Michael H J Barfuss | University of Vienna (original) (raw)

Papers by Michael H J Barfuss

Research paper thumbnail of Correction: Phylogeny, character evolution and spatiotemporal diversification of the species-rich and world-wide distributed tribe Rubieae (Rubiaceae)

PLOS ONE, Jan 25, 2019

The funding section is incorrect. The correct funding information is as follows: Open access fund... more The funding section is incorrect. The correct funding information is as follows: Open access funding provided by University of Vienna.

Research paper thumbnail of Phylogenetic relationships and evolutionary trends in <i>Tillandsia</i> subgenus <i>Diaphoranthema</i> and xerophytic species of subgenus <i>Phytarrhiza</i> (Bromeliaceae: Tillandsioideae)

Botanical Journal of the Linnean Society, Jul 23, 2022

Tillandsia subgenera Diaphoranthema and Phytarrhiza s.s. are closely related epiphytic and epilit... more Tillandsia subgenera Diaphoranthema and Phytarrhiza s.s. are closely related epiphytic and epilithic plants adapted to xeric habitats in the Americas. Several studies suggested that subgenus Diaphoranthema and the xerophytic species of subgenus Phytarrhiza should not be segregated into different subgenera. In recent phylogenetic analyses of Tillandsioideae, subgenus Phytarrhiza s.l. is revealed as highly polyphyletic, showing a xerophytic lineage closely related to subgenus Diaphoranthema. Moreover, the evolution of neotenic traits, a selfing breeding system and polyembryony have yet to be fully investigated in a phylogenetic context. Infrageneric classifications and phylogenetic relationships were inferred using the plastid matK-trnK and nuclear ETS markers and morphology. Phylogenetic analyses of individual and combined molecular data sets were performed under parsimony and Bayesian inference. Our results confirm subgenus Diaphoranthema as monophyletic if three xeric species of subgenus Phytarrhiza are included. In subgenus Diaphoranthema, aggregates Loliacea and Rectangula were recovered as monophyletic, whereas Myosura was paraphyletic and Caliginosa, Capillaris and Recurvata were polyphyletic. Alternative groupings and morphological trends are discussed. Analysis of morphological characters associated with heterochrony indicates that neoteny may not be the only paedomorphic process in the evolution of this group. Furthermore, paedomorphosis would occur in some vegetative structures, inflorescences and flowers, whereas some fruit and seed traits could be modified by peramorphosis.

Research paper thumbnail of Comparative phylogeography, morphological boundaries and climate envelopes of two sympatric widespread Bromeliaceae from the southern Andes

Botanical Journal of the Linnean Society, Mar 24, 2020

We studied the epiphytic species Tillandsia capillaris and T. virescens that grow as neotenic for... more We studied the epiphytic species Tillandsia capillaris and T. virescens that grow as neotenic forms with autogamous and cleistogamous flowers. They have expanded in Andean mountain environments, where they grow sympatrically across most of their ranges in arid regions of Peru, central-western Bolivia and north-central Argentina and Chile. We studied the relationships between the two species and populations within each of them, the morphological boundaries and the climatic characterization of distinct genetic entities. We analysed 125 and 102 individuals from 25 and 27 populations of T. virescens and T. capillaris, respectively, by means of haplotype networks, phylogenetic and genetic structure analyses, using the plastid gene ycf1 (c. 4500 bp) to define genetic groups. Morphological studies by discriminant analyses and correlation with climatic variables extracted from WorldClim were used to test differentiations among the genetic groups within each species. We confirm that T. capillaris is a distinct entity and that T. virescens consists of three groups that diverged by vicariance from widespread ancestors. The high genetic diversity found in both species is consistent with the complex geography of the Andes and the Pleistocene glaciation cycles that have driven ecological speciation in both species reflected in the timing of divergence of the clusters.

Research paper thumbnail of Disentangling Relationships among the Alpine Species of Luzula Sect. Luzula (Juncaceae) in the Eastern Alps

Plants, Feb 20, 2023

This article is an open access article distributed under the terms and conditions of the Creative... more This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY

Research paper thumbnail of Molecular phylogeny of<i>Gymnocalycium</i>(Cactaceae): Assessment of alternative infrageneric systems, a new subgenus, and trends in the evolution of the genus

American Journal of Botany, Nov 1, 2011

With ca. 124 genera and 1500-1800 species almost exclusively native to the Americas, Cactaceae is... more With ca. 124 genera and 1500-1800 species almost exclusively native to the Americas, Cactaceae is one of the most remarkable plant families in the world, showing extraordinary adaptations to hot, arid environments that include morphological (e.g., water-storage tissues, thick wax-covered epidermis, expanded root systems) as well as physiological (CAM photosynthesis) features (Barcikowski and Nobel, 1984 ; Nobel, 2002). No less notable than their biological characteristics is their beauty, making cacti popular plants among collectors. In the last years, molecular-based studies have provided evidence supporting classifi cations of cacti at the family, subfamily, tribe, and subtribal level (

Research paper thumbnail of Setting the evolutionary timeline: Tillandsia landbeckii in the Chilean Atacama Desert

Plant Systematics and Evolution, May 20, 2021

The Chilean Atacama Desert is among the oldest deserts of the world. Here, Tillandsia landbeckii ... more The Chilean Atacama Desert is among the oldest deserts of the world. Here, Tillandsia landbeckii is forming a unique vegetation type known as Tillandsia lomas. This vegetation consists in its typical configuration of one single vascular plant species only and forms regular linear structures in a sloped landscape and is largely depending on fog occurrence as dominant source of water supply. Without developing a typical root system, there are only few other terrestrial Tillandsia species growing on bare sand in Chile and Peru such as T. marconae, T. virescens, T. purpureaor T. latifolia. Although phylogenetic evidence is limited, convergent evolution of this unique growth behavior is evident. The predominantly arid and hyper-arid climate exists since the Early Miocene, which raises the question about timing of T. landbeckii evolutionary history. Phylogenomic analyses using whole plastome sequence data highlight the onset of diversification in T. landbeckii approximately 500,000 years ago. We also demonstrate subsequent secondary genetic contact with T. purpurea during the Late Pleistocene using DNA sequence data and genome size estimates, which resulted into the formation of T. marconae.

Research paper thumbnail of Pleistocene survival in three Mediterranean refugia: origin and diversification of the Italian endemic <i>Euphorbia gasparrinii</i> from the <i>E. verrucosa</i> alliance (Euphorbiaceae)

Botanical Journal of the Linnean Society, Feb 21, 2019

Applying molecular (AFLP and sequences of nuclear ribosomal ITS), karyological (relative genome s... more Applying molecular (AFLP and sequences of nuclear ribosomal ITS), karyological (relative genome size estimations and chromosome counts) and morphometric methods we explored the origin of Italian endemic Euphorbia gasparrinii. AFLP data inferred three phylogeographic groups corresponding to Balkan-Central European-northern Italian E. verrucosa, Italian E. gasparrinii and Iberian E. flavicoma. Genetic differentiation among the three species is weak, suggesting their relatively recent divergence. Karyological analyses revealed that populations of E. gasparrinii have 2n = 16 chromosomes, whereas the other two taxa have 2n = 14. This, with weak genetic differentiation and allopatric distribution, supports its recognition as an independent species, despite its weak morphological differentiation. Our study also revealed multiple autopolyploidization events within all three taxa. Contrary to the diploid Apennine populations, the single currently known Sicilian population of E. gasparrinii is tetraploid. It is critically endangered, as the Sicilian distribution has decreased significantly during the past century. Our study underlines the importance of southern European peninsulas as refugial areas during the Pleistocene and provides additional evidence that Mediterranean high mountain plants are suffering severe range contractions due to climate warming.

Research paper thumbnail of Systematics and evolution of the Old World Ebenaceae, a review with emphasis on the large genus<i>Diospyros</i>and its radiation in New Caledonia

Botanical Journal of the Linnean Society, Jan 25, 2019

In this review article, we provide an overview of the status of research on Old World Ebenaceae w... more In this review article, we provide an overview of the status of research on Old World Ebenaceae with an emphasis on the large genus Diospyros. The well-supported phylogenetic tree obtained from nucleotide sequences of multiple regions of plastid genome gave clear insights into the subfamilial classification of Ebenaceae. It supported inclusion of previously recognized genera such as Cargillia, Gunisanthus, Maba, Macreightia and Tetraclis in Diospyros. Endemic Diospyros spp. of New Caledonia have multiple origins. One of these clades has c. 21 species that are morphologically distinct and occupy different ecological niches, but they exhibit low genetic variation, leading to a lack of phylogenetic resolution. Analyses of whole plastid genome sequences did not greatly increase resolution or support for results of our previous plastid analyses. Geographical clustering of the individuals against a background of lower sequence divergence of the whole plastid genome could be due to transfer of plastid genomes during hybridization and introgression following secondary contact. However, > 8400 filtered single nucleotide polymorphisms (SNPs) from restriction site associated DNA sequencing (RADseq) confirmed species circumscriptions for this clade and produced well-supported phylogenetic relationships, pointing to an early regional clustering among populations and species. This supported allopatric speciation with respect to macrohabitat (i.e. climatic conditions) having had a role in the initial differentiation in the group. A later, more rapid radiation involved divergence with respect to microhabitat (e.g. soil preference). Although chromosome counts indicate that Diospyros spp. are consistently diploids with 2n = 30, extensive variation in genome size has been observed, which is due to an increase of repeat elements, including LTR/gypsy. In Ebenaceae, pollen is heterogeneous, and palynological synapomorphies are traced at different taxonomic levels. Several new Diospyros spp. have recently been identified and documented from India, Thailand, China, Africa and New Caledonia. Taxonomic revisions have been completed for the Australian species, and synonyms are reported for some New Caledonian Diospyros spp.

Research paper thumbnail of Plastome phylogenomics reveals an early Pliocene North- and Central America colonization by long-distance dispersal from South America of a highly diverse bromeliad lineage

Frontiers in Plant Science

Understanding the spatial and temporal frameworks of species diversification is fundamental in ev... more Understanding the spatial and temporal frameworks of species diversification is fundamental in evolutionary biology. Assessing the geographic origin and dispersal history of highly diverse lineages of rapid diversification can be hindered by the lack of appropriately sampled, resolved, and strongly supported phylogenetic contexts. The use of currently available cost-efficient sequencing strategies allows for the generation of a substantial amount of sequence data for dense taxonomic samplings, which together with well-curated geographic information and biogeographic models allow us to formally test the mode and tempo of dispersal events occurring in quick succession. Here, we assess the spatial and temporal frameworks for the origin and dispersal history of the expanded clade K, a highly diverse Tillandsia subgenus Tillandsia (Bromeliaceae, Poales) lineage hypothesized to have undergone a rapid radiation across the Neotropics. We assembled full plastomes from Hyb-Seq data for a dens...

Research paper thumbnail of Disentangling Relationships among the Alpine Species of Luzula Sect. Luzula (Juncaceae) in the Eastern Alps

Plants

Polyploidisation, agmatoploidy and symploidy have driven the diversification of Luzula sect. Luzu... more Polyploidisation, agmatoploidy and symploidy have driven the diversification of Luzula sect. Luzula. Several morphologically very similar species with different karyotypes have evolved, but their evolutionary origins and relationships are unknown. In this study, we used a combination of relative genome size and karyotype estimations as well amplified fragment length polymorphism (AFLP) fingerprinting to investigate the relationships among predominately (sub)alpine Luzula alpina, L. exspectata, L multiflora and L. sudetica in the Eastern Alps, including also some samples of L. campestris and L. taurica as outgroup. Our study revealed common co-occurrence of two or three different ploidies (di-, tetra- and hexaploids) at the same localities, and thus also common co-occurrence of different species, of which L. sudetica was morphologically, ecologically and genetically most divergent. Whereas agmatoploid L. exspectata likely originated only once from the Balkan L. taurica, and hexaploid...

Research paper thumbnail of Diploid Chastity vs. Polyploid Promiscuity – Extensive Gene Flow Among Polyploid Cytotypes Blurs Genetic, Morphological and Taxonomic Boundaries Among Dinaric Taxa of Knautia (Caprifoliaceae)

Research paper thumbnail of Phylogeographic patterns ofDeschampsia cespitosa(Poaceae) in Europe inferred from genomic data

Botanical Journal of the Linnean Society

The highly variable tufted hairgrass Deschampsia cespitosa is a tussock-forming plant especially ... more The highly variable tufted hairgrass Deschampsia cespitosa is a tussock-forming plant especially of cool and humid environments. Although common and widespread, its phylogeographic structure and the significance of polyploidy for its evolution are poorly understood. Here we used a phylogenomic approach to study the genetic structure of this species in Europe and how the polyploid cytotypes/subspecies are related to the diploids. Using genomic data (RADseq and whole plastid sequencing) we found a highly divergent Iberian group, including the Spanish Deschampsia cespitosa subsp. cespitosa (diploid and tetraploid) and the Macaronesian island endemic diploid Deschampsia argentea. Moreover, we found substantial divergence of pseudoviviparous tetraploids (Deschampsia cespitosa subspp. neoalpina and rhenana) from seminiferous tetraploids (except Deschampsia cespitosa subsp. littoralis) and all diploids of the remaining European samples. The divergent pseudoviviparous tetraploids (D. cespit...

Research paper thumbnail of Molecular phylogeny and chromosomal evolution of endemic species of Sri Lankan Anacardiaceae

Journal of the National Science Foundation of Sri Lanka, 2020

Family Anacardiaceae comprises 70 genera and approximately 985 species distributed worldwide. Sri... more Family Anacardiaceae comprises 70 genera and approximately 985 species distributed worldwide. Sri Lanka harbours 19 species in seven genera, among these 15 are endemics. This study focuses on regionally restricted endemics and native Anacardiaceae species, which have not been investigated before at molecular and cytological level. Nuclear rDNA ITS and plastid matK regions were sequenced for ten species, having nine endemics and one native, and incorporated into the existing sequence data for phylogenetic analyses. The topologies resulting from maximum parsimony, maximum likelihood and Bayesian inference are congruent. Family Anacardiaceae forms a monophyletic group having monophyletic subfamily Anacardioideae and paraphyletic subfamily Spondioideae. Tribes Anacardieae, Semecarpeae and Rhoeae form subclades within the major clade of Anacardioideae. All the endemic species occupy correct position in the molecular phylogeny as per the existing classifi cation except for Campnosperma zeylanica, which shows a close relationship to members of the subfamily Spondioideae. The position of non-native species Trichoscypha acuminata also remains unclear. Chromosome counts and karyograms were constructed for fi ve endemic species. The chromosome numbers incorporated in the tree range from 2n = 28-58. Species of tribe Rhoeae have the lowest chromosome number (2n = 24, 28, 30) while species of tribe Semecarpeae have the higher numbers (2n = 50, 52, 58). Chromosome numbers mapped on the phylogeny shows that dysploidy had played a role in the evolution of the species of the family Anacardiaceae in Sri Lanka.

Research paper thumbnail of The new locally endemic genus Yazdana (Caryophyllaceae) and patterns of endemism highlight the high conservation priority of the poorly studied Shirkuh Mountains (central Iran)

Journal of Systematics and Evolution, 2020

Although mountain ranges are often recognized as global biodiversity hotspots with a high level o... more Although mountain ranges are often recognized as global biodiversity hotspots with a high level of endemism, diversity and biogeographic connections of isolated and weakly explored mountains remain poorly understood. This is also the case for Shirkuh Mts. in central Iran. Here, Yazdana shirkuhensis gen. & spec. nov. (Caryophylleae, Caryophyllaceae) is described and illustrated from the high alpine zone of this mountain. Molecular phylogenetic analyses of nuclear and plastid DNA sequence data show that Y. shirkuhensis is related to Cyathophylla and Heterochroa (tribe Caryophylleae). The newly described genus and species accentuate Shirkuh Mts. as a center of endemism, which harbors a high number of narrowly distributed species, mostly in high elevations reaching alpine habitats. As this area is currently not protected, a conservation priority is highlighted for high elevations of Shirkuh Mts.

Research paper thumbnail of Phylogenetic analyses of plastid DNA suggest a different interpretation of morphological evolution than those used as the basis for previous classifications of Dipterocarpaceae (Malvales)

Botanical Journal of the Linnean Society, 2017

Phylogenetic and molecular clock analyses were performed including all genera except one (Pseudom... more Phylogenetic and molecular clock analyses were performed including all genera except one (Pseudomonotes) for the three subfamilies of Dipterocarpaceae. We also included representatives of Sarcolaenaceae and Cistaceae with Bixaceae as the ultimate outgroup. Three plastid regions (six markers), partial rbcL, trnK-matK-trnK (partial trnK intron including complete matK) and trnT-trnL-trnF (partial trnT, complete trnT-trnL intergenic spacer, complete trnL, complete trnL-trnF intergenic spacer and partial trnF), were analysed. We also investigated additional accessions for genome size and chromosome numbers. Our phylogenetic results differ in three important respects from previous interpretations of morphological characters, as reflected in recent classifications. First, our analyses strongly support assignment of Pakaraimaea (subfamily Pakaraimaeoideae) to Cistaceae. Second, the morphological concepts of Dipterocarpeae and Shoreeae in subfamily Dipterocarpoideae are not supported because Dipterocarpus is sister to Dryobalanops plus tribe Shoreeae. Our analysis revealed four clades: (1) Dipterocarpus; (2) Dryobalanops, for which tribal assignment has been contentious; (3) genera of Shoreeae; and (4) the remaining genera of Dipterocarpeae. Third, Shorea is not monophyletic. Monotoideae are weakly supported as sister to Dipterocarpoideae; Sarcolaenaceae (endemic to Madagascar) are sister to this pair. Divergence in extant Dipterocarpoideae occurred c. 55 Mya. Genome sizes for all accessions examined are small (0.3264-0.6724 pg), and the additional chromosome numbers we collected fit into the patterns previously observed for Dipterocarpaceae.

Research paper thumbnail of A preliminary phylogeny of the ‘didymocarpoid Gesneriaceae’ based on three molecular data sets: Incongruence with available tribal classifications

American Journal of Botany, 2009

The ‘didymocarpoid Gesneriaceae’ (traditional subfam. Cyrtandroideae excluding Epithemateae) are ... more The ‘didymocarpoid Gesneriaceae’ (traditional subfam. Cyrtandroideae excluding Epithemateae) are the largest group of Old World Gesneriaceae, comprising 85 genera and 1800 species. We attempt to resolve their hitherto poorly understood generic relationships using three molecular markers on 145 species, of which 128 belong to didymocarpoid Gesneriaceae. Our analyses demonstrate that consistent topological relationships can be retrieved from data sets with missing data using subsamples and different combinations of gene sequences. We show that all available classifications in Old World Gesneriaceae are artificial and do not reflect natural relationships. At the base of the didymocarpoids are grades of clades comprising isolated genera and small groups from Asia and Europe. These are followed by a clade comprising the African and Madagascan genera. The remaining clades represent the advanced Asiatic and Malesian genera. They include a major group with mostly twisted capsules. The much ...

Research paper thumbnail of CAM evolution is associated with gene family expansion in an explosive bromeliad radiation

1.The subgenusTillandsia(Bromeliaceae) belongs to one of the fastest radiating clades in the plan... more 1.The subgenusTillandsia(Bromeliaceae) belongs to one of the fastest radiating clades in the plant kingdom and is characterised by the repeated evolution of Crassulacean Acid Metabolism (CAM). Despite its complex genetic basis, this water-conserving trait has evolved independently across many plant families and is regarded as a key innovation trait and driver of ecological diversification in Bromeliaceae. By producing high quality genome assemblies of aTillandsiaspecies pair displaying divergent photosynthetic phenotypes, and combining genome-wide investigations of synteny, TE dynamics, sequence evolution, gene family evolution and temporal differential expression, we were able to pinpoint the genomic drivers of CAM evolution inTillandsia. Several large-scale rearrangements associated with karyotype changes between the two genomes and a highly dynamic TE landscape shaped the genomes ofTillandsia. However, our analyses show that rewiring of photosynthetic metabolism is mainly obtaine...

Research paper thumbnail of Molecular Phylogenetics and Trait Evolution in <i>Stigmatodon</i> (Bromeliaceae, Tillandsioideae), an Endemic Genus to Brazilian Rocky Outcrops

Systematic Botany, Jun 15, 2022

Research paper thumbnail of Phylogeny, character evolution and spatiotemporal diversification of the species-rich and world-wide distributed tribe Rubieae (Rubiaceae)

PLOS ONE, 2018

The Rubiaceae tribe Rubieae has a worldwide distribution with up to 1,000 species. These collecti... more The Rubiaceae tribe Rubieae has a worldwide distribution with up to 1,000 species. These collectively exhibit an enormous ecological and morphological diversity, making Rubieae an excellent group for macro-and microevolutionary studies. Previous molecular phylogenetic analyses used only a limited sampling within the tribe or missed lineages crucial for understanding character evolution in this group. Here, we analyze sequences from two plastid spacer regions as well as morphological and biogeographic data from an extensive and evenly distributed sampling to establish a sound phylogenetic framework. This framework serves as a basis for our investigation of the evolution of important morphological characters and the biogeographic history of the Rubieae. The tribe includes three major clades, the Kelloggiinae Clade (Kelloggia), the Rubiinae Clade (Didymaea, Rubia) and the most species-rich Galiinae Clade (Asperula,

Research paper thumbnail of Adaptive radiation, correlated and contingent evolution, and net species diversification in Bromeliaceae

Molecular Phylogenetics and Evolution, 2014

We present an integrative model predicting associations among epiphytism, the tank habit, entangl... more We present an integrative model predicting associations among epiphytism, the tank habit, entangling seeds, C 3 vs. CAM photosynthesis, avian pollinators, life in fertile, moist montane habitats, and net rates of species diversification in the monocot family Bromeliaceae. We test these predictions by relating evolutionary shifts in form, physiology, and ecology to time and ancestral distributions, quantifying patterns of correlated and contingent evolution among pairs of traits and analyzing the apparent impact of individual traits on rates of net species diversification and geographic expansion beyond the ancestral Guayana Shield. All predicted patterns of correlated evolution were significant, and the temporal and spatial associations of phenotypic shifts with orogenies generally accorded with predictions. Net rates of species diversification were most closely coupled to life in fertile, moist, geographically extensive cordilleras, with additional significant ties to epiphytism, avian pollination, and the tank habit. The highest rates of net diversification were seen in the bromelioid tank-epiphytic clade (D crown = 1.05 My À1), associated primarily with the Serra do Mar and nearby ranges of coastal Brazil, and in the core tillandsioids (D crown = 0.67 My À1), associated primarily with the Andes and Central America. Six large-scale adaptive radiations and accompanying pulses of speciation account for 86% of total species richness in the family. This study is among the first to test a priori hypotheses about the relationships among phylogeny, phenotypic evolution, geographic spread, and net species diversification, and to argue for causality to flow from functional diversity to spatial expansion to species diversity.

Research paper thumbnail of Correction: Phylogeny, character evolution and spatiotemporal diversification of the species-rich and world-wide distributed tribe Rubieae (Rubiaceae)

PLOS ONE, Jan 25, 2019

The funding section is incorrect. The correct funding information is as follows: Open access fund... more The funding section is incorrect. The correct funding information is as follows: Open access funding provided by University of Vienna.

Research paper thumbnail of Phylogenetic relationships and evolutionary trends in <i>Tillandsia</i> subgenus <i>Diaphoranthema</i> and xerophytic species of subgenus <i>Phytarrhiza</i> (Bromeliaceae: Tillandsioideae)

Botanical Journal of the Linnean Society, Jul 23, 2022

Tillandsia subgenera Diaphoranthema and Phytarrhiza s.s. are closely related epiphytic and epilit... more Tillandsia subgenera Diaphoranthema and Phytarrhiza s.s. are closely related epiphytic and epilithic plants adapted to xeric habitats in the Americas. Several studies suggested that subgenus Diaphoranthema and the xerophytic species of subgenus Phytarrhiza should not be segregated into different subgenera. In recent phylogenetic analyses of Tillandsioideae, subgenus Phytarrhiza s.l. is revealed as highly polyphyletic, showing a xerophytic lineage closely related to subgenus Diaphoranthema. Moreover, the evolution of neotenic traits, a selfing breeding system and polyembryony have yet to be fully investigated in a phylogenetic context. Infrageneric classifications and phylogenetic relationships were inferred using the plastid matK-trnK and nuclear ETS markers and morphology. Phylogenetic analyses of individual and combined molecular data sets were performed under parsimony and Bayesian inference. Our results confirm subgenus Diaphoranthema as monophyletic if three xeric species of subgenus Phytarrhiza are included. In subgenus Diaphoranthema, aggregates Loliacea and Rectangula were recovered as monophyletic, whereas Myosura was paraphyletic and Caliginosa, Capillaris and Recurvata were polyphyletic. Alternative groupings and morphological trends are discussed. Analysis of morphological characters associated with heterochrony indicates that neoteny may not be the only paedomorphic process in the evolution of this group. Furthermore, paedomorphosis would occur in some vegetative structures, inflorescences and flowers, whereas some fruit and seed traits could be modified by peramorphosis.

Research paper thumbnail of Comparative phylogeography, morphological boundaries and climate envelopes of two sympatric widespread Bromeliaceae from the southern Andes

Botanical Journal of the Linnean Society, Mar 24, 2020

We studied the epiphytic species Tillandsia capillaris and T. virescens that grow as neotenic for... more We studied the epiphytic species Tillandsia capillaris and T. virescens that grow as neotenic forms with autogamous and cleistogamous flowers. They have expanded in Andean mountain environments, where they grow sympatrically across most of their ranges in arid regions of Peru, central-western Bolivia and north-central Argentina and Chile. We studied the relationships between the two species and populations within each of them, the morphological boundaries and the climatic characterization of distinct genetic entities. We analysed 125 and 102 individuals from 25 and 27 populations of T. virescens and T. capillaris, respectively, by means of haplotype networks, phylogenetic and genetic structure analyses, using the plastid gene ycf1 (c. 4500 bp) to define genetic groups. Morphological studies by discriminant analyses and correlation with climatic variables extracted from WorldClim were used to test differentiations among the genetic groups within each species. We confirm that T. capillaris is a distinct entity and that T. virescens consists of three groups that diverged by vicariance from widespread ancestors. The high genetic diversity found in both species is consistent with the complex geography of the Andes and the Pleistocene glaciation cycles that have driven ecological speciation in both species reflected in the timing of divergence of the clusters.

Research paper thumbnail of Disentangling Relationships among the Alpine Species of Luzula Sect. Luzula (Juncaceae) in the Eastern Alps

Plants, Feb 20, 2023

This article is an open access article distributed under the terms and conditions of the Creative... more This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY

Research paper thumbnail of Molecular phylogeny of<i>Gymnocalycium</i>(Cactaceae): Assessment of alternative infrageneric systems, a new subgenus, and trends in the evolution of the genus

American Journal of Botany, Nov 1, 2011

With ca. 124 genera and 1500-1800 species almost exclusively native to the Americas, Cactaceae is... more With ca. 124 genera and 1500-1800 species almost exclusively native to the Americas, Cactaceae is one of the most remarkable plant families in the world, showing extraordinary adaptations to hot, arid environments that include morphological (e.g., water-storage tissues, thick wax-covered epidermis, expanded root systems) as well as physiological (CAM photosynthesis) features (Barcikowski and Nobel, 1984 ; Nobel, 2002). No less notable than their biological characteristics is their beauty, making cacti popular plants among collectors. In the last years, molecular-based studies have provided evidence supporting classifi cations of cacti at the family, subfamily, tribe, and subtribal level (

Research paper thumbnail of Setting the evolutionary timeline: Tillandsia landbeckii in the Chilean Atacama Desert

Plant Systematics and Evolution, May 20, 2021

The Chilean Atacama Desert is among the oldest deserts of the world. Here, Tillandsia landbeckii ... more The Chilean Atacama Desert is among the oldest deserts of the world. Here, Tillandsia landbeckii is forming a unique vegetation type known as Tillandsia lomas. This vegetation consists in its typical configuration of one single vascular plant species only and forms regular linear structures in a sloped landscape and is largely depending on fog occurrence as dominant source of water supply. Without developing a typical root system, there are only few other terrestrial Tillandsia species growing on bare sand in Chile and Peru such as T. marconae, T. virescens, T. purpureaor T. latifolia. Although phylogenetic evidence is limited, convergent evolution of this unique growth behavior is evident. The predominantly arid and hyper-arid climate exists since the Early Miocene, which raises the question about timing of T. landbeckii evolutionary history. Phylogenomic analyses using whole plastome sequence data highlight the onset of diversification in T. landbeckii approximately 500,000 years ago. We also demonstrate subsequent secondary genetic contact with T. purpurea during the Late Pleistocene using DNA sequence data and genome size estimates, which resulted into the formation of T. marconae.

Research paper thumbnail of Pleistocene survival in three Mediterranean refugia: origin and diversification of the Italian endemic <i>Euphorbia gasparrinii</i> from the <i>E. verrucosa</i> alliance (Euphorbiaceae)

Botanical Journal of the Linnean Society, Feb 21, 2019

Applying molecular (AFLP and sequences of nuclear ribosomal ITS), karyological (relative genome s... more Applying molecular (AFLP and sequences of nuclear ribosomal ITS), karyological (relative genome size estimations and chromosome counts) and morphometric methods we explored the origin of Italian endemic Euphorbia gasparrinii. AFLP data inferred three phylogeographic groups corresponding to Balkan-Central European-northern Italian E. verrucosa, Italian E. gasparrinii and Iberian E. flavicoma. Genetic differentiation among the three species is weak, suggesting their relatively recent divergence. Karyological analyses revealed that populations of E. gasparrinii have 2n = 16 chromosomes, whereas the other two taxa have 2n = 14. This, with weak genetic differentiation and allopatric distribution, supports its recognition as an independent species, despite its weak morphological differentiation. Our study also revealed multiple autopolyploidization events within all three taxa. Contrary to the diploid Apennine populations, the single currently known Sicilian population of E. gasparrinii is tetraploid. It is critically endangered, as the Sicilian distribution has decreased significantly during the past century. Our study underlines the importance of southern European peninsulas as refugial areas during the Pleistocene and provides additional evidence that Mediterranean high mountain plants are suffering severe range contractions due to climate warming.

Research paper thumbnail of Systematics and evolution of the Old World Ebenaceae, a review with emphasis on the large genus<i>Diospyros</i>and its radiation in New Caledonia

Botanical Journal of the Linnean Society, Jan 25, 2019

In this review article, we provide an overview of the status of research on Old World Ebenaceae w... more In this review article, we provide an overview of the status of research on Old World Ebenaceae with an emphasis on the large genus Diospyros. The well-supported phylogenetic tree obtained from nucleotide sequences of multiple regions of plastid genome gave clear insights into the subfamilial classification of Ebenaceae. It supported inclusion of previously recognized genera such as Cargillia, Gunisanthus, Maba, Macreightia and Tetraclis in Diospyros. Endemic Diospyros spp. of New Caledonia have multiple origins. One of these clades has c. 21 species that are morphologically distinct and occupy different ecological niches, but they exhibit low genetic variation, leading to a lack of phylogenetic resolution. Analyses of whole plastid genome sequences did not greatly increase resolution or support for results of our previous plastid analyses. Geographical clustering of the individuals against a background of lower sequence divergence of the whole plastid genome could be due to transfer of plastid genomes during hybridization and introgression following secondary contact. However, > 8400 filtered single nucleotide polymorphisms (SNPs) from restriction site associated DNA sequencing (RADseq) confirmed species circumscriptions for this clade and produced well-supported phylogenetic relationships, pointing to an early regional clustering among populations and species. This supported allopatric speciation with respect to macrohabitat (i.e. climatic conditions) having had a role in the initial differentiation in the group. A later, more rapid radiation involved divergence with respect to microhabitat (e.g. soil preference). Although chromosome counts indicate that Diospyros spp. are consistently diploids with 2n = 30, extensive variation in genome size has been observed, which is due to an increase of repeat elements, including LTR/gypsy. In Ebenaceae, pollen is heterogeneous, and palynological synapomorphies are traced at different taxonomic levels. Several new Diospyros spp. have recently been identified and documented from India, Thailand, China, Africa and New Caledonia. Taxonomic revisions have been completed for the Australian species, and synonyms are reported for some New Caledonian Diospyros spp.

Research paper thumbnail of Plastome phylogenomics reveals an early Pliocene North- and Central America colonization by long-distance dispersal from South America of a highly diverse bromeliad lineage

Frontiers in Plant Science

Understanding the spatial and temporal frameworks of species diversification is fundamental in ev... more Understanding the spatial and temporal frameworks of species diversification is fundamental in evolutionary biology. Assessing the geographic origin and dispersal history of highly diverse lineages of rapid diversification can be hindered by the lack of appropriately sampled, resolved, and strongly supported phylogenetic contexts. The use of currently available cost-efficient sequencing strategies allows for the generation of a substantial amount of sequence data for dense taxonomic samplings, which together with well-curated geographic information and biogeographic models allow us to formally test the mode and tempo of dispersal events occurring in quick succession. Here, we assess the spatial and temporal frameworks for the origin and dispersal history of the expanded clade K, a highly diverse Tillandsia subgenus Tillandsia (Bromeliaceae, Poales) lineage hypothesized to have undergone a rapid radiation across the Neotropics. We assembled full plastomes from Hyb-Seq data for a dens...

Research paper thumbnail of Disentangling Relationships among the Alpine Species of Luzula Sect. Luzula (Juncaceae) in the Eastern Alps

Plants

Polyploidisation, agmatoploidy and symploidy have driven the diversification of Luzula sect. Luzu... more Polyploidisation, agmatoploidy and symploidy have driven the diversification of Luzula sect. Luzula. Several morphologically very similar species with different karyotypes have evolved, but their evolutionary origins and relationships are unknown. In this study, we used a combination of relative genome size and karyotype estimations as well amplified fragment length polymorphism (AFLP) fingerprinting to investigate the relationships among predominately (sub)alpine Luzula alpina, L. exspectata, L multiflora and L. sudetica in the Eastern Alps, including also some samples of L. campestris and L. taurica as outgroup. Our study revealed common co-occurrence of two or three different ploidies (di-, tetra- and hexaploids) at the same localities, and thus also common co-occurrence of different species, of which L. sudetica was morphologically, ecologically and genetically most divergent. Whereas agmatoploid L. exspectata likely originated only once from the Balkan L. taurica, and hexaploid...

Research paper thumbnail of Diploid Chastity vs. Polyploid Promiscuity – Extensive Gene Flow Among Polyploid Cytotypes Blurs Genetic, Morphological and Taxonomic Boundaries Among Dinaric Taxa of Knautia (Caprifoliaceae)

Research paper thumbnail of Phylogeographic patterns ofDeschampsia cespitosa(Poaceae) in Europe inferred from genomic data

Botanical Journal of the Linnean Society

The highly variable tufted hairgrass Deschampsia cespitosa is a tussock-forming plant especially ... more The highly variable tufted hairgrass Deschampsia cespitosa is a tussock-forming plant especially of cool and humid environments. Although common and widespread, its phylogeographic structure and the significance of polyploidy for its evolution are poorly understood. Here we used a phylogenomic approach to study the genetic structure of this species in Europe and how the polyploid cytotypes/subspecies are related to the diploids. Using genomic data (RADseq and whole plastid sequencing) we found a highly divergent Iberian group, including the Spanish Deschampsia cespitosa subsp. cespitosa (diploid and tetraploid) and the Macaronesian island endemic diploid Deschampsia argentea. Moreover, we found substantial divergence of pseudoviviparous tetraploids (Deschampsia cespitosa subspp. neoalpina and rhenana) from seminiferous tetraploids (except Deschampsia cespitosa subsp. littoralis) and all diploids of the remaining European samples. The divergent pseudoviviparous tetraploids (D. cespit...

Research paper thumbnail of Molecular phylogeny and chromosomal evolution of endemic species of Sri Lankan Anacardiaceae

Journal of the National Science Foundation of Sri Lanka, 2020

Family Anacardiaceae comprises 70 genera and approximately 985 species distributed worldwide. Sri... more Family Anacardiaceae comprises 70 genera and approximately 985 species distributed worldwide. Sri Lanka harbours 19 species in seven genera, among these 15 are endemics. This study focuses on regionally restricted endemics and native Anacardiaceae species, which have not been investigated before at molecular and cytological level. Nuclear rDNA ITS and plastid matK regions were sequenced for ten species, having nine endemics and one native, and incorporated into the existing sequence data for phylogenetic analyses. The topologies resulting from maximum parsimony, maximum likelihood and Bayesian inference are congruent. Family Anacardiaceae forms a monophyletic group having monophyletic subfamily Anacardioideae and paraphyletic subfamily Spondioideae. Tribes Anacardieae, Semecarpeae and Rhoeae form subclades within the major clade of Anacardioideae. All the endemic species occupy correct position in the molecular phylogeny as per the existing classifi cation except for Campnosperma zeylanica, which shows a close relationship to members of the subfamily Spondioideae. The position of non-native species Trichoscypha acuminata also remains unclear. Chromosome counts and karyograms were constructed for fi ve endemic species. The chromosome numbers incorporated in the tree range from 2n = 28-58. Species of tribe Rhoeae have the lowest chromosome number (2n = 24, 28, 30) while species of tribe Semecarpeae have the higher numbers (2n = 50, 52, 58). Chromosome numbers mapped on the phylogeny shows that dysploidy had played a role in the evolution of the species of the family Anacardiaceae in Sri Lanka.

Research paper thumbnail of The new locally endemic genus Yazdana (Caryophyllaceae) and patterns of endemism highlight the high conservation priority of the poorly studied Shirkuh Mountains (central Iran)

Journal of Systematics and Evolution, 2020

Although mountain ranges are often recognized as global biodiversity hotspots with a high level o... more Although mountain ranges are often recognized as global biodiversity hotspots with a high level of endemism, diversity and biogeographic connections of isolated and weakly explored mountains remain poorly understood. This is also the case for Shirkuh Mts. in central Iran. Here, Yazdana shirkuhensis gen. & spec. nov. (Caryophylleae, Caryophyllaceae) is described and illustrated from the high alpine zone of this mountain. Molecular phylogenetic analyses of nuclear and plastid DNA sequence data show that Y. shirkuhensis is related to Cyathophylla and Heterochroa (tribe Caryophylleae). The newly described genus and species accentuate Shirkuh Mts. as a center of endemism, which harbors a high number of narrowly distributed species, mostly in high elevations reaching alpine habitats. As this area is currently not protected, a conservation priority is highlighted for high elevations of Shirkuh Mts.

Research paper thumbnail of Phylogenetic analyses of plastid DNA suggest a different interpretation of morphological evolution than those used as the basis for previous classifications of Dipterocarpaceae (Malvales)

Botanical Journal of the Linnean Society, 2017

Phylogenetic and molecular clock analyses were performed including all genera except one (Pseudom... more Phylogenetic and molecular clock analyses were performed including all genera except one (Pseudomonotes) for the three subfamilies of Dipterocarpaceae. We also included representatives of Sarcolaenaceae and Cistaceae with Bixaceae as the ultimate outgroup. Three plastid regions (six markers), partial rbcL, trnK-matK-trnK (partial trnK intron including complete matK) and trnT-trnL-trnF (partial trnT, complete trnT-trnL intergenic spacer, complete trnL, complete trnL-trnF intergenic spacer and partial trnF), were analysed. We also investigated additional accessions for genome size and chromosome numbers. Our phylogenetic results differ in three important respects from previous interpretations of morphological characters, as reflected in recent classifications. First, our analyses strongly support assignment of Pakaraimaea (subfamily Pakaraimaeoideae) to Cistaceae. Second, the morphological concepts of Dipterocarpeae and Shoreeae in subfamily Dipterocarpoideae are not supported because Dipterocarpus is sister to Dryobalanops plus tribe Shoreeae. Our analysis revealed four clades: (1) Dipterocarpus; (2) Dryobalanops, for which tribal assignment has been contentious; (3) genera of Shoreeae; and (4) the remaining genera of Dipterocarpeae. Third, Shorea is not monophyletic. Monotoideae are weakly supported as sister to Dipterocarpoideae; Sarcolaenaceae (endemic to Madagascar) are sister to this pair. Divergence in extant Dipterocarpoideae occurred c. 55 Mya. Genome sizes for all accessions examined are small (0.3264-0.6724 pg), and the additional chromosome numbers we collected fit into the patterns previously observed for Dipterocarpaceae.

Research paper thumbnail of A preliminary phylogeny of the ‘didymocarpoid Gesneriaceae’ based on three molecular data sets: Incongruence with available tribal classifications

American Journal of Botany, 2009

The ‘didymocarpoid Gesneriaceae’ (traditional subfam. Cyrtandroideae excluding Epithemateae) are ... more The ‘didymocarpoid Gesneriaceae’ (traditional subfam. Cyrtandroideae excluding Epithemateae) are the largest group of Old World Gesneriaceae, comprising 85 genera and 1800 species. We attempt to resolve their hitherto poorly understood generic relationships using three molecular markers on 145 species, of which 128 belong to didymocarpoid Gesneriaceae. Our analyses demonstrate that consistent topological relationships can be retrieved from data sets with missing data using subsamples and different combinations of gene sequences. We show that all available classifications in Old World Gesneriaceae are artificial and do not reflect natural relationships. At the base of the didymocarpoids are grades of clades comprising isolated genera and small groups from Asia and Europe. These are followed by a clade comprising the African and Madagascan genera. The remaining clades represent the advanced Asiatic and Malesian genera. They include a major group with mostly twisted capsules. The much ...

Research paper thumbnail of CAM evolution is associated with gene family expansion in an explosive bromeliad radiation

1.The subgenusTillandsia(Bromeliaceae) belongs to one of the fastest radiating clades in the plan... more 1.The subgenusTillandsia(Bromeliaceae) belongs to one of the fastest radiating clades in the plant kingdom and is characterised by the repeated evolution of Crassulacean Acid Metabolism (CAM). Despite its complex genetic basis, this water-conserving trait has evolved independently across many plant families and is regarded as a key innovation trait and driver of ecological diversification in Bromeliaceae. By producing high quality genome assemblies of aTillandsiaspecies pair displaying divergent photosynthetic phenotypes, and combining genome-wide investigations of synteny, TE dynamics, sequence evolution, gene family evolution and temporal differential expression, we were able to pinpoint the genomic drivers of CAM evolution inTillandsia. Several large-scale rearrangements associated with karyotype changes between the two genomes and a highly dynamic TE landscape shaped the genomes ofTillandsia. However, our analyses show that rewiring of photosynthetic metabolism is mainly obtaine...

Research paper thumbnail of Molecular Phylogenetics and Trait Evolution in <i>Stigmatodon</i> (Bromeliaceae, Tillandsioideae), an Endemic Genus to Brazilian Rocky Outcrops

Systematic Botany, Jun 15, 2022

Research paper thumbnail of Phylogeny, character evolution and spatiotemporal diversification of the species-rich and world-wide distributed tribe Rubieae (Rubiaceae)

PLOS ONE, 2018

The Rubiaceae tribe Rubieae has a worldwide distribution with up to 1,000 species. These collecti... more The Rubiaceae tribe Rubieae has a worldwide distribution with up to 1,000 species. These collectively exhibit an enormous ecological and morphological diversity, making Rubieae an excellent group for macro-and microevolutionary studies. Previous molecular phylogenetic analyses used only a limited sampling within the tribe or missed lineages crucial for understanding character evolution in this group. Here, we analyze sequences from two plastid spacer regions as well as morphological and biogeographic data from an extensive and evenly distributed sampling to establish a sound phylogenetic framework. This framework serves as a basis for our investigation of the evolution of important morphological characters and the biogeographic history of the Rubieae. The tribe includes three major clades, the Kelloggiinae Clade (Kelloggia), the Rubiinae Clade (Didymaea, Rubia) and the most species-rich Galiinae Clade (Asperula,

Research paper thumbnail of Adaptive radiation, correlated and contingent evolution, and net species diversification in Bromeliaceae

Molecular Phylogenetics and Evolution, 2014

We present an integrative model predicting associations among epiphytism, the tank habit, entangl... more We present an integrative model predicting associations among epiphytism, the tank habit, entangling seeds, C 3 vs. CAM photosynthesis, avian pollinators, life in fertile, moist montane habitats, and net rates of species diversification in the monocot family Bromeliaceae. We test these predictions by relating evolutionary shifts in form, physiology, and ecology to time and ancestral distributions, quantifying patterns of correlated and contingent evolution among pairs of traits and analyzing the apparent impact of individual traits on rates of net species diversification and geographic expansion beyond the ancestral Guayana Shield. All predicted patterns of correlated evolution were significant, and the temporal and spatial associations of phenotypic shifts with orogenies generally accorded with predictions. Net rates of species diversification were most closely coupled to life in fertile, moist, geographically extensive cordilleras, with additional significant ties to epiphytism, avian pollination, and the tank habit. The highest rates of net diversification were seen in the bromelioid tank-epiphytic clade (D crown = 1.05 My À1), associated primarily with the Serra do Mar and nearby ranges of coastal Brazil, and in the core tillandsioids (D crown = 0.67 My À1), associated primarily with the Andes and Central America. Six large-scale adaptive radiations and accompanying pulses of speciation account for 86% of total species richness in the family. This study is among the first to test a priori hypotheses about the relationships among phylogeny, phenotypic evolution, geographic spread, and net species diversification, and to argue for causality to flow from functional diversity to spatial expansion to species diversity.