of this study also provide the basis for a future formal classification, with two options: (1) splitting of the Eurystachys clade into 12 monophyletic genera, all of them based on pre-existing genus names and redefined to encompass additional taxa, but without clear morphological apomorphies; or (2) lumping of all segregates into a broadly defined Stachys, including widely recognized and well-defined segregates such as Prasium and Sideritis.

The Global Strategy for Plant Conservation (GSPC) was established by the Conference of Parties in 2002 to decrease the loss of plant diversity, reduce poverty and contribute to sustainable development. To achieve this overarching goal, the GSPC has established a series of targets, one of which is to ensure that plant diversity is well understood, so that it can be effectively conserved and used in a sustainable manner. Brazil hosts more than 46,000 species of plants, algae and fungi, representing one of the most biodiverse countries on Earth, and playing a key role in the GSPC. To meet the GSPC goals of Target 1 and facilitate access to plant diversity, Brazil committed to preparing the List of Species of the Brazilian Flora (2008-2015) and the Brazilian Flora 2020 (2016-present). Managing all the information associated with such great biodiversity has proven to be an extremely challenging task. Here, we synthesize the history of these projects, focusing on the multidisciplinary and collaborative approach adopted to develop and manage the inclusion of all the knowledge generated though digital information systems. We further describe the methods used, challenges faced, and strategies adopted, as well as summarize advances to date and prospects for completing the Brazilian flora in 2020.

Trichome biomineralization is widespread in plants but detailed chemical patterns and a possible influence of soil chemistry are poorly known. We explored this issue by investigating trichome biomineralization in 36 species of Mediterranean Brassicaceae from ultramafic and calcareous soils. Our aims were to chemically characterize biomineralization of different taxa, including metallophytes, under natural conditions and to investigate whether divergent Ca, Mg, Si and P-levels in the soil are reflected in trichome biomineralization and whether the elevated heavy metal concentrations lead to their integration into the mineralized cell walls. Forty-two samples were collected in the wild while a total of 6 taxa were brought into cultivation and grown in ultramafic, calcareous and standard potting soils in order to investigate an effect of soil composition on biomineralization. The sampling included numerous known hyperaccumulators of Ni. EDX microanalysis showed CaCO3 to be the dominant biomineral, often associated with considerable proportions of Mg—independent of soil type and wild versus cultivated samples. Across 6 of the 9 genera studied, trichome tips were mineralized with calcium phosphate, in Bornmuellera emarginata the P to Ca-ratio was close to that of pure apatite-calcium phosphate (Ca5(PO4)3OH). A few samples also showed biomineralization with Si, either only at the trichome tips or all over the trichome. Additionally, we found traces of Mn co-localized with calcium phosphate in Bornmuellera emarginata and traces of Ni were detected in trichomes of the Ni-hyperaccumulator Odontarrhena chalcidica. Our data from wild and cultivated plants could not confirm any major effect of soil chemistry on the chemistry of trichome biominerals. Hyperaccumulation of Ni in the plants is not mirrored in high levels of Ni in the trichomes, nor do we find large amounts of Mn. A comparison based on plants from cultivation (normal, calcareous and serpentine soils, Mg:Ca-ratios ca 1:2 to 1:20) shows at best a very weak reflection of different Mg:Ca-ratios in the mineralized trichomes. The plants studied seem to be able to maintain highly conserved biomineralization patterns across a wide range of soil chemistries.

The Atacama Desert harbors a unique arid-adapted flora with a high degree of endemism, the origin of which is poorly understood. In the Atacama Desert, Zygophyllaceae is represented by five endemic species: one member of Zygophylloideae: Fagonia chilensis; and four members of Larreoideae: Bulnesia chilensis and Porlieria chilensis, the only representatives in the Atacama Desert of genera with disjunct distributions between Argentina, Peru and Chile; and monotypic endemic genera Metharme lanata and Pintoa chilensis. Zygophyllaceae are thus a particularly suitable group for studying the historical assembly of the Atacama Desert flora as each of these species may represent independent biogeographical events. We made use of published as well as original plastid DNA sequences (rbcL, trnL-trnF & trnS-trnG) to reevaluate the phylogenetic relationships of the Atacama Zygophyllaceae. Bayesian divergence time estimates as implemented in BEAST2 and ancestral area reconstruction with the Dispersal Extinction Cladogenesis approach using BioGeoBEARS were applied to infer ancestral ranges. We compiled the most complete data set of Larreoideae to date with 25 of 28 species. Bulnesia rivas-martinezii from Bolivia forms a clade with Pintoa chilensis from the Atacama Desert, rendering the genus Bulnesia paraphyletic. Most representatives of Zygophyllaceae colonized the Atacama Desert during the Miocene, and only Fagonia dispersed more recently. The colonization history of the Atacama Desert in South America is reflected by three individual distribution patterns or floristic elements. The presence of Bulnesia, Pintoa, and Metharme is best explained by Andean vicariance, while the southern Atacama Desert representative, Porlieria chilensis, has a continuous distribution into central Chile from where it probably dispersed further north. The only South American Fagonia species (F. chilensis) likely colonized the Chilean-Peruvian Coastal Desert via long distance dispersal from North America.

Cynoglossinae with ca. 200 spp. is one of the taxonomically most challenging subtribes of tribe Cynoglosseae with regard to generic delimitations. Here, we present a phylogenetic analysis based on nuclear ITS and cpDNA trnL-F and rps16 sequences of 270 accessions including 102 newly sequenced ones, representing all currently recognized genera of Cynoglossinae to clarify relationships among Cynoglossum and allied genera. We conducted maximum likelihood, maximum parsimony, and Bayesian inference analyses on separate and combined datasets. Cynoglossinae is recovered as a monophylum of closely related genera. Subtribe Cynoglossinae falls into three major groups: (1) Clade I including Microparacaryum spp., (2) Clade II comprising two species of Lindelofia, several Cynoglossum and Paracynoglossum spp., and (3) a larger clade (Clades III + IV) consisting of Solenanthus, Rindera, Trachelanthus, Paracaryum, Pardoglossum, Mattiastrum, and the remainder of Cynoglossum. None of the genera were monophyletic in the combined nuclear-plastid and nuclear datasets, except for Microparacaryum. A close relationship between Solenanthus apenninus and Cynoglossum officinale is shown, which suggests transferring S. apenninus to Cynoglossum. Moreover, Pardoglossum atlanticum, the type of the genus, is placed within Mediterranean Cynoglossum corroborating its synonymy under Cynoglossum. Lindelofia olgae is distantly related to the type of the genus (L. longiflora), but more closely connected with Solenanthus core-group. The majority of species currently recognized as members of Rindera do not form a clade with the type of the genus (R. tetraspis). The genus Paracaryum as currently circumscribed is paraphyletic and includes some species of Mattiastrum and Rindera.

Extreme arid conditions in the Atacama Desert in northern Chile have created a unique vegetation almost entirely restricted to the desert margins along the coast of the Pacific Ocean and the Andean range. In this study we provide data on the desert vegetation along elevational gradients at four localities from the western Andean slopes, between 19° and 21° S. Additionally, zonation of floristic data was explored. Three altitudinal zones could be classified and described in detail for each locality. Conspicuously divergent floras in the Atacama Desert have been recorded in the coastal ‘lomas formations’ and in the Andean desert vegetation, separated by a narrow band of absolute desert. In this study, we investigate the floristic relationships between both regions by implementing similarity analyses for 21 localities from the coastal and Andean deserts in northern Chile. Our results show a drastic east-west divergence in pairwise floristic similarity, which is in stark contrast to a weaker north-south divergence. A biotic barrier, preventing plant exchange from east to west and vice versa, imposed by the hyperarid conditions of the absolute desert, is one possible explanation for this finding. Moreover, the coastal and Andean deserts likely represent ecologically divergent habitats, e.g., in rainfall seasonality. Essential differences in factors determining plant life between both regions have probably contributed to a divergent evolution of the floras. Both explanations–ecological divergence and ecogeographical isolation—are not mutually exclusive, but likely complementary. We also combined floristic data from northern Chile and southern Peru. Similarity analyses of this combined dataset provide first floristic evidence for the existence of a biotic north-south corridor along the western slope of the Andes. Sub-Andean distributions of several species are discussed in the light of floristic connectivity between the Peruvian and Chilean Andean floristic clusters.

Several plant clades show a characteristic disjunction between the southern Andes (SA) and southern Brazil (SB). This disjunction has been explained by Neogene geologic and climatic changes, especially due to the Andean uplift, the Mid-Miocene marine transgressions, and the development of Patagonian aridity. We compared phylogeny-derived divergence times and realized climatic niches for ten clades disjunct between the SA and the SB, and compared them with the timing proposed for the above-mentioned geologic and climatic events. Divergence times obtained for the node corresponding to the SA-SB split ranged from 27 to 1 Ma. Climatic niche overlap between SA and SB is generally low and niche equivalency (but not similarity) between SA and SB is significantly lower than expected by chance. Precipitation tends to be higher in SA and SB than in the intervening areas, where no species of the disjunct clades occur, but this trend does not apply to all clades. Clades ages correlate to differences in minimum temperature of coldest month and mean temperature of coldest quarter. Our results indicate that while different Neogene climatic and geological events can explain the disjunction of most clades, there is no single explanation for all of them: SA-SB disjunctions go back to different clade histories. Climatic niche shifts could explain the generally low climatic overlap and lack of niche equivalency between SA and SB, and might have played a central role in the biogeographic history of the SA-SB disjunction. A proper test of this hypothesis will require measuring differences in the fundamental niches of the species—which is beyond the scope of this study.

Tribe Marrubieae (Lamiaceae: Lamioideae) comprises about 91 species classified in four genera distributed from Europe to West and Central Asia and North Africa. Here, we present a phylogenetic analysis of cpDNA (rpl32-trnL, rps16, trnL intron, trnL-trnF spacer) and nuclear ribosomal (ITS) sequence data of 68 accessions representing all genera and most sections of Marrubieae. Our taxon sampling covers is dense for the genera Ballota and Marrubium, including a total of 60 of 85 currently recognized taxa (species and subspecies). Parsimony and Bayesian analyses of individual nrITS and cpDNA markers, as well as the combined plastid dataset, produced largely congruent trees. Our results confirm the monophyly of tribe Marrubieae. All analyses retrieved four well supported clades in the tribe. The phylogenetic position of the genera Acanthoprasium and Moluccella is consistent with previous studies and both are recovered as monophyletic. The most important result of the present study is the splitting of the genus Ballota, in its traditional concept, into two well-supported clades matching well with the genera Ballota and Pseudodictamnus. Moreover, Marrubium, in its current circumscription, is paraphyletic, but will become monophyletic by transferring Ballota deserti to this genus. In conclusion, we propose the recognition of five genera in tribe Marrubieae: Acanthoprasium, Ballota, Moluccella, Marrubium and the resurrected segregate Pseudodictamnus. The required new combinations are proposed.

Premise of the study Stinging hairs are striking examples of plant microengineering-the plant equivalent of the hypodermic syringe. The requisite mechanical properties are mostly achieved by cell wall mineralization. Stinging hairs of Urtica dioica (Urticaceae) are known to be mineralized with silica and calcium carbonate and those of Loasaceae also with calcium phosphate, but no comparative study has been provided across different taxa with stinging hairs. Methods Light microscopy and scanning electron microscopy (SEM) with cryo-SEM and energy-dispersive x-ray spectroscopy were used to analyze morphology and biomineralization of stinging hairs of 43 species from the families Caricaceae, Euphorbiaceae, Loasaceae, Namaceae, and Urticaceae. Key results Stinging hair morphology is similar across the taxa studied, in striking contrast to the divergent patterns of biomineralization. Trichome bases are mostly calcified, sometimes silicified, the shafts are mostly calcified, and the apices silicified (Urticaceae), and contain calcium phosphate (Caricaceae, Namaceae), both silica and calcium phosphate (some Loasaceae), or no minerals (Cnidoscolus, Euphorbiaceae). Some stinging hairs are superficially thinly coated with silica over a cell wall otherwise mineralized with calcium carbonate or calcium phosphate. Conclusions Mineralization patterns are surprisingly diverse and involve three different biominerals deposited in different parts of individual trichomes with calcium phosphate a common component. The physical properties of different wall regions of the stinging trichomes are thus fine-tuned to optimize their function via modulation of wall thickness and differential element deposition. Similar function is apparently achieved through divergent wall compositions.

Premise of research. Heliotropiaceae (Boraginales) are morphologically readily defined by their peculiar floral morphology, especially their conspicuous stigmatic heads. Heliotropium L. is the larg...

Plant volatile organic compounds (pVOCs) are being recognized as an important factor in plant–environment interactions. Both the type and amount of the emissions appear to be heavily affected by climate change. A range of studies therefore has been directed toward understanding pVOC emissions, mostly under laboratory conditions (branch/leaf enclosure). However, there is a lack of rapid, sensitive, and selective analytical methods, and therefore, only little is known about VOC emissions under natural, outdoor conditions. An increased sensitivity and the identification of taxon‐specific patterns could turn VOC analysis into a powerful tool for the monitoring of atmospheric chemistry, ecosystems, and biodiversity, with far‐reaching relevance to the impact of climate change on pVOCs and vice versa. This study for the first time investigates the potential of ion mobility spectrometry coupled to gas‐chromatographic preseparation (GC‐IMS) to dramatically increase sensitivity and selectivity for continuous monitoring of pVOCs and to discriminate contributing plant taxa and their phenology. Leaf volatiles were analyzed for nine different common herbaceous plants from Germany. Each plant turned out to have a characteristic metabolite pattern. pVOC patterns in the field would thus reflect the composition of the vegetation, but also phenology (with herbaceous and deciduous plants contributing according to season). The technique investigated here simultaneously enables the identification and quantification of substances characteristic for environmental pollution such as industrial and traffic emissions or pesticides. GC‐IMS thus has an enormous potential to provide a broad range of data on ecosystem function. This approach with near‐continues measurements in the real plant communities could provide crucial insights on pVOC‐level emissions and their relation to climate and phenology and thus provide a sound basis for modeling climate change scenarios including pVOC emissions.

Disjunct clades between the southern and the tropical Andes represent a biogeographical pattern that has not been studied. One of the plant groups showing this disjunction is the clade formed by the genera Archidasyphyllum, Arnaldoa and Fulcaldea (Asteraceae, Barnadesioideae). Archidasyphyllum is distributed in central and southern Chile and adjacent Argentina and is sister to the latter two, which in turn form a clade centered in northern Peru and southern Ecuador, with one species of Fulcaldea in Bahia, Brazil. The western American clades are separated by a distance of ca. 2500 km in direct line and have no representatives through the entire arid and hyperarid regions of the Pacific deserts and the dry Puna. We hypothesized that the Neogene origin of aridity in this intervening area might be responsible for this disjunction. To address this hypothesis, we estimated divergence times and ancestral range and quantified the climatic niches of the respective clades and compared them to each other and with the intervening area. Our results suggest a Miocene split of this clade from an ancestor previously distributed in the Central Andes, where the species of this clade are currently absent. Colonization of NE Brazil by Fulcaldea may have occurred during the Pliocene. The niche analysis rejects climatic niche conservatism and the intervening area is found to be climatically different from the current ranges occupied by either clade. We suggest that the global cooling trend in the Miocene and concomitant hyperaridity in the Pacific deserts and southern Central Andean highlands played a crucial role in the formation of this disjunct pattern and that the climatic niches of each clade have subsequently shifted in different directions.

Urtica L. (Urticaceae) is generally reported as a genus of monoecious and dioecious taxa. However, the gender information found in the literature does not at all reflect the actual diversity of gender patterns in Urtica. Dioecy appears to be truly absent from Urtica, but otherwise there has been a major diversification in the geometry of gender and no comparable patterns exist in other plant groups. Thus, we here define technical terms for all unique architectural types of monoecy found in Urtica and closely related genera and reconstruct the ancestral gender states in a Bayesian framework. Our studies are based on a near-comprehensive sampling, including 61 of the 63 Urtica species recognized. We report polygamy, two types of gynodioecy and five different architectural types of monoecy. A total of 15 switches appear to have taken place within the genus. Although gender characteristics have diversified strongly, they are relatively conserved within clades. Monoecy is the predominant sexual system within Urtica and specifically basiandrous monoecy (i.e. basal inflorescence branches of each individual male only, apical branches female) is the most widespread type, reported for 11 different clades. In particular, it characterizes the basally branching pilulifera-clade and the sister genus Zhengyia, and may thus represent the plesiomorphic condition for Urtica. Gender distribution and gross morphology appear to evolve largely independently from each other and gender distribution is largely independent of growth habit. However, polygamous taxa are most common amongst rhizomatous perennials (one-third of the taxa).

Made available in DSpace on 2020-12-10T17:39:01Z (GMT). No. of bitstreams: 0 Previous issue date: 2020-08-05 Royal Society International Exchanges Carlsberg Foundation Danish Research Council National Natural Science Foundation of China Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) US National Science Foundation National Parks Board, Singapore Deutsche Forschungsgemeinschaft Synthesys Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Edinburgh Botanic Garden (Sibbald) Trust M. L. MacIntyre Begonia Trust Scottish Government's Rural and Environmental Science and Analytical Services Division New Guinea is the world's largest tropical island and has fascinated naturalists for centuries(1,2). Home to some of the best-preserved ecosystems on the planet(3)and to intact ecological gradients-from mangroves to tropical alpine grasslands-that are unmatched in the Asia-Pacific region(4,5), it is a globally recognized centre of biological and cultural diversity(6,7). So far, however, there has been no attempt to critically catalogue the entire vascular plant diversity of New Guinea. Here we present the first, to our knowledge, expert-verified checklist of the vascular plants of mainland New Guinea and surrounding islands. Our publicly available checklist includes 13,634 species (68% endemic), 1,742 genera and 264 families-suggesting that New Guinea is the most floristically diverse island in the world. Expert knowledge is essential for building checklists in the digital era: reliance on online taxonomic resources alone would have inflated species counts by 22%. Species discovery shows no sign of levelling off, and we discuss steps to accelerate botanical research in the 'Last Unknown'(8). A catalogue of the vascular flora of New Guinea indicates that this island is the most floristically diverse in the world, and that 68% of the species identified are endemic to New Guinea. Royal Bot Gardens, London, England Nat Biodivers Ctr, Leiden, Netherlands Univ Michigan Herbarium, Ann Arbor, MI USA Univ Gottingen, Dept Systemat Biodivers & Evolut Plants, Albrecht von Haller Inst Plant Sci, Gottingen, Germany Royal Bot Garden Edinburgh, Edinburgh, Midlothian, Scotland Aarhus Univ, Dept Biosci, Aarhus, Denmark Univ Vermont, Dept Plant Biol, Pringle Herbarium, Burlington, VT USA Boise State Univ, Dept Biol Sci, Boise, ID 83725 USA Conservatoire & Jardin Bot Ville Geneve, Chambesy, Switzerland Univ Sydney, Sch Life & Environm Sci, Sydney, NSW, Australia Natl Museum Nat & Sci, Dept Bot, Tsukuba, Ibaraki, Japan Univ Minnesota, Dept Plant & Microbial Biol, St Paul, MN 55108 USA New Guinea Binatang Res Ctr, Madang, Papua N Guinea Guangxi Zhuang Autonomous Reg & Chinese Acad Sci, Guangxi Key Lab Plant Conservat & Restorat Ecol K, Guangxi Inst Bot, Guilin, Peoples R China Papua New Guinea Univ Technol, Lae, Papua N Guinea Herbarium Bogoriense, Bogor, Indonesia Univ Fed Mato Grosso do Sul, Tres Lagoas Campus, Tres Lagoas, Brazil Royal Bot Garden Sydney, Sydney, NSW, Australia Univ Papua, Fak Kehutanan, Manokwari, Indonesia Baden Penelitian & Pengembangan Daerah Prov Papua, Manokwari, Indonesia Missouri Bot Garden, St Louis, MO USA Papua New Guinea Forest Res Inst, Lae, Papua N Guinea Natl Bot Gardens Ireland, Glasnevin, Ireland Univ Zurich, Dept Systemat & Evolutionary Bot, Zurich, Switzerland Forest Res Inst Malaysia, Kepong, Malaysia Nat Hist Museum, Dept Life Sci, London, England Univ Bonn, Nees Inst Biodivers Plants, Bonn, Germany Martin Luther Univ Halle Wittenberg, Inst Biol, Herbarium, Halle, Germany Natl Pk Board, Nat Plants Ctr, Singapore, Singapore Natl Pk Board, Singapore Bot Gardens, Singapore, Singapore Univ Aberdeen, Sch Biol Sci, Aberdeen, Scotland Univ Philippines Manila, Coll Arts & Sci, Dept Biol, Manila, Philippines Kyoto Univ, Kyoto Univ Museum, Kyoto, Japan Sao Paulo State Univ, Inst Biosci, Rio Claro, Brazil Royal Bot Gardens Victoria, South Yarra, Vic, Australia Univ Melbourne, Sch BioSci, Parkville, Vic, Australia Univ Oxford, Dept Plant Sci, Oxford, England Queens Univ Belfast, Marine Lab, Portaferry, North Ireland Fern Res Fdn, Kerikeri, New Zealand Univ N Carolina, Wilmington, NC USA Museum New Zealand Papa Tongarewa, Wellington, New Zealand Western Michigan Univ, Dept Biol Sci, Kalamazoo, MI 49008 USA New Mexico State Univ, Dept Anim & Range Sci, Las Cruces, NM 88003 USA Univ Fed Vicosa, Dept Biol Vegetal, Vicosa, MG, Brazil Colorado State Univ, Dept Biol, Ft Collins, CO 80523 USA Trinity Coll Dublin, Sch Nat Sci, Dept Bot, Dublin, Ireland Univ Florida, Dept Biol, Gainesville, FL USA CSIRO Land & Water, Floreat, WA, Australia Univ Western Australia, Sch Biol Sci, Crawley, WA, Australia Chinese Acad Sci, Inst Bot, Beijing, Peoples R China Univ Minnesota, Bell Museum, St Paul, MN 55108 USA Univ Glasgow, Inst Biodivers Anim Hlth & Comparat Med, Glasgow, Lanark, Scotland Chinese Acad Sci, Chengdu Inst Biol, Chengdu, Peoples R China Leiden Univ, Inst Biol Leiden, Leiden, Netherlands Univ Zurich, Dept Evolutionary Biol & Environm Studies, Zurich, Switzerland Sao Paulo State Univ, Inst Biosci, Rio Claro, Brazil Royal Society International Exchanges: IE 170241 National Natural Science Foundation of China: 31570307 CNPq: 153430/2018-4 US National Science Foundation: DEB-1457366 US National Science Foundation: DEB-1754667 US National Science Foundation: DEB-0515678 US National Science Foundation: DEB-0816749 US National Science Foundation: DEB-0841885 Deutsche Forschungsgemeinschaft: LE 1826/3-1 Synthesys: GB-TAF-6305 FAPESP: 2015/13112-7 FAPESP: 2018/09379-6

Allium is a particularly species rich (more than 800 species) and economically important genus, with numerous taxonomic problems at all levels of classification. In this study, we try to uncover the phylogenetic relationships of the common leek Allium ampeloprasum based on selected samples of this species and its putative relatives in A. sect. Allium from Iran. The silica‐dried leaf samples of 56 accessions representing 23 species of Allium were sequenced; 53 sequences of nrDNA ITS, 35 sequences of plastid rps16 and 52 sequences of trnL–F were generated and additional accessions were extracted from GenBank in order to cover all recognized main lineages in the genus. Maximum Parsimony and Bayesian Inference generated similar trees, but the placement of A. ampeloprasum and its relatives differed slightly between the nuclear and the plastid phylogenies. In the nrITS tree, A. ampeloprasum is retrieved as a highly supported clade with A. iranicum, while in the combined plastid tree A. ampeloprasum formed a highly supported clade with A. vineale. This supports the hypothesis of a possible hybrid origin of A. ampeloprasum. Allium iranicum formed a clade in the plastid tree, but was resolved as paraphyletic in the nrITS tree, probably due to presence of multiple non‐concerted copies of nrITS. Close relationships are suggested between the following species: A. aznavense and A. wendelboi with A. talyschense, A. erubescens and A. rotundum with A. scorodoprasum and A. abbasii with A. phanerantherum.

The species-rich flora of Madagascar is well known for a range of unusual floral ecologies. One example is Impatiens section Trimorphopetalum with its unique combination of floral traits: small, spur-less, cup- or lip-shaped, greenish or brownish flowers. So far no hypotheses on floral function or pollination of this peculiar group have been proposed. We analysed six reproductive traits in relation to pollination syndromes for 34 Madagascan Impatiens species, including 18 species of section Trimorphopetalum plus six outgroup species, in a phylogenetic framework. Further, we present pollinator observations for one additional species of Trimophopetalum. All pollination syndromes occurring in the African species are also present in Madagascan Impatiens. In addition, species of Trimorphopetalum represent two unique floral types, possibly corresponding to two different types of fly pollination. The evolution of these flower types corresponds to a strong decrease in nectar production, flower display size, pollen grain and ovule number. Autogamy is found in one derived sub-clade of the otherwise largely pollinator-dependent Trimorphopetalum. We find evidence consistent with the evolution of brood-site deception and fungus mimicry in combination with fly pollination in one clade of Trimorphopetalum and the stepwise evolution of autogamy in the second clade. The evolution of these very different reproductive strategies may have been triggered by pollinator limitation in the dense, humid forest undergrowth of Madagascar.

The genus Urtica L. is subcosmopolitan, found on all continents (except Antarctica) and most extratropical islands and ranges from Alaska to Patagonia, Spitzbergen to the Cape and Camtschatka to the subantarctic islands. However, throughout its geographical range morphologically nearly indistinguishable species are found alongside morphologically quite disparate species, with the overall diversity of morphological characters extremely limited. The systematics of Urtica have puzzled scientists for the past 200years and no single comprehensive attempt at understanding infrageneric relationships has been published in the past, nor are species delimitations unequivocally established. We here provide the first comprehensive phylogeny of the genus including 61 of the 63 species recognized, represented by 144 ingroup accessions and 14 outgroup taxa. The markers ITS1-5.8S-ITS2, psbA-trnH intergenic spacer, trnL-trnF and trnS-trnG are used. The phylogeny is well resolved. The eastern Asian Zhengyia shennongensis T. Deng, D.G. Zhang & H. Sun is retrieved as sister to Urtica. Within Urtica, a clade comprising the western Eurasian species U. pilulifera L. and U. neubaueri Chrtek is sister to all other species of the genus. The phylogenetic analyses retrieve numerous well-supported clades, suggesting previously unsuspected relationships and implying that classically used taxonomic characters such as leaf morphology and growth habit are highly homoplasious. Species delimitation is problematical, and several accessions assigned to Urtica dioica L. (as subspecies) are retrieved in widely different places in the phylogeny. The genus seems to have undergone numerous dispersal-establishment events both between continents and onto different islands. Three recent species radiations are inferred, one in America centered in the Andes, one in New Zealand, and one in northern Eurasia which includes Urtica dioica s.str. sensu Henning et al. (2014). The present study provides the basis of a critical re-examination of species limits and taxonomy, but also of the dispersal ecology of this widespread plant group and an in-depth study of the three clades with recent radiations.

Aim To examine the historical biogeography of the Boraginales using molecular dating and ancestral area reconstruction. Location World-wide. Methods We constructed data sets that included all major clades of Boraginales and all orders of asterids using previously published sequences of four plastid markers (trnL-trnF, rps16, ndhF, rbcL). We estimated divergence times using a Bayesian uncorrelated, lognormal relaxed clock approach with four different fossil calibration schemes. Ancestral areas were reconstructed using maximum likelihood methods (Dispersal-Extinction-Cladogenesis). Results Boraginales originated during the Early to Late Cretaceous and started its diversification in the Late Cretaceous. The inferred ancestral area of Boraginales includes the Americas and Africa. The two major clades of Boraginales diversified during the Early Paleogene from African and American ancestors respectively. Early branching families in both clades (Codonaceae and Wellstediaceae in one clade and Hydrophyllacee and Namaceae in the other) may have remained restricted to their areas of origin. The other families started diversifying in several regions of the world during the Eocene (Boraginaceae s.str., Heliotropiaceae, Ehretiaceae) or later (Cordiaceae). Main conclusions Molecular dating and ancestral area reconstruction may be broadly consistent with the idea of a vicariant origin of the two major clades of Boraginales after the break-up of West Gondwana, followed by several independent trans-oceanic dispersal events into most areas of the world. However, uncertainty in both divergence times and ancestral area reconstruction do not rule out the possibility of an origin involving long-distance dispersal.

Autogamous species are usually distinguishable from xenogamous relatives by smaller flowers, fewer or even no floral rewards and lower pollen–ovule (P/O) ratios. Many Rhipsalis spp. are small flowered, selfing and include the most widespread species in Cactaceae. However, Rhipsalis also includes a large number of narrowly endemic species and is most diverse in the Atlantic rainforests of Brazil. To investigate the evolution of floral function and the correlation between floral function and range size, we analysed display size, floral reward and P/O ratios of Rhipsalis and its closest relatives, reconstructed ancestral traits and related these patterns to the distributions and range sizes of the species. Display size and sugar amount are reduced in subgenera Goniorhipsalis and Rhipsalis and secondarily increased in Phyllarthrorhipsalis, whereas the P/O ratio is decreased in subgenera Rhipsalis and Phyllarthrorhipsalis. We interpret this pattern as a switch from a predominantly xenogamous to an autogamous reproductive system, followed by a return to a predominantly xenogamous system. None of the floral parameters shows significant correlations with range size, except for display size. Nevertheless, those species with the smallest flowers, lowest sugar amounts per flower and lowest P/O ratios occur either outside southeastern Brazil and/or have comparatively large distribution ranges. Almost all Rhipsalis spp. occurring outside the Atlantic rainforests are restricted to the clade formed by subgenera Rhipsalis and Phyllarthrorhipsalis. Thus, we believe that the evolution of an autogamous reproduction system enabled this lineage of Rhipsalis to diversify and spread in the Atlantic rainforests, in the rest of the Neotropics and even spread to the Old World, where it is the only member of the family.

Pollen/ovule (P/O) ratios are often used as proxy for breeding systems. Here, we investigate the relations between breeding systems and P/O ratios, pollination syndromes, life history and climate zone in Balsaminaceae. We conducted controlled breeding system experiments (autonomous and active self-pollination and outcrossing tests) for 65 Balsaminaceae species, analysed pollen grain and ovule numbers and evaluated the results in combination with data on pollination syndrome, life history and climate zone on a phylogenetic basis. Based on fruit set, we assigned three breeding systems: autogamy, self-compatibility and self-incompatibility. Self-pollination led to lower fruit set than outcrossing. We neither found significant P/O differences between breeding systems nor between pollination syndromes. However, the numbers of pollen grains and ovules per flower were significantly lower in autogamous species, but pollen grain and ovule numbers did not differ between most pollination syndromes. Finally, we found no relation between breeding system and climate zone, but a relation between climate zone and life history. In Balsaminaceae reproductive traits can change under resource or pollinator limitation, leading to the evolution of autogamy, but are evolutionary rather constant and not under strong selection pressure by pollinator guild and geographic range changes. Colonisation of temperate regions, however, is correlated with transitions towards annual life history. Pollen/ovule-ratios, commonly accepted as good indicators of breeding system, have a low predictive value in Balsaminaceae. In the absence of experimental data on breeding system, additional floral traits (overall pollen grain and ovule number, traits of floral morphology) may be used as proxies.

Krameria lappacea (ratania) ha sido utilizada y reconocida como planta medicinal y tintura vegetal desde épocas precolombinas. En los últimos 15 años se han estudiado la biología, propagación y abundancia de ratania en Perú y desarrollado métodos que garanticen una recolección silvestre sostenible. El objetivo de este trabajo es investigar qué factores limitan la propagación natural y evaluar si una resiembra de semillas durante la recolección puede facilitar el establecimiento de plantas nuevas in situ. Al investigarse la germinación de las semillas y la propagación in situ pudo comprobarse mediante experimentos de invernadero que el entierro de las semillas es un factor limitante para la germinación de K. lappacea y que la tasa de germinación puede incrementarse significativamente (3 – 4x) al enterrar las semillas. Tanto la resiembra de las semillas en los hoyos realizados tras la cosecha, así como la resiembra más esparcida en parcelas seleccionadas, puede contribuir al reforzamiento de la regeneración natural. Las raíces secundarias que quedan en el suelo después de la cosecha forman un nuevo meristemo y dan lugar al desarrollo de nuevas plantas. El pre-tratamiento de las semillas (escarificación) no produce ningún efecto claro en el éxito de la germinación, sin embargo, esta práctica facilita la manipulación durante la resiembra, asegurando que únicamente semillas viables y completamente desarrolladas sean utilizadas posteriormente., Krameria lappacea (rhatany) is a well established medicinal and dye plant that is used since pre-Colombian times. In the last 15 years the biology, distribution and abundance of rhatany in Peru was investigated and methods for a sustainable wildcrafting were developed. The objective of this work was to investigate which factors limit the natural propagation and to explore whether the reseeding measures can contribute to the establishment of new plants in situ. Investigating the germination and in situ propagation, greenhouse trials showed that the seed burial is an important, limiting factor for the germination of K. lappacea und the germination rate can be increased (3—4x) by burying the seeds. Both the burial of the seeds during collection and re-sowing measures on selected lots can contribute to the rejuvenation of the population. Secondary roots, which are left in the ground during harvest, are showed to form new shoot meristems and develop into new plants. Pre-treatment (scarification) of the fruits does not clearly influence the germination success, but it facilitates the handling of the seeds during re-sowing and ensures the use of only fully developed, viable seeds.

Obvious movements of plant organs have fascinated scientists for a long time. They have been studied extensively, but few behavioural studies to date have dealt with them, and hardly anything is known about their evolution. Here, we present a large experimental dataset on the stamen movement patterns found in the Loasaceae subfam. Loasoideae (Cornales). An evolutionary transition from autonomous-only to a combination of autonomous and thigmonastic stamen movement with increased complexity was experimentally demonstrated. We compare the stamen movement patterns with extensive pollinator observations and discuss it in the context of male mating behavior. Thigmonastic pollen presentation via stamen movements appears to be a crucial component of floral adaptation to pollinator behaviour, evolving in concert with complex adjustments of flower signal, reward and morphology. We hypothesize that rapid adjustments of pollen presentation timing may play a significant role in the diversification of this plant group, representing a striking example for the evolutionary significance of plant behaviour.

Background and Aims Flowers of Geraniaceae and Hypseocharitaceae are generally considered as morphologically simple. However, previous studies indicated complex diversity in floral architecture including tendencies towards synorganization. Most of the species have nectar-rewarding flowers which makes the nectaries a key component of floral organization and architecture. Here, the development of the floral nectaries is studied and placed into the context of floral architecture. Methods Seven species from Geraniaceae and one from Hypseocharitaceae were investigated using scanning electron microscopy and light microscopy. Samples were prepared and processed using standard protocols. Key Results The development of the nectary glands follows the same trajectory in all species studied. Minor differences occur in the onset of nectarostomata development. The most striking finding is the discovery that a short anthophore develops via intercalary growth at the level of the nectary glands. This anthophore lifts up the entire flower apart from the nectary gland itself and thus plays an important role in floral architecture, especially in the flowers of Pelargonium. Here, the zygomorphic flowers show a particularly extensive receptacular growth, resulting in the formation of a spur-like receptacular cavity ('inner spur'). The nectary gland is hidden at the base of the cavity. Various forms of compartmentalization, culminating in the 'revolver flower' of Geranium maderense, are described. Conclusions Despite the superficial similarity of the flowers in Geraniaceae and Hypseocharitaceae, there is broad diversity in floral organization and floral architecture. While the receptacular origin of the spur-like cavity in Pelargonium had already been described, anthophore formation via intercalary growth of the receptacle in the other genera had not been previously documented. In the context of the most recent phylogenies of the families, an evolutionary series for the floral architecture is proposed, underscoring the importance of synorganization in these seemingly simple flowers.

BACKGROUND AND AIMS: Boraginales are often characterized by a dense cover of stiff, mineralized trichomes, which may act as a first line of defence against herbivores. Recent studies have demonstrated that the widely reported silica and calcium carbonate in plant trichomes may be replaced by calcium phosphate. The present study investigates mineralization patterns in 42 species from nine families of the order Boraginales to investigate detailed patterns of mineralization and the possible presence of a phylogenetic signal in different mineralization patterns. METHODS: The distribution of biominerals was analysed by scanning electron microscopy (SEM) including cryo-SEM and energy-dispersive X-ray analyses with element mapping. The observed distribution of biominerals was plotted onto a published phylogeny of the Boraginales. Three colours were selected to represent the principal elements: Si (red), Ca (green) and P (blue). KEY RESULTS: Calcium carbonate was present in the mineralized trichomes of all 42 species investigated, silica in 30 and calcium phosphate in 25; multiple mineralization with calcium carbonate and silica or calcium phosphate was found in all species, and 13 of the species were mineralized with all three biominerals. Trichome tips featured the most regular pattern – nearly all were exclusively mineralized with either silica or calcium phosphate. Biomineralization of the trichome shafts and bases was found to be more variable between species. However, the trichome bases were also frequently mineralized with calcium phosphate or silica, indicating that not only the tip is under functional constraints requiring specific patterns of chemical heterogeneity. The complete absence of either silica or phosphate may be an additional feature with systematic relevance. CONCLUSIONS: This study demonstrates that complex, site-specific and differential biomineralization is widespread across the order Boraginales. Calcium phosphate, only recently first reported as a structural plant biomineral, is common and appears to be functionally analogous to silica. A comparison with the phylogeny of Boraginales additionally reveals striking phylogenetic patterns. Most families show characteristic patterns of biomineralization, such as the virtual absence of calcium phosphate in Cordiaceae and Boraginaceae, the triple biomineralization of Heliotropiaceae and Ehretiaceae, or the absence of silica in Namaceae and Codonaceae. The complex chemical and phylogenetic patterns indicate that trichome evolution and functionalities are anything but simple and follow complex functional and phylogenetic constraints.

Long-distance dispersal seems to be the main biogeographic event responsible for intriguing distribution patterns in plant groups in which sister taxa are separated by thousands of kilometers of distance across oceans and continents. The biotic and abiotic mechanisms behind such dispersal events are poorly understood and many attempts have been made to explain how plants can manage to disperse and survive these long journeys. The biogeographic history of Boraginaceae, a subcosmopolitan plant family with many disjunct clades, is here addressed and analyzed in the context of the different dispersal modes exhibited by the species. The lack of a clear pattern between the main dispersal events in Boraginaceae and the phylogenetic distribution of the dispersal modes indicates that no single dispersal mechanism can be associated with the events of dispersal in the family. Moreover, adaptations to different dispersal agents and unassisted dispersal modes in some clades might have promoted the diversification of Boraginaceae in various habitats across several continents. Our study reveals that long-distance dispersal is a very complex process that needs to be analyzed in the context of climatic and environmental changes and the response of plants and their dispersal vectors to these variable conditions.

Internal sample conductivity in scanning electron microscopy can be a valuable alternative to metal coating. Proton conductivity may be used for this purpose. Many solid materials with active hydrogen atoms, such as hydrogen- and ammonium-salts, organic acids, and even ice, are protonic conductors or semiconductors. Here we present a method to generate proton conductivity in dry biological materials. A simple treatment with hydrogen chloride gas or hydrochloric acid vapour for a few minutes provides sufficient conductivity for many samples. After a removal of excess hydrogen chloride vapour with a vacuum desiccator, the objects may be examined in the SEM without metal coating. The use of internally conductive samples extends the range of easy-to-perform SEM preparation techniques. It is advantageous for material contrast imaging of uncoated samples, and it can be used in combination with metal coating to enhance conductivity on difficult samples with complex overlapping surfaces, where simple metal coating does not reliably eliminate charging problems.

Even in times of “big data”, the holdings of local herbaria worldwide are of increasing value for taxonomic discoveries and phytogeographic analyses. Based on our research in Peruvian herbaria we present new records for three species of Symplocos (Symplocaceae) for Peru, which were previously known only from Southern Ecuador., Actualmente, incluso en la era de las grandes bases de datos, las colecciones de los herbarios locales de todo el mundo siguen representando invaluables fuentes de información para los descubrimientos taxonómicos y los análisis fitogeográficos. Con base en la investigación llevada a cabo en diferentes herbarios del Perú, presentamos nuevos registros de tres especies de Symplocos (Symplocaceae) para el Perú, las cuales eran conocidas anteriormente sólo para el sur de Ecuador.

The Atacama Desert in western South America is considered as one of the driest places on earth, but is nevertheless characterized by surprisingly high species richness and levels of endemism. The plant genus Cristaria (Malvaceae), with ca. 21 species, is one of the most diverse genera of the Atacama Desert, while the much less diverse sister genus Lecanophora (7 species) is found east of the Andes. Here, we use DNA sequence data and divergence time estimates in order to investigate the biogeographical history of the Atacama species of Cristaria. We further investigate a possible influence of Andean uplift and the subsequent onset of hyperaridity in the Atacama Desert on diversification times in Cristaria. We sequenced three plastid markers (ndhF, trnK(matK) & rpl16) for 19 species of Cristaria and two species of Lecanophora from the Atacama Desert and Argentina, respectively. Further, we included sequences of the same plastid regions from GenBank in order to get a comprehensive dataset of Malvoideae. Phylogenetic relationships were inferred using maximum likelihood and Bayesian analyses, and divergence times were estimated with BEAST2. Our results place the monophyletic genera Cristaria and Lecanophora as sister groups in a clade sister to the rest of Malveae. The split between these two lineages (~20 Ma) correlates with Andean uplift during the early Miocene, indicating a vicariant event. During the late Miocene, two Mediterranean members of Cristaria separated from the major Atacama clade. The subsequent diversification of the latter one correlates with the onset and subsequent temporal expansion of hyperarid conditions in the Atacama Desert since the late Miocene and during the Quaternary climate oscillations.

Recent studies all indicated that both the affinities and subdivision of Boraginaceae s.str. are unsatisfactorily resolved. Major open issues are the placement and affinities of Boraginaceae s.str. in Boraginales and the major clades of the family, with especially the large tribes Cynoglosseae and Eritrichieae repeatedly retrieved as non-monophyletic groups, and the doubtful monophyly of several larger genera, especially Cynoglossum and Omphalodes . The present study addresses and solves these questions using two plastid markers ( trnL – trnF , rps 16) on the basis of a sampling including 16 outgroup taxa and 172 ingroup species from 65 genera. The phylogeny shows high statistical support for most nodes on the backbone and on the individual clades. Boraginaceae s.str. are sister to African Wellstediaceae, Wellstediaceae–Boraginaceae s.str. is sister to African Codonaceae. Echiochileae are retrieved as sister to the remainder of Boraginaceae s.str., which, in turn, fall into two major clades, the Boragineae–Lithospermeae (in a well-supported sister relationship) and the Cynoglosseae s.l. (including Eritrichieae). Cynoglosseae s.l. is highly resolved, with Trichodesmeae (incl. Microcaryum , Lasiocaryum ) as sister to the remainder of the group. Eritrichieae s.str. ( Eritrichium , Hackelia , Lappula ) are resolved on a poorly supported polytomy together with the Omphalodes -clade (incl. Myosotidium , Cynoglossum p.p.), and the Mertensia -clade (incl. O. scorpioides , Asperugo ). The Myosotideae ( Myosotis , Trigonotis , Pseudomertensia ) are retrieved in a well-supported sister-relationship to the core-Cynoglosseae, the latter comprising all other genera sampled. Cynoglossum is retrieved as highly para- and polyphyletic, with a large range of generic segregates embedded in Cynoglossum , but other species of Cynoglossum are sister to Microula or to the American “Eritrichieae” ( Cryptantha and allied genera). Representatives of the genus Cynoglossum in its current definition are segregated onto six independent lineages, members of Omphalodes onto three independent lineages. At least 11 of the genera here sampled are deeply nested in other genera. The data show that individual details of nutlet morphology (e.g., winged margins, glochidia) are highly homoplasious. Conversely, a complex of nutlet characters (e.g., characters of the gynobase and cicatrix together with nutlet orientation and sculpturing) tends to circumscribe natural units. Geographical distribution of major clades suggests that the family originated in Africa and western Asia and radiated to eastern Eurasia, with several independent dispersal events into Australia and the New World.

PREMISE OF THE STUDY Plants of the family Loasaceae are characterized by a usually dense indument of various trichome types, including two basically different types of mineralized, unicellular trichomes (stinging hairs or setae and scabrid-glochidiate trichomes). Mineralized trichomes have long been known to have silicified or calcified walls, but recent studies demonstrated that trichomes of Loasaceae may also contain calcium phosphate. The current study investigates the distribution of different biominerals in the mineralized trichomes across several different taxa. METHODS Plants from cultivation were studied with scanning electron microscopy including energy dispersive x-ray analyses and element mapping. KEY RESULTS The vast majority of the 31 species investigated had at least two different biominerals in their trichomes, and 22 had three different biominerals in their trichomes. Thirty of the species had calcium phosphate in their trichomes. Loasa was mostly free of silica, but contained calcium phosphate in trichome tips and barbs, whereas calcium phosphate and silica were found in representatives of other genera of the family (Blumenbachia, Caiophora, Nasa). CONCLUSIONS Biomineralization is remarkably diversified between species, different trichome types and parts of the same trichome. Individual genera largely had different patterns of biomineralization. The presence of three biominerals in the trichomes of the basally branching Eucnide urens indicates either an early evolution and subsequent loss or several independent origins of multiple biomineralization. Differential biomineralization of the parts of individual, unicellular trichomes clearly indicates an extraordinary degree of physiological control over this process.

Background and Aims The detailed relationships in Geraniales in their current circumscription have only recently been clarified. The disparate floral morphologies and especially the nectaries of the corresponding group have consequently not previously been studied in a phylogenetic context. Methods The present study investigates floral and especially nectary morphology and structure for representatives of 12 of the 13 currently accepted genera in the five families of the Geraniales. Flowers were studied using light microscopy and scanning electron microscopy. Key Results The data demonstrate the derivation of even the most disparate floral morphologies from a basic pentamerous and pentacyclic organization, with an obdiplostemonous androecium and receptacular nectaries associated with the antesepalous stamens. Divergent morphologies are explained by modifications of merosity (tetramerous flowers), symmetry (several transitions to zygomorphic flowers) and elaboration of the nectaries into variously shaped outgrowths and appendages, especially in Francoaceae. The divergent development of nectar glands ultimately leads to either a reduction in their number (to one in some Geraniaceae and Melianthaceae) or their total loss (some Vivianiaceae). Conclusions Floral morphology of the Geraniales shows a high degree of similarity, despite the variation in overall floral appearance and nectary morphology. A hypothesis on the transformation of the nectaries within the Geraniales is presented.

Biomineralization provides living organisms with various materials for the formation of resilient structures. Calcium phosphate is the main component of teeth and bones in vertebrates, whereas especially silica serves for the protection against herbivores on many plant surfaces. Functional calcium phosphate structures are well-known from the animal kingdom, but had not so far been reported from higher plants. Here, we document the occurrence of calcium phosphate biomineralization in the South-American plant group Loasaceae (rock nettle family), which have stinging trichomes similar to those of the well-known stinging nettles (Urtica). Stinging hairs and the smaller, glochidiate trichomes contained nanocrystalline hydroxylated apatite, especially in their distal portions, replacing the silica found in analogous structures of other flowering plants. This could be demonstrated by chemical, spectroscopic and diffraction analyses. Some species of Loasaceae contained both calcium phosphate and silica in addition to calcium carbonate. The intriguing discovery of structural hydroxylated apatite in plants invites further studies, e.g., on its systematic distribution across the family, the genetic and cellular control of plant biomineralization, the properties and ultrastructure of calcium phosphate. It may prove the starting point for the development of biomimetic calcium phosphate composites based on a cellulose matrix.

53 p., gráf., The use of DNA sequence data in plant systematics has brought us closer than ever to formulating well-founded hypotheses about phylogenetic relationships, and phylogenetic research keeps on revealing that plant genera as traditionally circumscribed often are not monophyletic. Here, we assess the monophyly of all genera of vascular plants found in Germany. Using a survey of the phylogenetic literature, we discuss which classifications would be consistent with the phylogenetic relationships found and could be followed, provided monophyly is accepted as the primary criterion for circumscribing taxa. We indicate whether and which names are available when changes in generic assignment are made (but do not present a comprehensive review of the nomenclatural aspects of such names). Among the 840 genera examined, we identified c. 140 where data quality is sufficiently high to conclude that they are not monophyletic, and an additional c. 20 where monophyly is questionable but where data quality is not yet sufficient to reach convincing conclusions. While it is still fiercely debated how a phylogenetic tree should be translated into a classification, our results could serve as a guide to the likely consequences of systematic research for the taxonomy of the German flora and the floras of neighbouring countries.

The Boraginales are now universally accepted as monophyletic and firmly placed in Lamiidae. However, a consensus about familial classification has remained elusive, with some advocating recognition of a single, widely variable family, and others proposing recognition of several distinct families. A consensus classification is proposed here, based on recent molecular phylogenetic studies, morphological characters, and taking nomenclatural stability into consideration. We suggest the recognition of eleven, morphologically well-defined and clearly monophyletic families, namely the Boraginaceae s.str., Codonaceae, Coldeniaceae fam. nov., Cordiaceae, Ehretiaceae, Heliotropiaceae, Hoplestigmataceae, Hydrophyllaceae, Lennoaceae, Namaceae, and Wellstediaceae. Descriptions, synonomy, a taxonomic key, and a list of genera for these eleven families are provided, including the new family Coldeniaceae (monogeneric) and Namaceae (segregated from Hydrophyllaceae and comprising Nama, Eriodictyon, Turricula, and Wigandia), the latter necessitating a revised circumscription of a more morphologically coherent Hydrophyllaceae.

SYNTHESYS project by the European Community Research Infrastructure Action under the FP6 "Structuring the European Research Area" GB-TAF-4514 FP7 "Capacities" Programmes ES-TAF-136 AT-TAF-2001 FR-TAF-1977 SE-TAF-2567

Krameria lappacea (Domb.) Burdet & B.B.Simpson is a shrub of the semi-desert in the Andes of Peru (Fig. 5.1 A–C), southern Ecuador, and northern Argentina, Chile and Bolivia, occurring at elevations from sea level to 3600 m a.s.l. (Simpson, 1989a; Simpson et al., 2004). In its native habitat K. lappacea is by far the largest root parasite, with individual shrubs reaching a diameter of 1.5 m and a height of up to 2 m. Inmost semi-desert areas of Peru it is also clearly the most common root parasite.

The origin of Geraniales (approximately 900 species in three families: Geraniaceae, Melianthaceae, and Vivianiaceae) is traced back to the Cretaceous of Gondwana, yet their geotemporal history is largely unknown because of a limited fossil record and incomplete phylogenies. In the present study, we provide the first fossil record of Vivianiaceae and a highly resolved molecular phylogeny for all extant Geraniales genera. Our results support the hypothesis that five (instead of three) families should be recognized in the order Geraniales: Francoaceae A. Juss. (Francoa, Greyia, Tetilla), Geraniaceae Juss. (Erodium, Geranium, Monsonia, Pelargonium), Hypseocharitaceae Wedd. (monogeneric), Melianthaceae Horan. (Bersama, Melianthus), and Vivianiaceae Klotzsch (Balbisia, Rhynchotheca, Viviania). The four major lineages (i.e. Geraniaceae, Francoaceae + Melianthaceae, Hypseocharitaceae, Vivianiaceae) all originated within a narrow time frame during the Eocene (36.9–49.9 Mya) based on the five fossil calibration points. The divergence of most of the extant genera occurred much later, from the Miocene onwards. The South American–South African disjunction in Francoaceae apparently goes back to long distance dispersal with an estimated divergence time of the lineages in the Middle Miocene [11.2 (5.9–17.7) Mya]. Diversification in Melianthus appears to be much more recent than previously assumed [starting approximately 3.4 (1.9–5.2) Mya rather than approximately 8–20 Mya]. However, divergence of the Andean Hypseocharis lineage [36.9 (31.9–42.8) Mya] significantly predates the main Andean uplift: Current distributions likely go back to northward migrations and subsequent extinctions in Patagonia. Similarly, Rhynchotheca, Balbisia, and Viviania have a current southern distribution limit > 10°N of the fossil finds, indicating a massive northward displacement. The present evidence suggests that niche conservatism likely played a major role in the historical biogeography of Geraniales. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ••, ••–••.

2 PREMISE OF THE STUDY: The Paleogene deposits of the Hamada of Meridja, southwestern Algeria, are currently dated as lower-to-middle Eocene in age based on fossil gastropods and charophytes. Here we report the presence of fruits that can be assigned to the Boraginaceae s.str., apparently representing the fifossil record for this family in Africa, shedding new light on the historical biogeography of this group. METHODS: Microscopic studies of the fossil nutlets were carried out and compared to extant Boraginaceae nutlets, and to types reported in the literature for this family. KEY RESULTS: The fossils are strikingly similar in general size and morphology, particularly in the fi ner details of the attachment scar and ornamentation, to nutlets of extant representatives of the Boraginaceae tribe Echiochileae, and especially the genus Ogastemma . We believe that these nutlets represent an extinct member of this lineage. CONCLUSIONS: The Ogastemma -like fossils indicate that the Echiochileae, which are most diverse in northern Africa and southwestern Asia, have a long history in this region, dating back to the Eocene. This tribe corresponds to the basal-most clade in Boraginaceae s.str., and the fossils described here agree well with an assumed African origin of the family and the Boraginales I, providing an important additional calibration point for dating the phylogenies of this clade.

The monophyletic Nasa ranunculifolia group (Nasa ser. Grandiflorae pro parte) is revised on the basis of extensive field studies and a revision of copious herbarium material. All species of this group are from the high Andes of Peru and Ecuador (mostly 3000–4300 m). The overall degree of morphological divergence is much lower than in other groups of Nasa and many of the various local ‘races’ are best recognized as ecogeographically isolated subspecies. Thus, six species and 13 subspecies are recognized on the basis of differences in leaf and floral morphology, growth habit, life history and distribution. Nasa ranunculifolia, N. macrantha, N. cymbopetala and N. macrorrhiza are united as subspecies under N. ranunculifolia. An additional five subspecies of N. ranunculifolia are newly described (ssp. pamparomasii, ssp. guzmangoensis, ssp. bolivarensis, ssp. patazensis and ssp. huanucoensis). Nasa rugosa is subdivided into four subspecies, three of them new (ssp. llaqtacochaensis, ssp. gracilipes and ssp. pygmaea). Nasa tulipadiaboli sp. nov. and N. basilica sp. nov. are described as new species. Eight taxa (seven subspecies and one species) are endemic to the Amotape–Huancabamba Zone, and seven of them (six subspecies and one species) are new to science. Only a single species ranges into the Amotape–Huancabamba Zone from the south, underscoring the high levels of endemism (and taxonomic novelty) in this area. Illustrations, distribution maps, a key and diagnoses are given for all taxa recognized. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167, 47–93.

The only Namibian species of Spiloxene is described and illustrated by drawings and photos as Spiloxene etesionamibensis U.Mull.-Doblies, Mark.Ackermann, Weigend &. D.Mull.-Doblies. It was hitherto misinterpreted as Spiloxene scullyi (Baker) Garside in Solch & Roessler (1969) in Merxmuller's Prodromus einer Flora von Sudwestafrika, but its closest ally is S. flaccida (Nel) Garside, found in the Western and the Eastern Cape. Spiloxene etesionamibensis, which was found independently by Giess & Merxmuller and by Muller-Doblies, is only known from two or three independent collections from a single locality. S. scullyi is no longer known from Namibia. In addition to the eight Spiloxene species already known from the Northern Cape Spiloxene namaquana U.Mull.-Doblies, Mark.Ackermann, Weigend & D.Mull.-Doblies is described and illustrated by drawings and photos. Its closest ally is Spiloxene trifurcillata (Nel) Fourc. from the Eastern Cape. For Spiloxene namaquana the morphological analysis of the branching pattern of an annual shoot taken in 1988 in the field is evaluated by a longitudinal and a horizontal diagram with explanatory remarks. The unusual innovation system of extreme anisoiteration is discussed. On Map 1 the quarter degree squares of the type localities (± only known localities) of both species is given. (© 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)