<figure><img src="image_284.png" /></figure>

Santiago Martín-Bravo², Tomomi Masaki¹,⁵,
Ana Morales Alonso¹⁶, Mónica Míguez⁴,
Georg Miehe¹⁷, Anton A. Reznicek¹⁸,
Eric H. Roalson¹⁹, Julian R. Starr²⁰, Irina
Shekhovtsova²¹, Polina A. Volkova²²,
Marcia J. Waterway²³, Martin Röser¹,
Matthias H. Hoffmann¹

1 Department of Systematic Botany, Martin Luthersh University Halle-Wittenberg, Geobotany and Botanical Garden, Halle (Saale), Germany. 2 Department of Molecular Biology and Biochemical Engineering, Universidad Pablo de Olavide, Seville, Spain. 3 Research Institute of Forests and Rangelands, Agricultural Research, Education, and Extension Organization, Tehran, Iran. 4 Department of Plant Biology and Ecology, Universidad de Sevilla, Seville, Spain. 5 Natural History Museum and Biological Sciences Faculty, Universidad Nacional Mayor de San Marcos, Lima, Peru. 6 Department of Medicinal Plant Science, Jungwon University, Goisan, South Korea. 7 Department of Vascular Plants, Botanische Staatssammlung München & Systematic Botany and Mycology, University of München, München, Germany. 8 Parc naturel régional de la Vallée de Trient, Salvan, Switzerland. 9 The Morton Arboretum, Lisle, USA. 10 The Field Museum, Chicago, USA. 11 Department of Biosphere-Geosphere Science, Faculty of Biosphere-Geosphere Science, Okayama University of Science, Okayama, Japan. 12 Zhejiang Provincial Key Laboratory of Forest Automatic Plants-based Healthcare Functions and School of Forestry and Biotechnology, Zhejiang A&F University, Hangzhou, Zhejiang, China. 13 Department of Biology, Sungshin University, Seoul, South Korea. 14 Lioukuei Research Center, Taiwan Forestry Research Institute, Kaohsiung County, Taiwan. 15 Nature Fieldwork Center, Okayama University of Science, Okayama, Japan. 16 Biodiversity and Conservation Area, Department of Biology and Geology, Physics and Inorganic Chemistry, Universidad Rey Juan Carlos, Madrid, Spain. 17 Department of Geography, Vegetation Geography, Philipps-Universität Marburg, Marburg, Germany. 18 University of Michigan Herbarium, Ann Abor, USA. 19 School of Biological Sciences, Washington State University, Pullman, USA. 20 Department of Biology, University of Ottawa, Ottawa, Canada. 21 Central Siberian Botanical Garden SB RAS, Novosibirsk, Russia. 22 Papanin Institute for Biology of Inland Waters RAS, Borok, Russia. 23 Plant Science Department, McGill University, Quebec, Canada

The mainly arctic-alpine Carex sect. Racemosae (>60
spp.) constitutes one of the larger, yet taxonomical-
ly most challenging groups within the megadiverse
genus Carex. Previous molecular studies have been
limited by incomplete sampling and have failed to
disentangle the taxonomic delimitation and phylo-
genetic placement of several controversial, doubtful,
or rare species. We achieved a more or less complete

species sampling thanks to a global collaborative ef-
fort, including field campaigns and herbarium work
as well as ongoing taxonomic revisions in Eurasia and
the Neotropics, Divergence-time estimation and mul-
tispecies coalescent (MSC) analyses were conducted
on a multi-accession Sanger-sequencing dataset (i.e.,
multiple samples per species) using nuclear (ITS, ETS)
and plastid (rps16, trnK-matK, trnL-trnF) DNA regions
(ca. 6,000 bp). Biogeographic analyses suggest a Late
Miocene (c. 10 Myr) origin within the forest and moun-
tain-steppe belts of the Northern Hemisphere, followed
by several, partly trans-continental range expansions,
mainly during the early diversification process (10–5
Myr). A trans-Cordilleran disjunction is also evident.
Overall patterns highlight the importance of Central
Asia and Western North America for the group’s diver-
sification. Obtained clades are more congruent with
biogeographic rather than by morphological patterns.
Diversification rate analyses showed an unexpected,
continuous and uniform slowdown in diversification
rates across all major clades, suggesting niche con-
servatism coupled with niche filling. Finally, several tax-
onomic and nomenclatural changes are suggested,
i.e., some regional taxa previously placed in synonymy
of other widespread species require revalidation (e.g.,
C. apodostachya) or remain “orphan species” with no
clear assignment to any specific clade (e.g., C. serre-
ana). In conclusion, our collaborative study provides
fundamental insights into the evolutionary dynamics
of a significant high mountain plant group and lays
the groundwork for a comprehensive global revision
of Carex sect. Racemosae.

S.121.6 Carex phylogenomics
(Cyperaceae) and MADS-
box gene evolution

Sangtae Kim¹, Se-Eun Jung¹, Seungyeon Lee¹, Bora Lee¹, Yanghoon Cho², Tamara Villaverde³, Pedro Jiménez-Mejías⁴, Étienne Léveillé-Bourret⁵, Carmen Benítez Benítez⁶, Bruce A. Ford⁷, Julian R. Starr⁸, Bishnu Adhikari⁹, Sangchul Choi¹, Sungsik Kong¹⁰

1 Department of Biology, Sungshin Women's University, Seoul, South Korea. 2 Uri Plant Research Institute, Kwangju, South Korea. 3 Department of Biology and Geology, Physics and Inorganic Chemistry, Universidad Rey Juan Carlos, Móstoles, Madrid, Spain. 4 Department of Molecular Biology and Biochemical Engineering, Universidad Pablo de Olavide, Seville, Spain. 5 Institut de recherche en biologie végétale and
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Département de sciences biologiques, Université de Montréal, Montréal, Canada. 6 Department of Plant Biology and Ecology, Universidad de Sevilla, Seville, Spain. 7 Department of Biological Sciences, University of Manitoba, Winnipeg, Manitoba, Canada. 8 Department of Biology, University of Ottawa, Ontario, Canada. 9 Department of Biological Sciences, Univ. of Alabama, USA. 10 The Wisconsin Institute for Discovery, University of Wisconsin-Madison, USA

With around 2,000 species, *Carex* (Cyperaceae) is the second-largest genus of monocots. Its great diversity means that extensive taxonomic sampling and genome-scale sequence data are necessary to unravel its detailed evolutionary history and evaluate its newly proposed infrageneric classification. In this study, we estimated both nuclear and plastid phylogenetic trees based on genome-scale data, incorporating 152 representative taxa. We recovered 490 single-copy nuclear gene loci using *Carex*-specific probes through the Hyb-Seq technique and sequenced 71 coding plastid genes through genome skimming. Results showed that four of the six subgenera proposed in the new classification system were highly supported (bootstrap support ≥ 95%) in nuclear and plastid

trees. However, their positions in the plastid and nuclear trees differed for many taxa or clades, suggesting a putative hybrid origin, which was assessed by HyDe and GHDet analyses. We also obtained high-quality genome sequences from five *Carex* species (*C. siderosticta*, *C. paxii*, *C. dickinsii*, *C. breviculmis*, and *C. capricornis*) and analyzed them with two previously reported *Carex* genomes and representatives of other Poales lineages. The genome sequences from *Amborella trichopoda*, *Oryza sativa*, and *Arabidopsis thaliana* were employed as references to detect MADS-box genes in *Carex*. The maximum-likelihood tree, constructed using an amino-acid-aligned DNA matrix, showed unique duplication and deletion events in the evolutionary history of MADS-box genes in *Carex*. Furthermore, we generated transcriptomes for each floral organ from two *Carex* species and compared them with those from *Arabidopsis* and rice. Our findings 1) highlight conflicting phylogenetic relationships among some subgroups, possibly due to the different evolutionary histories of plastid and nuclear genomes, and 2) shed light on floral structure and genome evolution in *Carex*.

**S.122 SOLANACEAE: BIOLOGY, SYSTEMATICS AND EVOLUTION. SESSION 2**

**S.122.1 Untangling the web of Solanum sections Herpystichum and Pteroidea**

Tepe, Eric¹, Jacob Bryant¹

1 Department of Biological Sciences, University of Cincinnati, Cincinnati, USA.

Solanum sections Herpystichum and Pteroidea are small sections of ten species each within the Potato Clade of the enormous genus Solanum. Both sections range from Central America to the central Andes and are heterogeneous groups of species that include herbs and vines, and species with simple and compound leaves. Pteroidea is defined by inflorescences in an axillary position, unifoliate sympodial units, and rugose, conical fruits in most species. This combination of characters is unique in Solanum and lends support to the monophyly of Pteroidea. Herpystichum, on the other hand, is more heterogeneous, but can be characterized by onion-shaped floral buds and distinctly flattened fruits in some species. Phylogenetic analyses

based on nuclear and plastid markers have shown that the two sections form a strongly supported clade, but an understanding of the relationship between the sections has remained elusive. Analysis of concatenated data supports the two sections as reciprocally monophyletic, or with Pteroidea nested within Herpystichum. Similarly, analysis of individual markers recovers Pteroidea as sister, in a polytomy with, or nested within Herpystichum. Our analyses of both nuclear and plastid markers, as well as nearly complete plastid sequence data suggest frequent shifts in leaf complexity in the entire clade and a single origin of sect. Pteroidea. Origins of the flattened fruits of Herpystichum, as well as the monophyly of the section itself, remain enigmatic. The additional data, however, reinforce the previous findings that the widespread species, S. anceps and S. mite are not monophyletic, and both morphology and molecular data suggest that there is at least one undescribed species in sect. Pteroidea. Additional sequence data using ddRAD-seq is underway, which may provide additional insights into the evolution and relationships among these groups.
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## S.122.2 Communities pulling together for 21st century taxonomy: the Solanum (Solanaceae) story

Knapp Sandra¹

¹ Natural History Museum, London, UK.

There are only a handful of genera of angiosperms with more than 1,000 species, and they are often left aside by botanists studying plant diversity. Their sheer size frightens us away from beginning to tackle them; they don't fit into a three- or five-year funding cycle, and making progress, let alone finding a way to begin, can be daunting. But what if we looked at working together in different ways? Not as lone taxonomists working on a group but as a collective – let's call it a "meta-lab" where different sets of people come together for different types of analysis, all with the goal of understanding something a single botanist could never do in a lifetime. Solanum is one such giant genus, with more than 1,400 currently recognised species distributed on all continents but Antarctica. Solanum was last treated as a whole on a global basis in the 19th century; we know a lot more today about both species diversity and relationships. I describe how the Solanum "meta-lab" has approached working on such a monster and show how the community approach facilitates significant progress in taxonomy, phylogeny and the production of identification tools; I also discuss some of our recent advances and where we are going next.

## S.122.3 Distinct assembly patterns of Solanum: insights from the globe and Australia

Ang Li¹, Tiina Särkinen², Brent D. Mishler³,

Li-Min Lu¹, Yun-Feng Li¹, Dan-Xiao Peng¹,

Hai-Hua Hu¹, Zhi-Duan Chen¹

¹ Institute of Botany, Chinese Academy of Sciences, Beijing, China. ² Royal Botanic Garden Edinburgh, UK. ³ University of California, Berkeley, California, USA

Solanum (Solanaceae) is one of the largest genera of angiosperms (containing more than 1,000 species) with worldwide distribution and great economic importance. However, few studies have been conducted on the global and continental distribution patterns of the genus. We here mea-

sured the diversity and endemism patterns of Solanum all over the world and using Australia as an example of the regional scale for its diverse species richness and insular geographical property. Around the globe, we found that Central America to South America has the highest species richness, species endemism, phylogenetic diversity, and phylogenetic endemism, whereas Australia has the second highest diversity and endemism, especially in its eastern coast. The distributions of neo- and paleo-endemism centers are generally congruent with the local volcanic activities and orogenic movements. Within Australia, the hotspots were distributed mainly in the tropical or temperate forests and grasslands and generally congruent with the vegetation types. Based on the clustering analyses, we further identified a certain linkage among Africa, Asia, and Australia, especially the clustering in Asia-Australia showing a pattern of multiple clusters in Philippines and Indonesia, suggesting that Solanum in tropical Asia possibly have multiple origins. By investigating the diversity patterns of Solanum through global to regional scale, our study suggests that diversity patterns of Solanum at the global scale are more likely shaped by the local orogenic movements, while those at regional scale (e.g., Australia) are more likely affected by the vegetation heterogeneity. The possible connection among Africa, Asia, and Australia was figured out to help to reveal the complex and controversial origin of Asian Solanum.

## S.122.4 Functional and ecological diversification of underground organs in Solanum

Edeline Gagnon¹,²,³, Ludwig Baldaszti²,⁴,
Peter Moonlight⁵, Sandra Knapp⁶,
Caroline E.R. Lehmann²,⁴, Tiina Särkinen²

¹ Department of Integrative Biology, University of Guelph, Guelph, ON, Canada. ² Tropical Diversity Section, Royal Botanic Garden Edinburgh, Edinburgh, UK. ³ TUM School of Life Sciences, Technical University of Munich, Freising, Germany. ⁴ School of GeoSciences, University of Edinburgh, Edinburgh, UK. ⁵ Botany, School of Natural Sciences, Trinity College Dublin, Dublin, Ireland. ⁶ Natural History Museum, London, UK.

The evolution of geophytes in response to different environmental stressors is poorly understood largely due to the great morphological variation in
<figure><img src="image_287.png" /></figure>

underground plant organs, which includes species with rhizomatous structures or underground storage organs (USOs). Here we compare the evolution and ecological niche patterns of different geophytic organs in *Solanum* L, classified based on a functional definition and using a clade-based approach with an expert-verified specimen occurrence dataset. Results from PERMANOVA and Phylogenetic ANOVAs indicate that geophytic species occupy drier areas, with rhizomatous species found in the hottest areas whereas species with USOs are restricted to cooler areas in the montane tropics. In addition, rhizomatous species appear to be adapted to fire-driven disturbance, in contrast to species with USOs that appear to be adapted to prolonged climatic disturbance such as unfavorable growing conditions due to drought and cold. We also show that the evolution of rhizome-like structures leads to changes in the relationship between range size and niche breadth. Ancestral state reconstruction shows that in *Solanum* rhizomatous species are evolutionarily more labile compared to species with USOs. Our results suggest that underground organs enable plants to shift their niches towards distinct extreme environmental conditions and have different evolutionary constraints.

**S.122.5 Taming a monster: taxonomic revision of the Torva clade of Solanum (Solanaceae) in the Andes**

Briggitthe Melchor-Castro¹,²,³, Sandra Knapp⁴, Tiina Särkinen¹

1 Royal Botanic Garden Edinburgh, Edinburgh, UK. 2 University of Edinburgh, Old College, South Bridge, Edinburgh, UK. 3 Departamento de Etnobotánica y Botánica Económica del Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Jesús María, Lima, Perú. 4 Department of Life Sciences, Natural History Museum, London, UK.

One third of *Solanum* species remain to be taxonomically revised (~250 of the total 1,200 species). The unrevised species are focused in five clades mostly Neotropical clades with highly plastic leaf morphology and indumentum that is difficult to understand without field observations, large number of specimens, and access to microscopic details. One of these clades is the Torva clade with ca. 68 species and 269 names and which includes the pea eggplant (*S. torvum*) used for its edible fruits. Regional accounts have helped to clarify taxonomy of the

group (e.g., Costa Rica, Africa and Asia), but a set of poorly understood species complexes remain in the tropical Andes especially that have been used as a dumping ground for complex morphological variation that may represent several distinct species. This talk presents a taxonomic overview of the Torva clade in the Andes. Based on a detailed morphological study of herbarium specimens, fieldwork, and nomenclatural revision, we recognise 36 species of the Torva clade in the Andes, of which 7 species are new and 5 are newly resurrected. The clade is most diverse in Peru where 21 species are found. Reassessment of the morphological traits reveals that trichome structure on stem, leaves and flowers is key to species delimitation. Other key characters include floral bud shape, calyx lobe shape, and corolla shape. The morphological species concepts will now be tested using target capture bait sequence data.

**S.122.6 Moving forward on Solanum (Solanaceae) systematics: phylogenomic insights from unarmed lineages**

Leandro Giacomin¹, Juan David Tovar D.², Tiina Särkinen³, Sandra Knapp⁴, Thiago André⁵, Clarisse Palma-Silva⁶, Edeline Gagnon⁷

1 Universidade Federal da Paraíba, João Pessoa, Brazil 2 Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil. 3 Royal Botanic Garden Edinburgh, Edinburgh, UK. 4 Natural History Museum, London, UK. 5 Universidade de Brasília, Brasília, Brazil. 6 Universidade Estadual de Campinas, Campinas, Brazil. 7 University of Guelph, Guelph, Canada

Building robust phylogenetic frameworks is the basis to pose any evolutionary question. Nevertheless, understanding relationships among taxa can also benefit society in many other ways, for example, in crop improvement. The economically important genus *Solanum* (*Solanaceae*) has had significant advances on its systematics in the past decades, where recent works have revealed strong incongruence among phylogenomic datasets and reticulate diversification in deep nodes. We present genomic insights from two unarmed lineages of the genus (clades Brevantherum and Geminata). These lineages together represent >20% of species diversity in *Solanum*. We generated target capture data (353 Angiosperm bait set) for 140 species and retrieved these regions from publicly available genome
<figure><img src="image_288.png" /></figure>

skimming data from another 5 species. Our results show that the 353 bait set can be useful to resolve relationships within genera, but more variable regions are desired for recently diversified lineages like Geminata. The heterogeneous length of recovered target regions seems to influence the support of recovered clades and, as this heterogeneity is related to DNA quality and sample preservation, it might remain as a challenge to any speciose group. Incongruences were found within the 353 bait set alone, and we argue a gene genealog

gy interrogation method should be employed in these cases. Capturing 353 targeted regions from genome skimming data yielded satisfactory amounts of data for these lineages, and it is a promising approach to compare different datasets and reduce bench work time related to sequence capture. We conclude that universal probe sets such as the 353 can be useful for big genera like *Solanum*, considering specific baits can be less cost-effective.

# S.123 EVOLUTION OF KEY INNOVATIONS IN LAND PLANTS REVEALED THROUGH THE LENS OF EVODEVO. SESSION 3

## S.123.1 Genetic underpinnings of growth regulation, flowering and floral organ identity in the holoparasitic Apodanthaceae

Natalia Pabón-Mora¹, Angie D. González², Andrea Ramírez-Ramírez¹, Juan F. Alzate³, ⁴ Elena M. Kramer⁵, Charles C. Davies⁵, Jeff P. Mower⁶, Favio González⁷

1 Instituto de Biología, Universidad de Antioquia, Medellín, Colombia. 2 Departamento de Biología, Facultad de Ciencias, Universidad Nacional de Colombia, Bogotá, Colombia. 3 Centro Nacional de Secuenciación Genómica-CNSG, Sede de Investigación Universitaria-SIU, Universidad de Antioquia, Medellín, Colombia. 4 Grupo de Parasitología, Facultad de Medicina, Universidad de Antioquia, Medellín, Colombia. 5 Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, USA. 6 Center for Plant Science Innovation, University of Nebraska, Lincoln, USA. 7 Universidad Nacional de Colombia, Facultad de Ciencias, Instituto de Ciencias Naturales, Bogotá, Colombia

Plant-plant parasitism has independently evolved 12 times in angiosperms. The ca. 4750 parasitic plants constitute only 2% of all angiosperm diversity; however, they exhibit some of the most outstanding genome reductions and extraordinary modifications in terms of development, structural adaptations, and reproductive strategies. The most dramatic devel

opmental reductions occur in the endoholoparasites, which grow completely inside of their hosts as parenchymatic cells, lack typical roots, shoots and leaves, and become visible only through emerging flowers and fruits. We have focused on documenting the atypical developmental transitions occurring in the Apodanthaceae, specifically in the Salicaceae parasite *Apodanthes* and the Fabaceae parasite *Pilostyles*. These plants have no root (RAM) or shoot (SAM) apical meristems; however, flowers initiate in contact to the host vascular tissue, completely inside the host, and emerge having completed organogenesis. Using comparative genomics, we are currently characterizing plastome reduction and are documenting extensive horizontal gene transfer in the mitogenomes. Using comparative transcriptomics (derived from the host, the parasite, and mixed tissues), we have addressed the mechanisms through which *Pilostyles* maintains stem cells in the absence of primary meristems, the signals perceived for reproductive transitions in the absence of typical photoperiod sensing leaves, and the endogenous capacity for organogenesis in the dark, fully embedded inside of the host. We present data on: 1) the substantial reduction in the SAMs genetic machinery; 2) the regulation of cell division during the endophytic phase until the completion of flower development, 3) potential recruitment of host flowering regulators by the holoparasites; and 4) expression analyses of endogenous *LEAFY* and MADS-box floral organ identity homologs in these atypical flowering plants. We discuss our results in comparison with emerging developmental data from other holoparasitic plants to determine convergent mechanisms in the evolution of parasitism.
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## S.123.2 How do ectopic cambia form? A case study from the legume family

Israel L. Cunha Neto¹, Anthony Snead², Jacob B. Landis³, Chelsea D. Specht³, Joyce G. Onyenedum¹

1 Department of Environmental Studies, New York University, New York, USA. 2 Department of Biology, New York University, New York, USA. 3 Section of Plant Biology and the L.H. Bailey Hortorium, Cornell University, Ithaca, USA

Secondary growth is a remarkable evolutionary innovation in vascular plants, representing a developmental process through which plants increase in thickness. Typically, secondary growth generates a core secondary xylem (wood) surrounded by a sheath of secondary phloem (inner bark), and this pattern is achieved through a delicate balance between cell initiation, proliferation, differentiation, and programmed cell death – all governed by a single stem-cell region called the vascular cambium. Thousands of seed plants share this conserved mode of radial growth, which is regulated by a common set of conserved genes. However, this typical modality of vascular radial growth has been disrupted in several lineages of gymnosperms and angiosperms that contain alternative vascular developmental pathways called ‘vascular variants’. In certain lineages, the presence of vascular variants becomes especially intriguing when the stem produces supplemental vascular cambia in unconventional locations, in addition to the initially formed original vascular cambium. This phenomenon is called ectopic cambia and generates distinct anatomical patterns, most commonly ‘successive cambia’ where concentric rings of cambium and its products encircle the stem/root. This process results in the development of stems or roots with multiple cambia. In this study, we investigate the evolution and development of ectopic cambia, with a particular focus on the Fabaceae. By comparing Phaseolus vulgaris (typical secondary growth) to Wisteria floribunda (ectopic cambia), we combine anatomy with comparative transcriptomes to elucidate the developmental and molecular mechanisms underlying successive cambia formation. We generated seventeen transcriptomes to explore variations in gene expression within and between species, focusing on both typical and ectopic cambia. Moreover, we explore the molecular

evolution of conserved genes related to vascular development (WOX4, CLE41, PXY, ER) to test for signatures of selection, gene duplication, and pseudogenization in lineages with and without successive cambia.

## S.123.3 Comparative single nuclei sequencing of the monocot gynoecium reveal candidate genes underlying septal nectary development in banana

Irene T. Liao¹, Lena C. Hileman², Karen E. Sears¹,³, Lachezar A. Nikolov⁴

1 Department of Molecular, Cell, and Development Biology, University of California – Los Angeles, Los Angeles, CA. 2. Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS. 3 Department of Ecology and Evolutionary Biology, University of California – Los Angeles, Los Angeles, CA. 4. Department of Biology, Indiana University, Bloomington, IN

Nectary function is central to many important ecological interactions. Across flowering plants, nectaries are diverse in positional location and morphology, yet few have used genomic approaches to examine how nectaries develop. Most monocot flowers form septal nectaries, which are located on the margins between the three fused carpels that comprise the gynoecium. While key transcription factors underlying nectary development in eudicots have been characterized, the genes driving monocot nectary development are virtually unknown. As a first step to identifying candidate genes, we are using single nuclei RNA sequencing to obtain cell-type specific expression profiles. We isolated and sequenced banana (Musa "Ice Cream") nuclei from two sets of tissues—one region enriched in nectary cells at the top of gynoecium (the "top") and the other that lacks nectary cells at the base of the gynoecium (the "base")—and asparagus (Asparagus officinalis) nuclei from whole gynoecia. We identified putative nectary cell clusters and genes, and a number of candidate genes will be spatially validated with fluorescence in-situ hybridization. These analyses will determine whether both species share a common core set of genes for nectary development. Results from this study will form the foundation for understanding how nectaries have been conserved in monocots.
S.123.4 A BLADE-ON-PETIOLE
orthologue regulates
corolla differentiation in the
proximal region in Torenia
fournieri

Shihao Su¹, Xuan Zhou¹, Tetsuya Higashiyama²

1 School of Agriculture, Sun Yat-sen University, Shenzhen, China. 2 Department of Biological Sciences, The University of Tokyo, Japan

The three-dimensional shape of a flower is integrated by morphogenesis along different independent axes. Differentiation along the petal proximodistal axis is tightly linked to the specification of pollinators; however, it is still unclear how a petal patterns its proximodistal axis. The corolla of *Torenia fournieri*, exhibits strong differentiation along the proximodistal axis, including a distal petal lobe region, an intermediate corolla tube, and a proximal corolla neck region that is controlled by an ALOG family homologue, TfALOG3. In this study, we report a floral-specific BOP gene, TfBOP2, which is predominantly expressed in the proximal region of the corolla. CRISPR knockout mutants of TfBOP2 have shorter proximal corolla tubes and longer distal lobe regions, with ectopic growth of conical cells in the fusion boundaries of the corolla tube, consistent with its expression pattern as a proximal regulator. Mutants of BOP homologues in *Arabidopsis thaliana* also show developmental defects in the petal proximal region, favouring a shared role of BOPs among different species. Genetic analysis demonstrates the interaction between TfBOP2 and TfALOG3 in the control of corolla proximodistal differentiation since double mutants have the shortest proximal corolla tubes while exhibiting normal boundary fusion. We further found that TfALOG3 physically interacts with TfBOP2 and can recruit TfBOP2 to the nuclear region, indicating its possible function as a complex. Our study favours a hypothetical shared BOP-ALOG complex across the core eudicots that is recruited to regulate corolla differentiation in the proximal region axis of *T. fournieri*.

S.123.5 How do neotropical orchids bloom? a comparative transcriptomic approach in orchids with different habits and storage organs

Yesenia Madrigal¹, Michael Scanlon²,
Marian Bemer³, Juan F. Alzate⁴, Natalia
Pabón-Mora¹

1 Instituto de Biología, Universidad de Antioquia, Medellín, Colombia. 2 School of Integrative Plant Science, Cornell University, Ithaca, US. 3 Plant Developmental Systems, Wageningen University and Research, Wageningen, the Netherlands. 4 Centro Nacional de Secuenciación Genómica, SIU, Universidad de Antioquia, Medellín, Colombia

Flowering in angiosperms occurs when vegetative meristems (VM) that form leaves become inflorescence meristems (IM) that form bracts and flowers. This process has been comprehensively studied in monocots, such as Oryza sativa, where the genetic regulatory network (GRN) includes promoter genes such as FT, COL and FUL that activate floral meristem identity genes. There are also repressors involved in the maintenance of vegetative phases that include TFL1 and AGL24/SVP, in addition to FLC and VRN2 that are important in response to cold. In contrast, little is known about how flowering is genetically controlled in neotropical orchids with different habits that colonize different micro-niches and respond to subtle environmental changes. We performed an exhaustive analysis of the morpho-anatomical and transcriptomic changes from VM to IM in three orchids: Epidendrum fimbriatum (minature terrestrial), Cattleya trianae (epiphytic with storage organs), and Elleanthus aurantiacus (a big size terrestrial). Our RNAseq results show a higher proportion of differentially expressed genes (DEGs) involved in flowering in terrestrial species compared to epiphytic taxa. Conversely, a greater number of genes related to the metabolism of sugars and lipids was found in the epiphytic species. The most important flowering transcription factors within the DEGs include the promoters FT2A, FD2A, FUL1B/1A, SOC1La, LFY and the repressors COL4A/4B, SVP2A, GHD7, AP2 and TFL1. We validated our results using spatiotemporal expression by ISH and by protein-protein interaction using Y2H. We found important differ-
<figure><img src="image_290.png" /></figure>

ences between the flowering GRN in orchids and model crops in the Poaceae: 1) high duplication rates for flower integrators in orchids but a low percentage of transcriptionally active homologs; 2) the retention of canonical flowering integra-

tors, at the expense of low expression, loss of key protein interactions, and possibly pseudogenization of some homologs; and 3) changes in transcriptomic profiles in different orchids according to their morphological adaptations and habits.

# S.124 ETHNOBOTANY AND CONSERVATION OF MEDICINAL PLANTS. SESSION 3

## S.124.1 Medicinal plants profile of the Mapulana people of Ehlanzeni District in Mpumalanga Province, South Africa

Shalom P. Mashile¹, Milingoni P. Tshisikhawe¹, Ndivhaleni A. Masevhe¹

1 Department of Biological Sciences, University of Venda, Thohoyandou, South Africa

South African government has declared public health care free for all citizens. Traditional medicine is preferred as supplement to avoid challenges faced by the public health care system; most people rely on both health-care systems. A semi-structured questionnaire was administered to participants who included elderly people, community adults and youths. Data was analysed by calculating the use value and informants' consensus factor. The study revealed that the elderly are custodians of traditional medicine. Hundred and six plant species were identified in treatment of 50 different ailments. The recorded medicinal plant species were distributed amongst 55 plant families with Fabaceae (20%), Crassulaceae (5%) and Cucurbitaceae (5%) as dominating families. Plant parts such as roots (48%), leaves (26%) and barks (15%) were collected from herbs (45%), trees (37%) and shrubs (18%). *Aloe zebrina* (0.38), *Aloe marlothii* (0.29), *Sclerocarya birrea* subsp. *caffra* (0.28) and *Siphonochilus aethiopicus* (0.28) use values were the highest. Gonorrhea (1.80), stomach cleansing (1.40) chicken diseases (1.19), bad luck (0.88), flu (0.84), and diarrhea (0.80) received the highest informants' consensus factor (ICF). Plant species with low use value and ICF were known by the elder and few community adults.

## S.124.2 Ethnobotanical Exploration: Assessing antifeedant and phytochemical properties of selected plants against Spodoptera frugiperda

K.C. Shilaluke, D.T Ndinteh, A. Moteetee

University of Johannesburg, Johannesburg, South Africa

Environmental and health hazards linked to synthetic pesticides have prompted a growing interest in botanical insecticides as safer and environmentally friendly alternatives for controlling insect pests in agriculture. Following the historical utilization of plants in folk remedies and their continued application in diverse communities, this study explores the insecticidal potential of nine traditionally recognized plants as insecticides against the *Spodoptera frugiperda* larvae. The feeding deterrence activity of methanol and hexane extracts was evaluated using a choice test bioassay where maize leaves were used as test food. *Podalyria sericea* and *Cassytha ciliolata* hexane extracts were the most toxic to the larvae, with LC₅₀ values of 0.779 and 0.993 mg/ml, respectively, as compared to the methanol extracts LC₅₀ values of 1.853 and 1.461 mg/ml, respectively. Subsequently, the most efficient insecticidal extracts underwent further chemical analysis for compound isolation, purification, and structural elucidation of secondary metabolites. The efficacy of the isolated compounds with insecticidal activity will be evaluated and compared with synthetic insecticides, paving the way for a comprehensive assessment of their potential use as sustainable alternatives in pest management.
<figure><img src="image_291.png" /></figure>

### S.124.3 Phase 5: Status of ethnobiology and study perspectives in Albania

Ani Bajrami¹, Anyla Saraci², Besnik Rexhepi³, Robert Damo⁴

1 Museum of Natural Sciences "Sabiha Kasimati", University of Tirana, Tirana, Albania. 2 Department of Language and Literature, "Fan S. Noli" University, Korce, Albania. 3 Department of Biology, University of Tetova, Tetova, Republic of North Macedonia. 4 Department of Agronomy, "Fan S. Noli" University, Albania

The science of ethnobiology has been influenced by theories and methods belonging to various scientific disciplines, giving it an interdisciplinary nature. In this article, we address the main trends and new directions in ethnobiology, focusing on the contribution of evolutionary theories that have altered the conceptual and interpretative framework of ethnobiology today. Additionally, we outline the main phases of ethnobiology in Albania, with a focus on ethnobotany, from its beginnings to the present day. From here on, in accordance with the theoretical development of contemporary ethnobiology in Europe and the USA, we attempt to create a local research framework and explore the ways in which ethnobiological studies may develop in the future in Albania.

### S.124.4 In vitro culture and hairy root induction in Solanum virginianum: an important component of 'Dashamoola' an ayurvedic multi-ingred

Charushila M. Dalavi¹, Jalaja Naravula², P.B. Kavi Kishor³, Pratik S. Kamble¹, Swaroopa A. Patil¹

1 Shivaji University, Kolhapur, India 2 Vignan's Foundation for Science, Technology and Research (Deemed to be University), Guntur, India 3 Osmania University, Hyderabad, India

Solanum virginianum L. (Solanaceae) is one among the Dashamoola group of plants used in Ayurveda. Dasmul Asava and Dasmularishta are well known ayurvedic formulation used as analgesic and anti-inflammatory. The roots of S. virginianum are used

in these formulations and therefore, there is a need to develop an efficient protocol for *in vitro* multiplication of this species so that roots can be obtained on a large scale within a short time span. Accordingly, we standardized the *in vitro* protocol for this species using nodal explants. MS (Murashige and Skoog) medium supplemented with different concentrations and combinations of cytokinins was used for shoot initiation. MS supplemented with 0.75 mg l⁻¹ BA ideally yielded 31.7±1.33 shoots/explant (p<0.05) with a response of 95%. Combination of BA (0.75 mg l⁻¹) and TDZ (1 mg l⁻¹) produced the highest number of shoots (63.91±5.19 shoots/explant) with 90% response. Subculturing the shoots on different concentrations of BA led to 100% shoot multiplication with an average shoot number of 26.25±1.31 shoots/explant and average shoot length 4.43±0.26 cm. MS augmented with IBA (2.5 mg l⁻¹) and IAA (1.5 mg l⁻¹) was used for *in vitro* rooting of shoots that yielded 26.83±1.27 roots/shoot with 92% rooting response and the average root length 6.25±0.33 cm. Agrobacterium rhizogenes mediated hairy root induction was successfully done in leaf and stem explants of S. virginianum using ATCC 15834 strain. The transformation was confirmed by Polymerase Chain Reaction (PCR). Initiation of hairy roots from both leaf and stem explants was recorded but in varied frequencies (leaf 37.77%, 5.88±0.22 hairy roots/explant and stem-21.33% with 3.18±0.29 hairy roots/explant). proliferative growth of hairy roots was achieved in the 9th passage (188.94±3.65 hairy roots/leaf explant) and in 6th passage (115.77±6.62 hairy roots/stem explant).

### S.124.5 The bitter chemicals in the tip shoots of Bambusa oldhamii Munro (Green bamboo)

Jiao, Yulian

Zhejiang Institute of Subtropical Crops, Zhejiang Academy of Agricultural Sciences

When the young bamboo shoots of Bambusa oldhamii have their best taste, they are rich in nutrition and widely used in traditional Chinese cuisines. But the shoots erupted out of the soil surface with sunlight for several days, the flavor would be changed to a little bitterness in the tip shoot which would decrease the price in the market.&nbsp; Our research was aimed at mining out the bitter chemicals that would influence the bitter flavor when the shoot of
green bamboo grew from underground to above ground. In our previous research based on three developing bamboo shoot phases, we mined several bitter chemicals counting for the flavor transformation, such as Solanidine, Amygdalin, Salicin, Arbutin, and others from metabolome data, and the AP2/ ERF transcription factor family and the pathway of Biosynthesis of phenylpropanoids might play a key role in the biological process based on the correlation analysis with gene expression data corresponded with each phases. To explain the biological question more comprehensively and with multiple perspectives, the design and experiment proposal was improved. The shoot development phases of Bambusa oldhamii were extended to seven phases,

and the transcriptome data was assembled based on the genome reference, moreover, the detection of the metabolites has a different method, finally, we mined more than 30 bitter chemicals. The bitter chemicals were distributed in the family of Flavonoids, Terpenoids, Amino acids, Purines and derivatives, Carbohydrates, and others. With the correlation analysis between the data of metabolome and transcription factor expression data with corresponding phases, we found that bHLH and HB would be the major transcription factor family, AUX/IAA would be the major transcription regulation factor family, and Leucine Rich Repeat Kinase XI &amp; XII would be the major protein kinase family participate in the biological process of flavor taste transition.

# S.125 PLANT DIVERSITY, BIOGEOGRAPHY AND EVOLUTION IN THE TROPICS FOR CONSERVATION, RESTORATION AND SUSTAINABLE USE. SESSION 4

## S.125.1 The key role of the seasonally inundated forest in the conservation of biodiversity in southeastern Mexico

Alexis Herminio Plasencia-Vázquez<fnref n="1" />,
Alfredo García-González<fnref n="1" />,
Annery Serrano Rodríguez²,
Frander B. Riverón-Giró³,
Yarelys Ferrer-Sánchez⁴,
Anay Serrano Rodríguez<fnref n="1" />,
Griselda Escalona-Segura⁵,
José Domingo Cú-Vizcarra⁶,
Karla Paola Borges-Jesús⁶,
Pedro Zamora Crescencio<fnref n="1" />,
Jorge Albino Vargas-Contreras²

Wetlands are one of the most productive ecosystems in the world, supporting a large biodiversity. Tintal (seasonally inundated forest dominated by *Haematoxylum campechianum*) is a characteristic wetland of the Southeastern Mexico and have a long history of commercial exploitation. Due to its structural characteristics and the conditions of the areas where it develops, tintal could work as a critical habitat for many plants and animal species, and as a refuge, mainly in highly anthropized landscapes. Furthermore, as the tintal develops at sites where water disappears totally or partially during determined times of the year, and the characteristic vegetation favors the formation of open spaces, these areas can be easily accessed to carry out sampling. Therefore, the tintal is the ideal case study in order to demonstrate the impact of human activities on wetland ecosystems. We analyzed the variations in the richness and abundance of different plants (vascular epiphytes) and animals (macroinvertebrates, amphibians, reptiles and bats) between preserved and human-disturbed tintales. The study was carried out at 27 tintal sites in the Southeastern Mexico, 10 preserved and 17 human-disturbed, and for each group different sampling techniques were used. The species composition and structure differ between preserved and human-disturbed tintales. In the case of vascular epiphyte species, the rich-

<footnote n="1">1 Centro de Investigaciones Históricas y Sociales, Universidad Autónoma de Campeche, Campeche, Mexico. 2 Facultad de Ciencias Químico Biológicas, Universidad Autónoma de Campeche, Campeche, Mexico. 3 El Colegio de La Frontera Sur (ECOSUR), Universidad Tapachula, Tapachula, Chiapas, Mexico. 4 Facultad de Ciencias de la Ingeniería, Universidad Técnica Estatal de Quevedo, Quevedo, Ecuador. 5 El Colegio de La Frontera Sur (ECOSUR), Unidad Campeche, Ciudad Industrial, Lerma, Campeche, México. 6 Instituto de Ecología, Xalapa Enríquez, Mexico.</footnote>
<figure><img src="image_292.png" /></figure>

ness and abundance were related to a degree of conservation, and the tintal can be an important reservoir of these plants. Most of the researched tintales were highly anthropized, but the tolerance of several animal and plant species to these landscape modifications has allowed them to make use of such areas. The function of tintales as islands and sources of diversity in degraded land matrices and *H. campechianum* must be considered as a key species. This allows as to value the importance of tintal from an ecological and biodiversity conservation point of view.

## S.125.2 Epiphytic bryophytes as indicators of successional changes in tropical lowland rainforests in Sri Lanka

H.A.C. Dilrukshi¹, S.C.K. Rubasinghe², N.C.S. Ruklani²

1 Postgraduate Institute of Science, University of Peradeniya, Kandy, Sri Lanka. 2 Department of Botany, Faculty of Science, University of Peradeniya, Kandy, Sri Lanka

Epiphytic bryophytes stand out as vital indicators of ecosystem transformations in rainforests due to their heightened sensitivity to environmental shifts. Succession, the gradual evolution of plant communities over time leading to the establishment of a mature and stable ecosystem, can be effectively monitored by tracking the presence and abundance of specific species. This study aims to investigate the response of epiphytic bryophytes to forest succession in three lowland rainforests in the Southern Province of Sri Lanka; Wilpita, Kottawa, and Pituwala. The assessment of bryophytes on 135 trees across three succession stages (early, intermediate, late) in each forest site was conducted, and the impacts of forest succession stage, tree species richness, and diameter at breast height on epiphytic bryophyte species richness per plot were analyzed using generalized linear mixed models. Results revealed the identification of 73 bryophyte species across the three forests. The diversity of epiphytic bryophytes exhibited a pattern, being highest in late succession stages and lowest in early succession stages across all three forest sites. The families Calymperaceae, Sematophyllaceae, and Jungermanniaceae emerged as the most abundant among bryophytes in the three lowland rainforests. A significant revelation of

this study is the identification, for the first time, of the pivotal role played by tree species richness in influencing the richness of epiphytic bryophytes in the Kottawa, Pituwala, and Wilpita forests. Notably, higher tree species diversity in late succession forests is found to be a key factor contributing substantially to the conservation of epiphytic bryophytes in lowland rainforests. This study underscores the importance of exploring epiphytic bryophytes in tropical lowland rainforests in Sri Lanka, offering valuable insights into successional processes and ecosystem dynamics. The obtained information holds crucial implications for the formulation of effective conservation and management strategies, particularly in light of the persisting environmental changes.

## S.125.3 Cascading effects of defaunation on vertebrate-dispersed plant communities in Madagascar

Yuanshu Pu¹,², Alexander Zizka⁴, Renske E. Onstein²,³

1 Univeristy of Leipzig, Leipzig, DE (yuanshu.pu@evobio.eu). 2 German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, DE. 3 Naturalis Biodiversity Center, Leiden, NL. 4 Philipps-University Marburg, Marburg, DE

Madagascar's biodiversity faces various threats leading to defaunation and biodiversity loss. However, how defaunation affects biotic interactions and hence ecosystem functions, such as seed dispersal of fleshy fruited plants by vertebrate (frugivory), remains unclear. We used structural equation modelling to test whether the loss of trait matching between fruit-eating and seed dispersing animals (frugivores) and fleshy-fruited vertebrate-dispersed plants (i.e., dysfunctional seed dispersal) has led to high plant extinction risk, and whether protected areas in Madagascar conserve seed dispersal interactions. We integrated distribution data, dispersal traits, and IUCN Red List status of 2,852 vertebrate-dispersed plant species and 129 frugivore species in 649 communities across Madagascar. We detected a loss of seed size to body size trait matching in non-protected areas, suggesting that many large-seeded plants do not co-occur with frugivores that are able to swallow and effectively disperse the seeds. Interestingly, such 'dysfunctional' communities also showed the largest proportion of
<figure><img src="image_293.png" /></figure>

threatened plants, but only when they were deple-
ed of all large-bodied frugivorous lemurs, with birds
or bats left as the largest seed dispersers. We also
found that protected areas conserve lemur-dom-
inated communities with the highest proportion of
threatened plants, but not the highest seed disper-
sal dysfunctionality. In these communities higher
dysfunctionality did not lead to higher threat. This
suggests that increased plant extinction risk is linked
to defaunation as well as other stressors. Our results
highlight the importance of biotic interactions for
ecosystem functioning and plant extinction risk in
Madagascar, emphasizing how timely monitoring
ecosystem functioning of degraded ecosystems
may help conserve biodiversity.

S.125.4 Biodiversity of
terrestrial chlorophytes
in Yogyakarta Province,
Indonesia

Ludmilla F. Untari¹*, Dita Rahmadani², Faiq
I. Majid², Juntria R. Khasanah², Japareng
Lulang³, Ari Satia Nugraha⁴

1 The Plant Systematics Laboratory, Department of Tropical Biology, Faculty of Biology, Universitas Gadjah Mada, Jl. Teknika Selatan, Sekip Utara, Yogyakarta, Indonesia. 2 Department of Tropical Biology, Faculty of Biology, Universitas Gadjah Mada, Jl. Teknika Selatan, Sekip Utara, Yogyakarta, Indonesia. 3 School of Industrial Technology, Universiti Sains Malaysia, Jalan Sungai 2, Gelugor, Pulau Pinang, Malaysia. 4 The Faculty of Pharmacy, University of Jember, Krajan Timur, Sumbersari, Kec. Sumbersari, Kabupaten Jember, Jawa Timur, Indonesia. *Corresponding author. ludmilla.untari@ugm.mail.ac.id

Algae can be found in a variety of habitats, including terrestrial habitats. Chlorophytes are algae that are often found as the main component of terrestrial algal populations. Molecular data also explain that the diversity of terrestrial algae is much higher than the others. However, several problems in studying terrestrial algae make the study of terrestrial green algae challenging. As a result, it is difficult to find data on terrestrial green algae and their species diversity, especially in Indonesia. More research is needed on the diversity of terrestrial Chlorophyta species in Indonesia. Therefore, this research was conducted to determine the diversity of terrestrial Chlorophyta species in Indonesia, especially in the Yogyakarta Province. At the time of sampling, physicochemical parameters of the growth environment were also

measured. The algae samples were examined un-
der a light microscope and morphological iden-
tification was analysed. Diversity and Evenness In-
dex were also calculated and presented. Species
descriptions and identification keys of twenty-one
species grouped into eleven families are reported
here. This paper is the first report on terrestrial algae
from Yogyakarta Province. The environmental fac-
tors such as temperature, humidity, wind speed and
light intensity affect the growth and density of ter-
restrial algae. However, each species has different
tolerance to these environmental factors.

S.125.5 Floristic wealth of
North Eastern Region (NER)
of India - an aura of nature

Debjyoti Bhattacharyya¹

1 Department of Life Science & Bioinformatics, Assam University, Silchar, Assam, India

North Eastern Region (NER) of India comprises eight States viz. Arunachal Pradesh, Assam, Manipur, Meghalaya, Mizoram, Nagaland, Sikkim and Tripura. The region covers an area of 2,62,179 sq. km, of which, about 66% is under forest cover. Occupying only 7.7% of the total geographical area of India, the region supports about 50% of the total flora of the country. Blessed with tropical humid climate, vast altitudinal variation, dynamic ecosystems and geographical isolation, the area serves as a rich repository of wild floras. Two important biodiversity hotspots, the Himalaya and the Indo Burma cover the region which contains a globally significant array of unique flora with high endemism. Diverse forest types ranging from tropical to sub-tropical broad-leaved forests, temperate, sub-alpine to alpine meadows host various plant species. These include gigantic trees and lianas in the foot hills of Himalayas to rhododendron scrubs at the high-altitude alpine region. Composition of about 8000 flowering plants with high percentage of endemic taxa has made the region an area of interest for the conservationists and nature lovers. The area has been identified as the home for a wide range of Orchids, Grasses and Bamboos, Gingers, Rhododendrons, Primulas, *Impatiens*, Begonias, *Pedicularis* and so many. Many commercially valuable hardwood trees, medicinal and aromatic species, ethno-medicinally important taxa used in the traditional systems are profusely found in the region. Still many areas are underexplored and even unexplored due to dense forests and tough terrain.
<figure><img src="image_294.png" /></figure>

Exploration to the areas in future may open up new vistas for discovery of many new taxa. India has set up a wide network of conservation programmes for sustenance of biodiversity in the region along with prioritization of preserving ecosystems and their natural resources. In this paper, floristic diversity of the region is discussed with their challenges and opportunities of conservation.

**S.125.6 Climatic determinants of Andean forests functional composition**

Báez Selene¹, Hans Verbeeck², Marijn

Bauters², Sonia Palacios³, Sebastián Tello⁴,

Homeier Jürgen⁵,⁶

¹ Escuela Politécnica Nacional, Facultad de Ciencias, Quito, Ecuador. ² Department of Environment, CAVELab - Computational and Applied Vegetation Ecology, Ghent University, Gent, Belgium. ³ Universidad Agraria La Molina, Facultad de Ciencias Forestales, Lima, Perú. ⁴ Missouri Botanical Garden, MO, USA. ⁵ HAWK Göttingen, Faculty of Resource Management, Göttingen, Germany. ⁶ Georg August University Göttingen, Plant Ecology and Ecosystems Research, Göttingen, Germany

Trait composition of tropical forests are closely related to climatic factors. Andean forests are extremely diverse and occur over large areas with wide climatic variation. There is a general understanding of how trait composition of Andean forests varies across climatic gradients. However, detailed studies relating key plant functional traits -that could provide information about forest sensitivity to changing climatic conditions- are inexistent thus far. In this study, we used 86 permanent forest plots established in Ecuador, Perú and Bolivia. For each plot, we calculated community-weighted means (CWM) for the following traits: Leaf nitrogen, Specific Leaf Area (SLA) and Specific stem density (SSD). We used linear models to predict how trait varied in response to climatic variables obtained from extrapolated databases (i.e., mean annual temperature (MAT), mean annual precipitation (MAP), and vapor pressure deficit (VPD). Our results indicated that Andean forests trait expression is tightly linked to MAT and VPD, but the role of MAP was unclear. These findings confirm the critical role of climate on Andean forest's functional configuration.

**S.126 ALPINE PLANT ECOLOGY AND DIVERSITY: FROM SPECIES TO ECOSYSTEMS. SESSION 3**

**S.126.1 Can alpine species take the heat?: Impacts of increased temperatures on early life stages**

Alexandra Seglias¹

¹ Department of Research and Conservation, Denver Botanic Gardens, Denver, USA

Climate change threatens plant biodiversity worldwide. Alpine plant species are particularly vulnerable to climate change, as temperature fluctuations are projected to be most severe in high elevation areas. Even small shifts in climate can have major consequences on phenology, reproduction, fitness, and community composition. Early life stages (seed germination and seedling survival) are arguably the most important processes in not only the fitness of an individual plant, but also in the dynamics of

persistence of plant populations. These initial developmental stages are expected to be more vulnerable to changes in climate than adult life stages and thereby represent a major bottleneck to recruitment and population responses as climate change increases in severity. To understand how the early life stages of alpine plant species will respond to warming temperatures, I used climate-controlled incubators at three different temperature regimes. Temperatures were based on current conditions and projected future conditions under moderate and severe scenarios. I found that two rare alpine species (endemic to the Rocky Mountains of the USA), Physaria alpina and Townsendia rothrockii, performed better under warmer temperatures as compared to control temperatures at both the germination and seedling stage. Germination proportion, number of leaves, and the length of the longest leaf were all higher on average under warm conditions for both species. The results show that early life stages of al-
<figure><img src="image_295.png" /></figure>

pine plants may not be at high risk from warming temperatures, however there are many other shifting climatic factors resulting from climate change beyond temperature alone. Precipitation, snowpack, freeze/thaw cycles, and length/variability of seasons will all impact alpine species. Furthermore, it is important to understand the entire life cycle of the species and how conditions may impact plants at later life stages. Future studies aim to tease apart these considerations.

## S.126.2 Daily functional dynamics of alpine plants at chloroplast level

Miren I. Arzac¹, Jon Miranda-Apodaca¹, Laura González-Cosío¹, Usue Pérez-López¹, Antonio Hernández¹, Beatriz Fernández-Marín¹, José I. García Plazaola¹

1 Department of Plant Biology and Ecology, University of the Basque Country, Leioa, Spain

Alpine plants are exposed to very contrasting conditions of irradiance and UV radiation imposed by a day/night regime during the summer period. To maintain their photosynthetic efficiency under these conditions, alpine plants need to dissipate the excess energy and scavenge the reactive oxygen species (ROS) formed, via different photoprotective mechanisms. At chloroplast level, some carotenoids such as those involved in the xanthophyll cycle directly participate in the dissipation of excess energy. Some of them (such as zeaxanthin) are additionally strong antioxidants that act together with other antioxidant molecules (e.g. lutein, β-carotene and α-tocopherol). These lipophilic antioxidants are associated with plastoglobules, lipoprotein structures attached to thylakoid membranes that respond to stress conditions by changing their size and number. While daily modulation of some protective mechanisms such as the xanthophyll cycle are rather well known, daily oscillations in other molecules (tocopherols) and structures (plastoglobules) are virtually unexplored. In the present study, we aim at unraveling how the leaves of alpine plants are able to thrive under such strong diurnal oscillations, and in particular, what the role of those lipophilic antioxidants and their holding structures (plastoglobules) is on such responses. For this purpose, we studied five alpine species during a day/night cycle (Ranunculus glaucis, Saxifraga oppositifolia, Veronica alpina, Geum reptans and Doronicum grandiflorum) natu-

rally growing in a talus field at 2630 m a.s.l. near Col du Galibier (French Alps). Combining physiological, biochemical and ultrastructural analyses our results show daily changes in plastoglobule size and number. The analysed antioxidants also presented oscillations, mainly the xanthophyll cycle. However, the changes were species-dependent, indicating possible different strategies among alpine plants living under similar conditions. We conclude that thylakoid membranes and their components present the ability to adapt according to daily changes of high irradiance in different species of alpine plants.

## S.126.3 Adapting to extremes: cold stress tolerance mechanisms in montane plant communities of the Pyrenees

Melanie Morales¹,², Sergi Munné Bosch¹,²

1 Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals, Universitat de Barcelona, Barcelona, Spain. 2 Institut de Recerca en Biodiversitat, Universitat de Barcelona, Barcelona, Spain

Montane plant communities like those growing in the Pyrenees must face multiple and contrasting abiotic stresses throughout their life cycle. Particularly, during winter and cloudless sunny days, these species have to cope harsh low temperatures combined to high solar radiation, both promoting photo-oxidative damage. Extreme temperatures are a main factor limiting plant growth in montane habitats, besides the high risk of frost damage especially in open sites where wind may have a severe drying effect, often in snow-free sites. Here, we studied the cold stress tolerance mechanisms of four diverse perennial herbs inhabiting the Pyrenees: *Anthyllis vulneraria* L, *Leontodon hispidus* L, *Plantago lanceolata* L. and *Scabiosa columbaria*, with an emphasis on photoprotective and antioxidant, and their association with stress-related phytohormonal responses. The study was performed in a Pyrenean plant community located in a south-facing slope near Sispony (Andorra, Spain) at 1660 m.a.s.l., and a set of functional traits, including leaf gas exchange, photosynthetic pigments and photo- and antioxidant molecules (anthocyanins, carotenoids and α-tocopherol) were measured together with stress-related phytohormones (ABA, salicylic acid and jasmonates), during autumn (before), winter (during) and spring (after cold stress). All species
<figure><img src="image_296.png" /></figure>

studied were efficient in the prevention of photoin-
hibition during winter, as indicated by values of the
maximum efficiency of PSII (Fv/Fm) above 0.75 during
winter. However, they distinctly respond to cold-in-
duced desiccation stress, as indicated by reduc-
tions in the relative water content (RWC), which
could recover during spring in A. vulneraria and L.
hispidus, but not in P. lanceolata and S. columbaria.
ABA and jasmonates confirmed the differential be-
havior and increased stress sensitivity of A. vulnerar-
ia and L. hispidus. compared to that of P. lanceolata
and S. columbaria. This knowledge is very valuable
for predicting how global warming may impact the
Pyrenean plant communities.

transport and retention capacity. Thus, they could
be considered an adaptation to drought. Plasticity
varied according to donor populations, suggesting
a region-specific adaptive potential. However, we
could not demonstrate phenotypic plasticity when
transplanting samples to lower altitudes, but only to
drought conditions. Extended warm and dry sum-
mers compromise Sphagnum hummocks in the cli-
matic edge of distribution. The outcome of our study
suggests that Sphagnum capillifolium would have
the potential for adaptation to future climatic sce-
narios to some extent. Furthermore, the experiment
showed how hummock restorations are feasible us-
ing near populations as donors.

S.126.4 Phenotypic plasticity to warming and drought in a Sphagnum species

Eulàlia Pladevall-Izard¹, Pau Montané¹,
Jaume Espuny¹, Aaron Pérez-Haase¹

1 University of Barcelona, Barcelona, Spain

Phenotypic plasticity enables species to adapt to
fast environmental changes. Therefore, it may pro-
vide a fitness advantage at a short-term response
and may evolve through natural selection at long-
term scales. Studying phenotypic plasticity in a cli-
mate change simulation experiment would allow
predicting the viability of populations in their cur-
rent distribution limits facing global change. Mires
are sensitive ecosystems and peat mosses growth
determine the ecosystem functioning. In Sphagnum
hummocks, maintaining moisture in the apical part
of shoots compromises Sphagnum survival, growth,
and sexual reproduction. Therefore, many anatomi-
cal and morphological traits are key to retain water
from precipitation. We carried out a field experiment
on Sphagnum capillifolium hummocks in the Pyre-
nees. We simulated global warming by transplanting
Sphagnum shoots from two mires at high altitudes
(2190 m and 2340 m a.s.l.) to a subalpine mire (1770
m). We simulated drought by placing transparent
roofs and digging ditches around hummocks. After
two growing seasons, we analysed the performance
of nine anatomical functional traits of Sphagnum
shoots from the experiment and of other Sphagnum
samples from the donor populations. Sphagnum
shoots showed significant changes under drought
conditions: leaf hyaline cells were wider and lon-
ger, and they had more but smaller pores. These
changes coincide with a greater water absorption,

S.126.5 Plant diversity survey and progress in the flora of Pan-Himalaya

Qiang Wang¹, De-Yuan Hong¹

1 Institute of Botany, Chinese Academy of Sciences

Pan-Himalaya covers the northeastern Hindu-Kush,
Karakoram, Himalaya, and Hengduan mountains.
These majestic mountain ranges form the high-
est region in the world, which is called ‘the roof of
the world’. Three biodiversity hotspots are wholly or
partially included in Pan-Himalaya, i.e., Himalaya,
Mountains of Southwest China, and Indo-Burma.
However, although this region is an important and
indispensable source of biodiversity, the plant diver-
sity of Pan-Himalaya is poorly known. There is even
no complete record of the flora for this most unique
region. The aim of the Flora of Pan-Himalaya (FPH)
project is to understand the plant diversity of Pan-Hi-
malaya and present the first complete record of the
flora of Pan-Himalaya. For the last ten years, the FPH
project has organized 15 comprehensive expeditions
(>10 taxonomists) and more than 400 teams (each
consisting of 3–5 taxonomists) and has investigated
nearly all areas of Pan-Himalaya (except those ar-
eas in Northeast Afghanistan). Members of these ex-
peditions have collected more than 150, 000 speci-
mens, and have taken more than 200, 000 pictures of
vascular plants in Pan-Himalaya. Observation data,
living seeds, and silica gel-dried leaves have been
collected as well. New or updated illustrations have
been drawn for more than 1, 000 representative spe-
cies. Based on extensive field work and integrative
phylogenetic analysis, one new family, 10 new gen-
era, and more than 100 new species from Pan-Hi-
malaya have been described. A sound knowledge of
<figure><img src="image_297.png" /></figure>

the plant diversity of Pan-Himalaya is the only way to achieve sustainability, and it is also essential for ecological civilization development, which is now an important national policy of China. The FPH project is the first and key step in improving our knowledge of the plant diversity in Pan-Himalaya.

S.126.6 Variable
phenological responses
of High Andean species
to artificial warming in
Antisana Volcano

Priscilla Muriel¹, Gabriel Sáenz¹, Carolina Tovar²

1 Laboratorio de Ecofisiología, Escuela de Ciencias Biológicas, Pontificia Universidad Católica del Ecuador, Ecuador. 2 Department of Biodiversity Informatics and Spatial Analysis, Royal Botanical Gardens, Kew, Surrey, UK

Climate warming globally affects plant communi-
ties, affecting their distribution, development, pro-
ductivity and phenological patterns. However, stud-
ies of phenological responses to warming in species
of equatorial regions are very scarce. In our investi-
gation at Antisana Volcano, Ecuador, we character-
ized the reproductive phenophases (buds, flower-
ing, and fruiting) of 12 selected plants in high altitude
grasslands (páramo) under passive warming using
Open Top Chambers (OTC). Objectives included
assessing if reproductive seasons coincide among
12 selected páramo species with different growth
forms, determining if artificial warming induces phe-
nological shifts, and understanding its impact on re-
productive structure abundance. We monitored the
phenology of 12 species every 22 days over a year in
23 control plots, and 23 plots under passive warm-
ing. We used circular statistics to characterize and
compare the phenology of the different species,
and we used non-parametric variance analyses
to assess if there is a significant phenological shift
related to warming for the selected species. Results
revealed diverse seasonality patterns, with sea-
sonal, aseasonal, and species with mixed patterns.
Seasonal species exhibited a higher abundance of
reproductive structures between May and August,
indicating a "reproductive season," independent of
growth form. Contrary to temperate regions, where
temperature increase led to earlier flowering and
fruiting of many species, we found no significant
phenological shift in plants under passive warm-

ing in comparison to control. However, significant
differences in reproductive structure abundance
emerged between control and warming treatments
among species, with five producing more structures
in control plots and three within chambers. This
suggests that warming in equatorial regions may
consistently affect the production of reproductive
structures in páramo species, instead of their flow-
ering and fruiting seasons, thus potentially altering
community composition in the future. The hetero-
geneous response underscores the urgency of as-
sessing a larger number of species for conservation
measures in this highly endemic and diverse region.
# S.127 FUNCTIONAL ASPECTS OF NECTAR-MEDIATED PLANT-POLLINATOR INTERACTIONS

## S.127.1 Where the bee sips: the perception of nectar by eusocial bees

Geraldine A. Wright¹

1 Department of Biology, University of Oxford, Oxford, UK

Floral nectar and pollen are the currencies of the transactions that occur among flowering plants and their animal pollinators. The sense of taste allows pollinators to make rapid decisions about food quality. Specialist feeders like bees, however, that feed on nectar and pollen may have adaptations that permit them to have greater acuity for compounds like sugars that are relevant to their survival. We know surprisingly little about the way that the chemical composition of nectar influences floral choice by pollinators. In this talk, I review what we know about the bee's sense of taste. My lab has discovered that bees have specialized mechanisms for the perception and encoding of sugar molecules in food which make them capable of discriminating among sugar compounds. We have also found that bee species have specific preferences for nectar cocktails, indicating that bees may exert selective pressure on plants to produce nectar suited to their metabolic needs.

## S.127.2 Geographic variation of nectar properties in generalist plants: pollinated-mediated selection or adaptive wandering?

Marcin Zych¹, Katarzyna Roguz¹

1 Botanic Garden, University of Warsaw, Warsaw, Poland

Nectar is one of the important floral characters that shape plant-pollinator interactions. Some nectar properties,

e.g. sugar profiles, seem relatively conservative across different populations of the same species, whereas there is reported considerable variation in others. For instance, in some members of the generalist family Apiaceae nectar of individual plants from certain populations includes amino acids (AAs) that are absent in representatives of other populations. In species of *Rhododendron* producing toxic nectar such variation was reported for toxic grayanotoxine I. Such divergence may be correlated with habitat properties but also may reflect pollinator-mediated selection exerted by local insect communities. The paper discusses plausible evolutionary scenarios that arise from the spatial variation of nectar characters in plants, with the special focus on generalist pollination systems.

## S.127.3 Nectar thieves are costlier than robbers to animal-pollinated plants

Laura Leal¹,², Matthew Koski², Rebecca Irwin³, Judith Bronstein⁴

1 Universidade Federal de São Paulo, Diadema, São Paulo, Brazil. Contact: laura.leal@unifesp.br. 2 Clemson University, Clemson, South Carolina, USA. 3 North Carolina State University, Raleigh, North Carolina, USA. 4 University of Arizona, Tucson, Arizona, USA

Mutualistic interactions are biological markets where different species exchange commodities to mutual benefit. Because of that, mutualisms are susceptible to exploitation, with individuals taking without reciprocating. While models predict fitness costs inflicted by exploiters, empirical knowledge is limited, highlighting the need for broader quantification. Pollination serves as a case study since animal-pollinated flowers are commonly exploited by larcenists—non-pollinating animals consuming flower rewards without reciprocating. The impact of larcenists on plant reproduction varies widely, indicating differing costs and factors influencing them. In the context of nectar flowers, the behavior of larcenists, classified as robbers or thieves, plays a crucial role. Robbers damage flowers while extracting nectar, po-
<figure><img src="image_298.png" /></figure>

tentially affecting flower attractiveness and female re-productive function. Thieves, though not damaging, can disrupt flower visitation patterns. Plant mating systems further mediate these costs, with self-incompatible or strongly selfing plants experiencing greater impacts. The study aims to fill gaps in understanding by employ-ing a meta-analytical approach, focusing on prima-ry nectar robbing and theft and considering different datasets capturing various pollination process steps. We found that both robbers and thieves negatively in-fluence flower visitation patterns, but only thieves im-pact nectar quality and availability. Surprisingly, robbers have neutral effects on nectar traits and even improve male reproductive success. On the contrary, thieves consistently reduce male and female reproductive performance. Importantly, these effects remain consis-tent across plant mating systems, challenging previous generalizations. The study provides a comprehensive evaluation of larcenists' costs on animal-pollinated plants, revealing that nectar theft is more detrimental than robbing, regardless of the plant's mating system. This pioneering evidence enhances our understanding of the intricate dynamics of mutualistic interactions in ecological and evolutionary contexts.

### S.127.4 Nectaries changes across the flower lifetime ensure multiple ecological interactions

Hannelise Balduino¹, Elza Guimarães¹, Silvia Machado¹

1 São Paulo State University, Botucatu, Brazil

Nectar is an essential trophic resource for many ani-mals, playing an important role in plant-animal interac-tions. Amphilophium mansoanum (DC.) LG. Lohmann (Bignoniaceae) is a liana with extranuptial nectaries (ENN), at the calyces' margins, and nuptial annular nectariferous disk (NN) below the ovary, at the bottom of the floral tubes. The energetic content, the chemical composition, and the dynamics of nectar from both nectary types are different. These traits seem to be un-der selective pressure by the interacting animals since ENN is visited by ants, wasps, flies, and cockroaches, and the latter mainly by large-sized pollinating bees. The aim of this study was to understand the functioning of these nectaries based on ultrastructural evidences. The ENN released nectar since the bud stage, while the NN released nectar only during the anthesis. These pat-terns matched the changes in the cell's machinery: For ENN, in the young bud stage, amyloplasts were scarce

and since the pre-anthesis bud, the starch grains pre-sented degradation signs and became almost absent, while cytoplasmic oil bodies, mitochondria, endoplas-mic reticulum, and leucoplasts became more evident. For the NN, in the young bud stage amyloplasts were also scarce, but there were lipophilic bodies in plastids and vacuoles. However, at pre-anthesis stage there was a massive increase in amyloplasts abundance, Golgi bodies (fewer than in ENN), mitochondria and plastids with different contents. At anthesis, we ob-served an increase in cytoplasmic oil bodies, protein in the cytosol and starch grain degradation. These ul-trastructural changes in ENN, evince a transition from hydrophilic to lipophilic secretion. In NN, in contrast, the population and morphology of organelles were consis-tent with hydrophilic secretion during the whole flower's lifetime. These differences in nectaries functioning may be linked to nectar features that attract specific animal groups, acting on protection and pollination mutualism throughout the flower's lifetime.

### S.127.5 Nectariferous and nectarless Prosthechea Knowles & Westc. (Laeliinae; Orchidaceae)

Małgorzata Stpiczyńska¹, Kevin L. Davies², Emerson R. Pansarin³

1 Faculty of Biology, University of Warsaw, Botanic Garden, Warszawa, Poland. 2 School of Earth and Environmental Sciences, Cardiff University, Main Building, Cardiff, UK. 3 Department of Biology, FFCL-RP, University of São Paulo, São Paulo, Brazil

In the flowers of Prosthechea Knowles & Westc. (Laeliinae; Orchidaceae), there is an internal spur or reser-voir formed by the fusion of the labellum base and the column. This structure acts as a nectary in nectar-se-creting species. However, not all species in this genus produce nectar. In order to determine the structural adaptations for secretory activity or any constraints leading to the cessation of nectar production, the in-ner floral spur (cuniculus) of selected nectar-secret-ing and nectarless flowers was examined at the an-atomical and cellular level. Our results indicate that the presence of nectar in the studied species is unre-lated to the structure and character of the epidermis that covers inside of the spur, e.g. occurrence of the trichomes or papillae, and cuticle surface. Additional-ly, the ultrastructure of epidermal and subepidermal cells, distribution of plasmodesmata and the occur-rence of reserve materials are similar in both types of
flowers. Nectar, if present, becomes visible during the late bud stage, one day prior to flower opening, and is solely accessible during the first day of anthesis. It is then likely reabsorbed. Limited amount of accessible nectar in the flowers of *Prosthechea* can represent a mechanism for pollinator manipulation, enabling plants to decrease the energetic costs of the interaction while still ensuring consistent pollinator visitation.

## S.127.6 The evolution of nectarary anatomy: structural and developmental modifications associated to pollinator switch in Malvaceae

Cristina R. Marinho¹, Lluís García-Mir¹,
Lucía del Castillo¹, Elisa Román¹, Magui
Olangua², Olga Fernández-Palacios²,
Rosa Febles², Yannick M. Staedler³, Jürg
Schönenberger³, Javier Fuertes-Aguilar¹

1 Real Jardín Botánico de Madrid (CSIC), Spain 2 Jardín Botánico Canario Viera y Clavijo (UA CSIC), Spain. 3 Department of Botany and Biodiversity Research, University of Vienna, Austria

The presence of floral nectaries (FN) composed of a carpet of multicellular secretory trichomes, is one of the most conspicuous features of Malvaceae. Although their location within the flowers is virtually constant in the Malvoideae subfamily, they display considerable morphological differences associated

with specific pollination syndromes. Here, we aimed at understanding the evolution of the anatomy underlying the pollinator-mediated nectar secretion comparing the FN structure and development between three phylogenetically-related Malva species pollinated by insects (Malva wigandii, Malva canariensis) or birds (Navaea phoenicea). Buds and flowers were collected along seven developmental stages, from which FNs were dissected and processed for optical microscopy, SEM and CT-scan. Morphological parameters were digitally measured using ImageJ and analyzed with RStudio suite. We found changes in growth and differentiation patterns, anatomy, and starch metabolism relevant to nectar secretion. 1) Trichome inception developed synchronically among the five sepals in all species, but monocentrically in M. canariensis and M. wigandii, and polycentrically in N. phoenicea. 2) N. phoenicea FN takes longer to develop than other species, which may be related to its greater structural complexity. As a result, bird-pollinated N. phoenicea FN are much larger and show a nectariferous parenchyma that is more compact, thicker, and with abundant phloem vascularization. On the contrary, entomophilous species show smaller nectaries, with a parenchyma with intercellular spaces and few or no phloem terminations. 3) Trichomes have a similar shape, but they are shorter and tightly packed in ornitophilous N. phoenicea, which also accumulated more starch in the parenchyma over time than the other species. This study provides evidence for the intimate relationship between FN morphology and nectar production, showcasing the pivotal role of FN size and structure in determining nectar output across distinct pollination strategies.

## S.128 PLANTS IN FOSSIL LAGERSTÄTTEN

### S.128.1 The Umm Irna flora, a latest Paleozoic melting pot from the Dead Sea Region, Jordan

Hans Kerp¹, Benjamin Bomfleur¹

1 Institute of Geology and Palaeontology, University Münster, German

With some 60 species the Umm Irna flora from the Dead Sea region in Jordan ranks among the most late

Permian diverse floras known worldwide. During the latest Permian the present-day Dead Sea region was situated at a paleolatitude of c. 15°S at the western end of the Paleotethys in a monsoonal floodplain setting with meandering rivers, abandoned channels, ephemeral lakes and backswamps of various size. The Umm Irna Formation, exposed on the eastern side of the Dead Sea, is characterized by rapid vertical and lateral facies changes. About a dozen localities have yielded a very diverse flora reflecting the different depositional settings. Not only the high diversity but also the excellent preservation, particularly the cuticles, and the composition make this flora unique. The Umm Irna flora com-
<figure><img src="image_299.png" /></figure>

prises elements of at least three of the four major flo-
ral provinces, Euramerica, Cathaysia and Gondwana. Not only the mixed nature of this flora is of interest, but
also the earliest appearances of extinct plant groups
traditionally regarded as typically Mesozoic, i.e., cory-
stosperms and bennettitaleans, and the still existing
podocarpalean conifers, which became major, some
even dominant constituents of early Mesozoic floras.
Moreover, the Umm Irna flora includes the last appear-
ances of long-ranging Paleozoic groups, including no-
eggerathialean progymnosperms, lepidodendrids, as
well as lyginopteridalean, callistophytalean and gigan-
topteridalean pteridosperms, glosopterids, and wal-
chian conifers. The Umm Irna flora contributes to a bet-
ter understanding of evolution, distribution, extinction
and survival patterns in a crucial period in life history,
shortly before the largest mass extinction on Earth.

S.128.2 The Fossillagerstätte
Kühwiesenkopf of the
Middle Triassic of the
Dolomites (N Italy)

Evelyn Kustatscher¹, Johanna H. A. van Konijnenburg - van Cittert²

1 Naturmuseum Südtirol, Bolzano, Italy; email: Evelyn.Kustatscher@naturmuseum.it. 2 Laboratory of Palaeobotany and Palynology, Utrecht, The Netherlands; email: j.h.a.vankonijnenburg@bio.uu.nl

The Triassic was characterized by greenhouse condi-
tions, but experienced major environmental changes
linked to several short but severe climate shifts. It is in
this climatic scenario that a vegetation turnover took
place with the appearance and radiation of mod-
ern plant families and the rapid diversification of the
gymnosperms, which then became ecologically dom-
inant for most of the Mesozoic era. In 1999 a rich fos-
sil horizon has been discovered within the Anisian ter-
rigenous-carbonatic succession of Kühwiesenkopf /
Monte Prà della Vacca (Dolomites, N-Italy). The fossil
horizon yielded abundant marine fossils, such as bra-
chiopods, bivalves, fishes, and ammonoids, but yield-
ed also abundant terrestrial plant fossils and a small
reptile skeleton called *Megachirella wachtleri* Renesto
et Posenato, 2002, which represents the oldest known
squamate. The main fossiliferous level represents a
rapid burial event caused by submarine flows with-
in a marine basin, triggered by heavy storm events in
the terrestrial domain, but plant debris and scattered
plant remains occur in other layers along the section
as well. The flora is composed of both sterile (shoots,

roots, stems, leaves) and fertile organs (fructifications,
seeds, fertile fern leaves) attributed to at least 35 taxa.
Among those are some of the oldest representants of
the Caytoniales seed ferns as well as conifers that are
more closely related to the Permian forms than to Me-
sozoic ones. This demonstrates how this period of time
was important for the evolution of plants.

S.128.3 A snapshot into
the exceptionally well-
preserved Jurassic
vegetation of the
Mahajanga Basin,
Madagascar

Vivi Vajda¹, Sylvain Razafimandimbison²,
Nicolas Rakotosolofo³, Hery L.
Ranarijaona³

1 Department of Paleobiology, Swedish Museum of Natural History, Svante Arrhenius v. 9, Stockholm, Sweden. 2 Department of Botany, Swedish Museum of Natural History, Svante Arrhenius v. 7, Stockholm, Sweden. 3 École Doctorale sur les Ecosystèmes Naturels, Faculté des Sciences, Université de Mahajanga, Mahajanga, Madagascar.

Here we present results originating from a collabora-
tive project involving paleontologists, geologists, and
botanists from the Swedish Museum of Natural Histo-
ry and Mahajanga University, Madagascar. Madagas-
car's geological history is central to interpreting the ori-
gins of its flora and fauna as the island separated from
continental Africa in the Jurassic ca. 165 Mya, leaving
Madagascar and its biota to develop in isolation from
other continental landmasses. Paleobotanical studies
are few, most dating back to the 1960s and 1970s by
Appert, who described Upper Jurassic (Oxfordian) as-
semblages from the Manamana massif (southwestern
Madagascar). Dipteridaceous ferns, conifers belonging
to the genera *Brachyphyllum*, *Elatacladus*, *Cupressino-
cladus*, *Cyparissidium*, and *Araucarites*, as well as mi-
nor amounts of "seed ferns", make up the majority of
these impression floras. Our palynological results from
two outcrop sections within the Mahajanga Basin, re-
veal well-preserved palynological assemblages of en-
tirely continental aspect. While the older assemblage
is highly dominated by fern- and lycophyte spores,
including *Cyathidites*, *Baculatisporites*, *Dictyophyllid-
ites*, *Ceratosporites* and *Retritiletes*, the younger as-
semblage is dominated by gymnosperm pollen, which
constitutes c. 70%. Within the gymnosperms *Classopol-
<figure><img src="image_300.png" /></figure>

lis, Araucariacites, Perinopollenites, and Calliasporites are the most abundant spore genera. Based on these taxa, we assign the studied assemblage to the Early Jurassic (Pliensbachian–early Toarcian) *Corollina torosa* Zone of Helby et al. (1987). The floras are very similar to the coeval floras of, e.g., Sweden and Australia, revealing a highly homogenous global vegetation pattern at the time growing in a stable ecosystem for many million years, as similar floras are present in the Late Jurassic, as outlined by Appert.

## S.128.4 Paleoecophysiology of Early Cretaceous Crato Flora (Brazil)

Gobo, William Vieira¹, Fradret, Alice²;
Nascimento Jr., Daniel Rodrigues³, Silva
Filho, Wellington Ferreira³, Iannuzzi,
Roberto¹; Kunzmann, Lutz⁴, Coiffard

1 Universidade Federal do Rio Grande do Sul, Porto Alegre, Brazil. 2 ENS de Lyon, Lyon, France. 3 Universidade Federal do Ceará, Fortaleza, Brazil. 4 Senckenberg Naturhistorische Sammlungen, Dresden, Germany. 5 Freie Universität, Berlin, Germany

The Crato Formation of the Araripe Basin, northeast of Brazil is one of the best *Fossil Konservat Lagerstätte* to access Early Cretaceous paleoecosystems in equatorial regions of Gondwana. There, the apparent dominance of gymnosperms splits attention with the emerging group of angiosperms. Despite intensive geological and taxonomic studies made in this area, approaches to elucidate the paleoecophysiological aspects of fossil plants are still rare. Interestingly, the gnetaleans, a gymnosperm lineage, which convergently evolved similar features with many angiosperms, might have occurred the same environments together with some early angiosperms. Until now, it is still lacking clear evidence showing the reason for intense diversification and radiation of both angiosperms and gnetaleans in the early to middle Cretaceous. Both plant lineages may be functionally similar and then subjects to similar selective pressures. One of the big questions is what caused the subsequent success of angiosperms and the coeval decline of gnetaleans. Did gnetalean taxa already differ, in the Crato flora, from angiosperms in terms of ecophysiological adaptations and tolerance ranges? Could this have influenced on the subsequent success of the angiosperms? In addition, does the ecological role of these plants differ significantly? To address these questions, 35 specimens of 17 angiosperm taxa and 70 specimens of 20 gnetalean taxa were prelimi-

narily selected to analyze their paleoecophysiological leaf traits. The functional types of fossils and living taxa were included in our analysis, as well as the type of fos- sil association according to the mode of transport. The leaf energy balance model was used to infer the leaf temperature and transpiration from measurements of vein density and leaf width. As a result, the ecological requirements of fossil taxa of angiosperms and gne- talean gymnosperms were estimated and compared with those of related living plants in different habitats.

## S.128.5 How does fossilisation change the anatomy of wood: awareness and consequences?

Ronny Roessler¹,²

1 Museum für Naturkunde Chemnitz, Chemnitz, Germany, roessler@naturkunde-chemnitz.de. 2 TU Bergakademie Freiberg, Department of Palaeontology, Geological Institute, Freiberg, Germany

Fossil wood belongs to the most frequent and insightful records of ancient terrestrial life. Although cellular anatomical preservation often provides excellent detail, various alterations of organic matter occur during fossilisation and modify tissue properties. Due to manifold taphonomic pathways, plant tissues experience multiple changes, which may cause a tissue-volume reduction, including desiccation, microbial decay, charcoalification, coalification and mineralisation. A few case studies concerning different geological ages, preser- vational backgrounds and palaeogeographic occurrences offer evidence of widely distributed but vastly underestimated shrinkage phenomena in fossil woods. Samples from several Paleozoic to Cenozoic fossil la- gerstätten representing diverse geologic settings are analysed to comprehend putative shrinkage reasons. As a result, volume reduction is more widespread in petrified wood than commonly thought and not easy to recognise. Unexpectedly, tissue contraction varies even in wood from the same fossil site, the same strata and reflecting the same taphonomy. Accordingly, quantita- tive data on cellular anatomy gained as a routine may be misleading, and cell size and shape modifications affect morphometric purposes, like the interpretation and identification of fossil species. Differences in pres-ervation demonstrate the profound effects of microbial degradation, such as lack of tracheid secondary walls and decayed tissue constituents. Differential shrinkage of silicified wood and stromatolite encrustation help
<figure><img src="image_301.png" /></figure>

quantify shrinkage by providing graphically measurable, one- and two-dimensional parameters to assess percental volume loss during fossilisation. This study reveals essential shrinkage-related alterations independent of palaeoenvironment, plant material and host rock. The results demonstrate that shrinkage must be considered when comparing fossil and modern wood morphometric data.

**S.128.6 The hyperdiverse conifer flora of the Baltic amber forest and its palaeoecological significance**

Lutz Kunzmann¹, Alexander R. Schmidt², Eva-Maria Sadowski³

1 Senckenberg Naturhistorische Sammlungen Dresden, Dresden, Germany. 2 Georg August University of Göttingen, Göttingen, Germany. 3 Museum für Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany

Late Eocene Baltic amber constitutes the largest amber deposit on Earth and one of the most important Cenozoic fossil lagerstätten worldwide, however, the picture of vegetation and habitat diversity of its source area has been very fragmentary. Notions concerning the Baltic amber flora were mainly based on palaeobotanical data published in the 19th century and the early 20th century, and conclusions concerning habitat types and

palaeoclimate were predominantly drawn from the ecology of closest living extant relatives of fossil animals from Baltic amber. Here, we summarize our revision of historic monographic treatments and investigation of conifer inclusions from historic and newly discovered amber specimens, and show that the conifer flora of the source area of Baltic amber was hyperdiverse. Our study revealed five families, 17 genera and 26 fossil-species of conifers, including eight genera and 14 fossil-species of Cupressaceae (Calocedrus, aff. Glyptostrobus, Cryptomeria, Cupressinanthus, Cupressinocladus, Quasisequoia, Sequoia, Taxodium); six genera and nine fossil-species of Pinaceae (Abies, Cathaya, Nothotsuga, Pinus, Pseudolarix, aff. Tsuga), and one fossil-species each of the families Geinitziaceae (Cupressospermum), Cephalotaxaceae (Cephalotaxus), and Sciadopityaceae (Sciadopitys). Our data show that the Baltic amber conifer flora is unique and clearly distinct from both central European paratropical or subtropical floras and polar cool-temperate floras of the Eocene. Instead, assemblages from Neogene coastal lowland vegetation of mid-latitudinal Europe exhibit a similar conifer composition. Evaluation of the autecology of individual fossil-species unraveled a heterogeneous vegetation with a mosaic of forests and open habitats in alluvial plains, coastal swamps and mesic hinterland. Our data challenge previous notions that Baltic amber derives from tropical or 'subtropical' forests. Compilation of all currently available botanical data implicates a warm-temperate and humid palaeoclimate in the source area.

**S.129 SYSTEMATICS, FLORISTICS, AND CONSERVATION: FACILITATING DATA INTEGRATION TO PROMOTE SOUND SCIENCE.**
**SESSION 1**

**S.129.1 The intersection of taxonomy and conservation in the United States and Canada**

Wesley M. Knapp¹

1 NatureServe, Arlington, Virginia, USA

NatureServe is a conservation non-profit based out of Arlington, VA, USA that has a foundational role in North American conservation. NatureServe works with 60+ Member Programs (usually but not exclusively referred to as Natural Heritage Programs) to answer the three questions of conservation: What species exist, where are these species found, and how are these species doing?
<figure><img src="image_302.png" /></figure>

NatureServe is a Red List partner but maintains a distinct ranking methodology that has been used to assess over 100,000 plants, animals, fungi, and natural communities. The NatureServe Network maintains a robust geospatial database that includes over 1,000,000 on-the-ground locations of rare species. NatureServe partners with numerous other organizations to maintain taxonomic currency (including but not limited to the Flora of North America committee, USDA-Plants, and Missouri Botanical Garden's Tropicos Database). NatureServe's ranking system of G-, N-, and S- ranks are integral to conservation as these ranks are a valuable prioritization complement to legal status designations assigned by government agencies such as the U.S. Fish and Wildlife Service and the National Marine Fisheries Service in administering the U.S. Endangered Species Act (ESA), and the Canadian Ministry of Environment and Climate Change in administering the Species At Risk Act (SARA). Integrating the best available taxonomy presents a pressing conservation challenge. The intersection of taxonomy and conservation is often misunderstood, difficult to understand, and confusing for the non-expert. This can lead to the mislocation of precious conservation resources, inappropriate decisions about what species should be conserved, and what natural communities should be restored or managed. Great conservation liabilities exist when there are taxonomic disagreements and misalignments. In this presentation, I give examples of how disagreements or misalignments have caused negative conservation impacts, discuss how NatureServe presents a multi-tiered taxonomy, and propose possible solutions to the botanical community.

### S.129.2 The Flora of North America: a hard copy and online continental flora

Geoffrey A. Levin¹

1 Research and Collections, Canadian Museum of Nature, Ottawa, Canada

The Flora of North America North of Mexico (FNA) is a 30-volume work that includes taxonomic treatments for all the native and naturalized plants growing in the continental United States, Canada, Greenland, and St. Pierre and Miquelon. Treatments include accepted names and important synonyms, detailed morphological descriptions, dichotomous keys, phenology and habitat data, text- and map-based distributions, and line drawings of about 20% of the species. The project is a truly international effort involving a collaboration among about 1,000 artists, reviewers, and editors from

throughout the world. FNA is a vital tool for those concerned with conserving the continent's flora for several reasons. For many groups, FNA provides the only comprehensive, modern North American treatments available anywhere. Although continental in scope, the treatments are reviewed by a team of about 75 botanists with expertise in their state or provincial floras, ensuring that important local variation and distributions are accurately reflected. The combination of keys and detailed technical descriptions improves plant identification. Furthermore, because taxonomic opinions vary, both among sources and over time, the included synonymy affords the opportunity for nomenclatural and taxonomic cross-walks. Because the treatments are made available online soon after they are published, they not only are widely accessible, but the finely parsed data are also made available in formats that improve data sharing and cross-linking. Efforts currently are underway to improve coordination among FNA, NatureServe, and the USDA-NRCS National Plant Data Team, organizations that play major roles in providing conservation data for North American plants. Looking to the future, we will be working to develop tools for moderated and versioned authoring and editing of treatments at FNA's website (floranthorhamerica.org), allowing FNA to remain up-to-date by continuously incorporating results of new research.

### S.129.3 Reinventing the "flora" as a 21st century tool for science and conservation

Alan S. Weakley¹, Michael T. Lee¹, Scott G. Ward¹, J. Christopher Ludwig², Katie Gibson³

1 North Carolina Botanical Garden (Herbarium NCU), University of North Carolina, Chapel Hill, USA.
2 Seedbox Consulting, Maidens, USA.
3 High Country Apps, Bozeman, USA

The flora ("a treatise on or list of the plant life of a particular region") has been a centuries-old basic tool in understanding, studying, and using the plant biodiversity of a particular geographic area. The contents and format of floras have evolved through time, but typically include a list organized by taxonomy of the species native or naturalized in the region covered, with descriptions of the morphology, habitats, phenology, and synonyms of each, including keys for their identification, and sometimes including other information (illustrations, maps, synonyms, taxonomic discussion,
<figure><img src="image_303.png" /></figure>

chromosome numbers, etc.). Floras are difficult to develop, expensive to publish and purchase, often written for a limited and technical audience, and infrequently revised, hampering the utility and currency of this necessary resource. The Southeastern United States Flora project covers a biodiverse area of in southeastern North America, with about 11,000 vascular plant species. We are using modern technology to reinvent the flora as a dynamic, digital resource for modern biodiversity science and conservation by developing and maintaining a database, FloraManager. FloraManager supports easy revision by multiple, production of products in diverse platforms (traditional paper, web app, installable mobile app), and customization of those products. Digital flora products enable us to have nearly unlimited imagery, taxonomic concept-mapping to over 7000 other taxonomic sources, insertion of diagnostic illustrations as "popups" into keys, development of multiple-access graphic keys, geographic autosubsetting of keys, autocreation of "on-the-fly" dichotomous keys (decision trees) for any set of species in the flora, and inclusion of species-specific conservation information (imperilment status, wetland indicator status, heliophy, ecological conservatism, etc.). Goals include engaging a broader audience (general public, land managers, environmental consultants, etc.) for information on the plants of the region, using modern technology to make plant identification easier, and bringing current taxonomy to bear on locating and conserving plant biodiversity.

### S.129.4 Taxonomic discovery from a regional perspective: current approaches and conservation implications

Derick B. Poindexter¹, Alan S. Weakley¹

1 North Carolina Botanical Garden (Herbarium NCU), University of North Carolina, Chapel Hill, USA

Despite its long history of botanical exploration, the Southeastern United States still has much novel biodiversity to yield, as evidenced by the large number of new taxa described over the past 50 years – nearly 10% of the recognized native flora. Contemporary molecular tools, in conjunction with traditional systematic techniques, have made this continued advancement possible, particularly concerning difficult species complexes. Furthermore, many of these ongoing studies focus on long-recognized problematic groups, historically neglected due to the perceived need for extensive time and financial investments. As

these rate-limiting factors lessen (e.g. NGS sequencing costs decrease), we are encouraged to pursue these "sidelined" studies with a collective evidence approach. This effort utilizes multiple data sources (molecular, cytogenetic, ecological, biogeographic, etc.) to help guide our ultimate taxonomic interpretations. While this approach is not conceptually novel, the power of current technologies to help aid this process is, and an approach that regionally targets known research needs is atypical in a world in which research funding tends to drive research by plant family. Our lab is actively pursuing resolution in a number of clades within taxonomic disparate genera including *Allium* (Alliaceae), *Astragalus* (Fabaceae), *Blephilia* (Lamiaceae), *Carex* (Cyperaceae), *Eryngium* (Apiaceae), *Helianthus*, *Marshallia*, and *Packera* (Asteraceae), *Micranthes* (Saxifragaceae), and *Thalictrum* (Ranunculaceae). We use select examples from these groups to illustrate the case-by-case nuances encountered during our systematic assessments, as well as discuss the regional and global implications of our findings.

### S.129.5 Collaboration is the key to documenting and conserving biodiversity: A case study on Colorado's Front Range

Jennifer R. Ackerfield¹, Mit E. McGlaughlin²

1 Jennifer R. Ackerfield, Denver Botanic Gardens, Denver, CO, U.S.A. 2 Mit E. McGlaughlin, University of Northern Colorado, Greeley, CO, U.S.A.

Conservation management requires time, personnel, and funding to accomplish effectively. However, these resources are often in short supply. It is therefore important for government agencies to efficiently spend their time, targeting the most vulnerable rare plant species and communities for protection. Additionally, which species agencies target for conservation relies on a foundation of accurate taxonomic classification. One taxon of conservation concern on the Front Range of Colorado is referred to as *Physaria* X1 by the Colorado Natural Heritage Program (CNHP). *Physaria* X1 is narrowly restricted in range and is hypothesized to be a hybrid between the rare *Physaria bellii* (G2S2) and more widespread *P. vitulifera* (G3S3). Previous research done nearly 30 years ago indicated that *Physaria* X1 may be genetically distinct and deserve recognition as a species. Based on these findings, *Physaria* X1 is tracked as a taxon
of conservation concern, despite formal species recognition. However, designation at the species level would have significant management consequences for several agencies. A collaboration of researchers, government, and non-profit agencies determined if *Physaria* X1 deserved species recognition using population-level sampling and next-generation sequencing. Our results showed conclusively that *Physaria* X1 is not a unique species, but a phenotypically unusual *P. vitulifera*. However, surprisingly, one population of *P. vitulifera* was shown to be quite genetically distinct from other populations. Upon further investigation, we found populations of these *Physaria* to also be morphologically distinct from *P. vitulifera* and uniquely substrate specific. Therefore, we recommend that populations of this *Physaria* be treated as a new species, *Physaria pamsmithiae*, for the CNHP botanist who diligently tracked *Physaria* X1 and advocated for this project. These results highlight the importance of government and non-profit agencies working collaboratively with researchers to provide efficient, evidence-based conservation management strategies. Ultimately, these results also inform our understanding of regional biodiversity.

**S.129.6 Using genomic data to inform taxonomy and conservation of rare plants**

Mitchell McGlaughlin¹, Samantha Naibauer¹

1 University of Northern Colorado, Greeley, Colorado, USA

Accurate taxonomic identification of rare plant species is essential for defining elements of biodiversity and directing conservation funding to protect that diversity. Recent advances in genetic data collection, particularly those utilizing next-generation DNA sequencing technologies, have increased the accuracy and speed of genetic studies, and has lowered the cost to generate robust genomic datasets. Genetic data provides a method to determine if species are distinct from close relatives, genetic structure of populations, and levels of genetic diversity within populations, independent of challenges associated with phenotypic plasticity, which can complicate analyses of plant morphology. However, many of these modern genetic tools remain focused on evolutionary studies and are not directly leveraged to support conservation. In this presentation, We discuss collaborations between the University of Northern Colorado, the U.S. Fish and Wildlife Service, and the Colorado Bureau of Land Management as a model to support the collection of conservation focused genetic data. Supported studies have examined taxonomic uncertainty, defined management units, and increased our understanding species life history. Examples will focus on how genetic data has improved the allocation of conservation resources in Colorado, with studies that determined that *Draba weberi* (Brassicaceae) is not a distinct species, that endangered *Sclerocactus glaucus* (Cactaceae) should be recognized as two species, and that *Erythranthe gemmipara* (Phrymaceae) is entirely clonal with a very uneven distribution of genetic diversity across the species range. Collectively, these results illustrate how agencies and researchers can collaborate to ensure that the taxonomy of plant species of concern is accurate and that conservation activities are targeted to species most in need.

**S.130 MALVACEAE: PROGRESS & PROSPECTS IN EVOLUTIONARY RESEARCH IN THE COTTON & CACAO FAMILY**

**S.130.1 What have we learned from the Malvaceae fossil record?**

Cédric Del Rio¹

1CR2P, Centre de Recherche en Paléontologie-Paris, MNHN, Sorbonne Université, CNRS, Paris, France

The Malvaceae is a family of flowering plants that consists of 244 genera and 4,225 species. This family is found on all vegetated continents and exhibits substantial morphological and habitat diversity. The fossil record has been abundant since the Cretaceous period, but many fossils, especially pollen and leaf remains, are ambiguous. As a result, some subfamilies, such as Malvoideae, may have a challenging-to-understand fossil record. Nevertheless, Malvaceae fossils
<figure><img src="image_304.png" /></figure>

are distinguished by the variety of questions they
have helped answer throughout the history of paleo-
botany. What have we learned from the fossil record
of Malvaceae? This presentation highlights three fo-
cus of study that have been developed: (i) Origins:
Although partial, Malvaceae fossils have helped clar-
ify the temporal origin of the family and the diversi-
fication of its subfamilies, particularly by supporting
molecular phylogenies. The study of new fossils has
constantly made it possible to test the ages pro-
posed by molecular analysis and has often extended
the previously determined ages. (ii) Biogeography:
Malvaceae serves as a model for the biogeography
of subtropical floras. For instance, the genus *Craigia*,
a relict group from East Asia, has an extensive fossil
record that has helped elucidate the history of floras
at various geographic levels in the Northern Hemi-
sphere, especially exchanges between North Amer-
ica, Asia, and Europe. (iii) Morphological evolution:
Fossils exhibit forms that are little or not documented
in the present diversity, allowing us to understand the
evolution of certain organs over time. Examples in-
clude the inflorescence bracts of the genus *Tilia* and
the evolution of wood in the Sterculioideae subfamily,
which have revealed morphological changes over
time. From these three axes, it follows that the study
of Malvaceae fossils addresses many key questions
related to plant evolution. New fossils, for example,
those from less explored regions, could refine both
paleobotanical and neobotanical studies.

**S.130.2 Reconstructing the**
**phylogeny of Malvaceae**

Rebeca Hernández-Gutiérrez¹

1 Department of Evolution, Ecology, and Organismal Biology, University of California, Riverside, USA

The Malvaceae family outstands for its diversity in
species richness, morphology, growth form, and geo-
graphic distribution. Unraveling the drivers behind
such a high variation demands an integrative ap-
proach that leverages multiple data sources to yield
a comprehensive picture of the evolution of this im-
portant family. It is known that numerous events of in-
trogression occurred in the deep past of Malvaceae,
which not only explains the pervasive phylogenetic
discordance among genes trees, but also explains flo-
ral evolution in some subclades. However, the broader
implications of introgression on floral evolution remain
largely unexplored. This study aimed to quantify the ex-
tent of introgression among genera within Malvaceae
and to assess its role in shaping floral morphologies.

By examining a comprehensive molecular dataset
of hundreds of single-copy nuclear genes across all
nine subfamilies, we employed a maximum likelihood
framework to ascertain introgression levels and direc-
tion. Floral evolution was examined applying a model
that explicitly tests morphological evolution through
introgression. This study illuminates the complex inter-
play between introgression and floral evolution, pro-
viding novel insights into the evolutionary processes
that have shaped the phylogenetic relationships and
diversity in Malvaceae.

S.130.3 Hibiscus sect.
Lilibiscus: Expanding
perspectives on the tempo
and mode of extreme long-
distance dispersal and
rapid radiations on

Brock Mashburn¹,², Margaret M. Hanes³,
Lex A. J. Thomson⁴, Kenneth M. Olsen¹,
Christine E. Edwards²

1 Washington University in St. Louis, St. Louis, MO, USA. 2 Missouri Botanical Garden, St. Louis, MO, USA. 3 Eastern Michigan University, Ypsilanti, MI, USA. 4 Australian Centre for Pacific Islands Research, Queensland, Australia

The last three decades have seen an explosion
of studies showing long-distance dispersal to
be a major force in determining the patterns of
species distributions around the globe. The in-
creasing prevalence of molecular phylogenies
with widespread species sampling, coupled with
improved dating methods, has revealed that dis-
persal and subsequent adaptive radiations are
common and occur more rapidly than previ-
ously thought. Recent studies in the genus *Hibis-
cus* (Malvaceae) have demonstrated repeated
dispersal events out of Madagascar alone, in-
dicating that the island is not only a hotspot of
biodiversity but also a source of colonizers to nu-
merous continents. Perhaps most striking are the
species in *Hibiscus* section *Lilibiscus*, a group that
arose in Madagascar within the last ~2 million
years. Members of *Lilibiscus* are largely endemic
to oceanic islands, with 19 of the 30 species (63%)
occurring on three volcanic archipelagos (Mas-
carene, Fijian, Hawaiian) spread across the globe.
A parsimonious explanation for this distribution
might be a single dispersal event out of Mada-
<figure><img src="image_305.png" /></figure>

gascar to the Mascarenes, from there to Fiji, and finally to Hawaii. We tested this hypothesis using a 2bRAD next-generation sequencing approach with over 700 samples and produced a phylogeny that includes every species in sect. *Lilibiscus* and closely related outgroups. The resulting species relationships revealed an unexpected colonization pattern, with at least three independent dispersal events out of Madagascar and two-way dispersal between islands in the Pacific and Indian Oceans. This suggests that we may still be underestimating the ability of species to regularly disperse across incredibly long distances and that species disjunctions could be much younger than expected. In addition, we uncovered a surprising biogeographical connection between volcanic archipelagos in the Pacific and Indian Oceans that requires further investigation.

## S.130.4 Towards an updated classification of Malvales and Malvaceae: insights from phylogenomics and morphology

Matheus Colli-Silva¹, Oscar A. Pérez-Escobar¹, Carlos D.M. Ferreira², Maria T.R. Costa², Samuele Gerace², Thales Silva-Coutinho³, Vania Nobuko Yoshikawa⁴,⁵, Higor Antonio-Domingues¹, Rebeca Hernández-Gutiérrez⁶, Martin Cheek¹, Mark W. Chase¹, Michael F. Fay¹, William S. Alverson⁷, Massimo Bovini², Laurence J. Dorr⁸, William J. Baker¹, Alexandre Antonelli¹,⁹,¹⁰,¹¹

¹ Royal Botanic Gardens, Kew, Richmond, Surrey, UK.

2 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Diretoria de Pesquisas, Rio de Janeiro, Rio de Janeiro, Brazil.
3 Instituto de Biociências, Universidade Federal do Mato Grosso, Cuiabá, Mato Grosso, Brazil.
4 Laboratório de Sistemática Vegetal, Núcleo de Ciências Ambientais, Programa de Pós-Graduação em Biotecnologia, Universidade de Mogi das Cruzes, Mogi das Cruzes, São Paulo, Brazil.
5 Departament of Botany, Institute of Biosciences, University of São Paulo, São Paulo, Brazil (Present affiliation).
6 Department of Evolution, Ecology, and Organismal Biology, University of California, Riverside, California, USA.
7 WIS Herbarium, University of Wisconsin-Madison, Madison, Wisconsin, USA.
8 Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington DC, USA.

9 Gothenburg Global Biodiversity Centre, Gothenburg, Sweden.
10 Wuhan Botanical Garden, Chinese Academy of Sciences, Wuhan, China.
11 Department of Biology, University of Oxford, Oxford, UK.

The Malvales, a diverse order of flowering plants spanning various global biomes, boasts significant species diversity and morphological variability. Despite its ecological and economic importance, Malvales classification has been largely neglected since the late 1990s. This study addresses this gap by revisiting Malvales classification, emphasizing its largest family Malvaceae. By integrating molecular and morphological datasets, our primary goal is to map key traits for classification onto an extensive phylogeny and reassess taxonomic boundaries as necessary. This investigation enhances our evolutionary understanding of Malvales, bringing providing valuable insights for a more contemporary taxonomy. We generated a molecular dataset based on 196 nuclear genes for 188 genera within Malvales, including genera from all recognized families and subfamilies, almost all tribes, and all subtribes. Phylogenetic trees were inferred using maximum-likelihood and a coalescence approach to reconcile conflicts. A morphological matrix with 52 characters relevant for to suprageneric classification was compiled to identify diagnostic features within specific clades. Character state distributions were mapped onto the final phylogenetic topology to aid the search of for diagnostic features for of suprageneric taxa. Phylogenetic relationships and monophyly for most groups aligned with prior research. Significant variations in diagnostic morphological traits were observed across clades, including growth forms, leaf structures, pollen types, flower floral attributes, and fruit/seeds. Incongruities in relationships among some Malvales families and within well-supported Malvaceae clades were attributed to discrepancies observed in gene trees. For Malvaceae, uncertainties in basal Malvoideae were resolved. Incongruities among nuclear loci, while not impacting clade definitions, suggest caution in interpreting the current Malvaceae classification. We propose maintaining Malvaceae sensu lato and introducing two new subfamilies (Matisioideae and Durionoideae) for several tropical genera. Pollen characteristics define Malvoideae as a monophyletic subfamily. Four new tribes within Malvoideae improve taxonomic placement for genera previously of uncertain classification, aligning with updated findings.
<figure><img src="image_306.png" /></figure>

### S.130.5 A new phylogeny and classification for tribe Hibisceae (Malvaceae)

Margaret M. Hanes¹,¹³, Orland J. Blanchard Jr.², Janice Valencia-D.³,⁴, Todd McLay⁵,⁶,⁷, J. Richard Abbott⁸, Stuart F. McDaniel⁹, Russell L. Barrett¹⁰,¹¹, Sarah Mathews⁶,¹², Kurt M. Neubig³

1 Eastern Michigan University, Ypsilanti, Michigan, USA. 2 Florida Museum of Natural History, University of Florida, Gainesville, Florida, USA. 3 Southern Illinois University, Carbondale, Illinois, USA. 4 Universidad Nacional de Colombia, Bogotá D.C., Colombia. 5 Royal Botanic Gardens Victoria, Melbourne, Australia. 6 Centre for Australian National Biodiversity Research, CSIRO, Canberra, Australia. 7 The University of Melbourne, Melbourne, Australia. 8 University of Arkansas at Monticello, Monticello, Arkansas, USA. 9 University of Florida, Gainesville, Florida, USA. 10 National Herbarium of New South Wales, Botanic Gardens of Sydney, Australia. 11 University of New South Wales Sydney, Australia

Sydney, Australia 12 Louisiana State University, Baton Rouge, Louisiana, USA

Tribe Hibisceae represents a very iconic group of plants with extraordinary horticultural, agricultural and cultural significance. No studies, however, have broadly sampled across the dozens of taxa within the tribe, leading to uncertainty in the relationships among genera. The non-monophyly of the genus *Hibiscus* is infamous and challenging, whereas the monophyly of most other genera in the tribe has yet to be assessed, including the large genus *Pavonia*. We significantly increase taxon sampling and assess monophyly of most currently recognized genera in the tribe. We also include many newly sampled species and sections of *Hibiscus* and *Pavonia*. Our phylogenetic trees demonstrate that *Hibiscus*, as traditionally defined, encompasses at least 20 additional genera. The status of *Pavonia* emerges as comparable in complexity to *Hibiscus*. Our phylogeny offers clarity in the phylogenetic placement of several taxa of uncertain affinity (e.g., *Helicteropsis*, *Hibiscadelphus*, *Jumelleanthus*, and *Wercklea*). We introduce a new classification for the tribe and clarify the boundaries of *Hibiscus* and *Pavonia*.

## S.131 THE ROLE OF AGMATOPLOIDY AND SYMPLOIDY FOR DIVERSIFICATION OF FLOWERING PLANTS WITH HOLOCENTRIC CHROMOSOMES

### S.131.1 Evolutionary significance of holocentric chromosomes

František Zedek¹, Petr Bureš¹

1 Department of Botany and Zoology, Faculty of Science, Masaryk University, Czechia

Holocentric chromosomes, in contrast to the commonly known monocentric ones, exhibit a remarkable tolerance for fragmentation. They have independently evolved multiple times in both plants and animals. Despite their rarity, holocentrics might have significantly influenced the colonization of land half a billion years ago, while also gaining relevance as significant agricultural pests in contemporary times. Furthermore, these chromosomes are increasingly becoming focal

points in studying chromosomal speciation and comprehending the impact of recombination on species' success within dynamic or stable environments.

### S.131.2 The architecture of chromosome fission and fusion in holocentrics

Marcial Escudero¹

1 Department of Plant Biology and Ecology, University of Seville, Seville, Spain

Chromosome evolution is a major driver of diversification in angiosperms. Whereas polyploidy has been investigated at a great deal in angiosperms, dysploidy (chromosome number changes by fission/agmatoploidy and fusion/symploidy without a significant
<figure><img src="image_307.png" /></figure>

icant change in DNA content) has been, in comparison, mostly ignored. The clade of family Cyperaceae, with holocentric chromosomes -kinetochoric activity is distributed along the whole chromosome-, and consequently a large range of chromosome number variation, is an ideal study system to investigate the consequences of chromosome fission and fusion. By fitting models of chromosome evolution in the phylogeny of the Cyperaceae, we have inferred that the rate of dysploidy, and also polyploidy, is not homogeneous across Cyperaceae, instead there are clades where rates of chromosome evolution accelerates or decelerates. Using comparative genomics of closely and distantly related species, we have also observed the pull of the present for chromosome evolution. The same way that the rates of molecular, morphological, fossil and speciation seem to accelerate towards the present in the tree of life, we have also found how the rates of fission and fusion seem to be much faster at microevolutionary scale than at macroevolutionary scale. Comparative genomics suggest that there is a strong genomic constraint that limits where fission and fusion may occur which results in parallel events of fission and fusion across different lineages and leads to underestimate the rates of dysploidy at the macroevolutionary scale. The inferred hotspots of dysploidy are very rich in repeat DNA, that is, the repeat landscape may determine the rates of fission and fusion and their location in the genome. Finally, at microevolutionary scale, we shed light into the crucial question of how new karyotypes become established. New chromosome variants may become fixed in the populations by both, stochastic processes like drift or selection of locally adapted karyotypes.

### S.131.3 Cryptic evolution and diversification of agmatoploid-polyploid Luzula sect. Luzula (Juncaceae) in the Eastern Alps

Valentin Heimer¹,², Jonas Geurden², Andreas Hilpold¹, Mingai Li³, Claudio Varotto³, Peter Schönswetter², Božo Frajman²

1 Eurac Research, Bozen/Bolzano, Italy. 2 University of Innsbruck, Innsbruck, Austria. 3 Fondazione Edmund Mach, S. Michele all'Adige, Italy.

The diversification of flowering plants has been influenced by changes in the number and structure of

chromosomes. While the evolutionary consequences of chromosome duplications (polyploidy) are well documented, much less is known about the effects of chromosome fragmentations (agmatoploidy). *Luzula* (Juncaceae) is a genus with common incidence of both phenomena, leading to a variety of cytotypes. The most variable and taxonomically intricate group within the genus is *Luzula* sect. *Luzula*, of which eight species with six karyotypes, including di-, tetra- and hexaploids, have been reported for the Eastern Alps. However, due to weak morphological differentiation among taxa, their distributions and phylogenetic relationships are insufficiently known. By using an integrated approach employing extensive vegetation surveys, relative genome size estimations, chromosome counts, RAD- and plastome sequencing, we aim at disentangling the evolutionary history and ecological niche segregation within this species complex. Combining a variety of different methods will allow us to reconstruct phylogenetic relationships among species as well as unravel the roles of chromosome duplication and fragmentation for speciation and ecological divergence in *Luzula*. In this talk we present preliminary results of ecological and genomic data. Our findings include common co-occurrence of cytotypes at the same locality and a varying degree of ecological differentiation among taxa. In the (sub)alpine belt, we found tetraploid species to be more common on silicates while diploid *L. exspectata* occurs mostly on calcareous bedrock and *L. sudetica* is restricted to humid habitats.

### S.131.4 Initial insights about mono- and holocentromeres in carnivorous Drosera species

Laura Ávila Robledillo¹, Aaryan Bhatia¹, André Marques¹

1 Max Planck Institute for Plant Breeding Research, Cologne, Germany

Sundews are carnivorous plants that grow in nutrient-poor habitats, such as bogs, swamps, and sand dunes. They are distributed globally, with more than 200 species described. In addition to carnivory, sundews are reported to have great diversity of genomic and chromosomal features. The chromosomes of sundews have been studied for over a century revealing extraordinary variability in chromosome number and ploidy across species. But perhaps, one of the most fascinating features of sundews chromosomes is the fact that several species analyzed within
<figure><img src="image_308.png" /></figure>

the genus *Drosera* lack a visible primary constriction, leading to the assumption that holocentricity is common in the genus. This is very exciting as it points to the occurrence of both monocentric and holocentric species in a single genus. Combining genomics and cytogenetics we are analyzing the genome of *Drosera* species differing in centromere organization. We assembled the genome of 5 selected species from the main clades using a combination of long-read sequencing technologies. In addition we made use of low-coverage Illumina short reads for the characterization of the repetitive fraction of the genome in additional species. Our preliminary data reveals the different genome architecture of mono- and holocentric *Drosera* and suggest that satellite repeats are associated with centromeres in both types. However, satDNAs are rarely conserved among *Drosera* species contrary to other reported holocentric species. Moreover, the development of specific centromere markers allowed us to verify the centromere type of our set of species. In addition our data gives insights into the evolution of the karyotypes of *Drosera* species. Synteny analyses among *Drosera* and its sister clade such as *Nepenthes* showed that events of fusion and fusion are common in the group. This work could provide the bases for the understanding of the genomic changes associated with holocentricity as well as the drivers of this transition.

### S.131.5 Combining chromosome evolution models and phylogenomic data to disentangle polyploidy and single chromosome evolution in *Drosera*

Rebekah Mohn¹,², Rosana Zenil-Ferguson³, Thilo Krueger⁴, Andreas Fleischmann⁵,⁶, Adam Cross⁴,⁷, Ya Yang²

1 Center for Tree Science, Morton Arboretum, Lisle, IL, United States. 2 Department of Plant and Microbial Biology, University of Minnesota-Twin Cities, St. Paul, MN, United States. 3 Department of Biology, University of Kentucky, Lexington, KY, United States. 4 School of Molecular and Life Sciences, Curtin University, Bentley, WA, Australia. 5 Botanische Staatssammlung München (SNSB-BSM), Munich, Germany. 6 GeoBio-Center LMU, Ludwig-Maximilians-University, Munich, Germany. 7 EcoHealth Network, Brookline, MA, United States

*Drosera* L. (Droseraceae, Caryophyllales) has been long thought to exhibit large amounts of chromosome evolution due to both single chromosome number change and polyploidy. We used the BiChrom model to test whether rates of single chromosome number increase and decrease, and chromosome number doubling differed between *D. subg. Ergaleium* and the other subgenera and between self-compatible and self-incompatible lineages. The best model for chromosome evolution among subgenera had equal rates of chromosome number doubling but higher rates of single chromosome number change in *D. subg. Ergaleium* than in the other subgenera. Contrary to expectation, self-incompatible lineages had a significantly higher rate of single chromosome loss than self-compatible lineages. The findings of different rates in different subgenera were further supported by phylogenomic analyses not finding evidence for any polyploid events in the *D. subg. Ergaleium* while finding evidence of polyploidy in the other subgenera. We found no evidence for an association between differences in single chromosome number changes and diploidization after polyploidy or centromere type. This study highlights the complexity of factors influencing rates of chromosome number evolution. Further work should explore whether meiotic and specifically holocentric drive might explain heterogeneity in chromosome number evolution in *Drosera*. This work is in part published in Evolution at https://doi.org/10.1093/evolut/qpad153.

### S.131.6 Detecting shifts in the mode of chromosomal speciation across the cosmopolitan plant lineage *Carex*

Carrie M. Tribble¹, José Ignacio Márquez-Corro²,³, Michael R. May⁴, Andrew L. Hipp⁵, Marcial Escudero⁶, Rosana Zenil-Ferguson⁷

1 School of Life Sciences, University of Hawai'i at Mānoa, Honolulu, USA. 2 Royal Botanic Gardens, Kew. Richmond, Surrey, UK. 3 Department of Molecular Biology and Biochemistry Engineering, Universidad Pablo de Olavide, Sevilla, Spain. 4 Department of Evolution and Ecology, University of California Davis, Davis, USA. 5 Herbarium and Center for Tree Science, The Morton Arboretum, Lisle, USA. 6 Department of Plant Biology and Ecology, University of Sevilla, Sevilla, Spain. 7 Department of Biology, University of Kentucky, Lexington, USA
Recent studies of angiosperm diversification have focused on the role of polyploidy as a driver of diversification. However, we know far less about the effects of single changes in chromosome number—dysploidy, which can mediate lineage diversification by affecting recombination rates, linkage, and/or reproductive isolation. Modeling the effects of dysploidy on diversification is mathematically and computationally challenging because many states and parameters are required to track changes in individual chromosomes, especially in lineages that have high variability in chromosome number. Additionally, we expect the processes of diversification and dysploidy to vary across clades, which requires modeling process variation to disentangle the effects of the observed trait (chromosome number) from the effects of unobserved traits on diversification. In this work, we propose a new state-dependent diversification model of chromosome evolution that

includes numerous character states and explicitly models heterogeneity in the diversification process. Our model includes parameters that functionally link diversification rates to dysploidy rates and differentiate between anagenetic and cladogenetic changes. We apply this model to *Carex* (Cyperaceae), a model lineage for understanding dysploidy and diversification, leveraging chromosome number information and the most recent time-calibrated phylogenetic tree for over 700 species and subspecies. We recover distinct modes of chromosomal speciation across *Carex*. In one mode, dysploidy occurs very frequently and drives faster diversification rates. In the other mode, dysploidy is rare and diversification is driven by other factors, unmeasured in our analysis. This study is the first to demonstrate that dysploidy drives diversification in plants while considering unmeasured factors affecting diversification.

# S.132 BIOGEOGRAPHIC AND EVOLUTIONARY PATTERNS AND PROCESSES OF THE TROPICAL TEMPERATE BIOME TRANSITION

## S.132.1 Taxonomy, biogeography, and sectional boundaries of the tropical blueberries (Vaccinieae, Ericaceae) of Malesia

Maverick N. Tamayo¹, Wendy A. Mustaqim², John D. Horner¹, Matthew C. Hale¹, Peter W. Fritsch³

1 Department of Biology, College of Science and Engineering, Texas Christian University, Fort Worth, USA. 2 Program Studi Biologi, Fakultas Teknik, Universitas Samudra, Langsa, Indonesia. 3 Botanical Research Institute of Texas, Fort Worth, USA

The tribe Vaccinieae (Ericaceae) is a large group of more than 1000 species mostly found in the tropics, and a third of which are found in Malesia. Vaccinium is the largest genus of Vaccinieae in Malesia, comprising ca. 250 species, more than half of which are endemic to the island of New Guinea. Malesian species of Vaccinium are mainly adapted to high-elevation zones, exposed mountain ridges and peaks, montane mossy rain

forests, nutrient poor habitats, and rarely littoral areas. They are often rare in the wild; hence, few studies on its taxonomy have been conducted in Malesia. A revision was included in the account for Ericaceae in the *Flora Malesiana*. However, limited specimen availability and utilization of ambiguous and poorly defined morphological characters hindered resolution of many species complexes. Moreover, sectional boundaries of Malesian Vaccinium are poorly defined, and the morphological characters overlap with those of other sections, thus limiting their utility. Furthermore, a comprehensive phylogeny for the Malesian Vaccinium is lacking. A preliminary phylogenetic analysis of Ericaceae reveals that Vaccinium is non-monophyletic, with the genera *Agapetes* and *Rigiolepis* nested within it. We present recent findings from field and herbarium work helping to clarify the systematics and evolution of Malesian Vaccinieae. A taxonomic revision of Philippine Vaccinium has resulted in the discovery of 11 new species, three new island records, rediscoveries, and clarification of species complexes. In Indonesia, we have discovered a new species and a variety of Vaccinium, including three new species of *Rigiolepis*. We provide assessment of the morphologi-
<figure><img src="image_309.png" /></figure>

cal basis for the sectional boundaries of Malesian Vaccinium in the context of ongoing molecular analysis. The biogeography, distribution, and morphological distinctions among Agapetes, Rißiolepis, and Vaccinium will be discussed.

## S.132.2 The tropical-temperate transition in a World without clear-cut climatic boundaries: evolutionary implications

Medina, N.G.¹,², Coca de la Iglesia, M.¹, Gallego, A.¹, Wen, J.³, Valcárcel, V.¹,²

1. Departamento de Biología, Universidad Autónoma de Madrid, Madrid, Spain. 2. Centro de Investigación en Biodiversidad y Cambio Global, Universidad Autónoma de Madrid, Madrid, Spain. 3. Department of Botany/MRC 166, Smithsonian Institution, Washington, DC, USA.

The so-called latitudinal diversity pattern has often been explained invoking the clear-cut climatic differences between tropical-temperate latitudes. However, this hypothesis assumes that moving from the tropics to the temperate latitudes implies drastic changes in the climatic preferences of the species. Yet the climatic transitions between regions are often gradual and overlooking their importance can deeply affect how we perceive the impact of climate on evolutionary patterns. We examined the nature and distribution of the transitions between climatic regions worldwide and evaluated their impact for inferring the past and present climatic preferences in the Asian Palmates, a highly diverse and climatically complex lineage of Araliaceae. We find consistent geographic patterns in the climatic transitions worldwide. Particularly, we detect a consistent higher extent and spatial complexity of the transitions between tropical-temperate climates in the Old World. By considering climatic transitions we identified that the early ancestors of the Asian Palmates had strong affinities for the subtropical-temperate transitions suggesting an initial diversification under such climate. This leads us to propose that the Asian Palmates diversified under these transitional climates and its evolution was interspersed with occasional shifts towards more pronounced tropical or temperate climates. Such results emphasized the significance of transitional climates in understanding niche diversity and evolutionary patterns and calls for a (re)

evaluation of the role of transitional areas in promoting biodiversity across the Tree of Life.

## S.132.3 A diversification-transition model between tropical and temperate biomes explains the Northern Hemisphere biodiversity

Ze-Long Nie¹, Ying Meng¹, Meng-Hua Zhang¹, Luke Sparreo², Alicia Talavera³, Jun Wen³

1. College of Biology and Environmental Sciences, Jishou University, Jishou, Hunan, China. 2. Department of Botany, Connecticut College, New London, CT, U.S.A. 3. Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A.

The evolutionary patterns and processes driving ecological transition and diversification between tropical and north temperate biomes are complex and remain poorly understood in the Northern Hemisphere flora. Moreover, it is becoming clear that introgression is an important driving force of speciation in plant diversity. Here, we applied phylogenetic and biogeographic analyses to account for both introgression and incomplete lineage sorting based on genomic data from two charismatic lineages of the Northern Hemisphere, the tribe Viteae (the *Ampelocissus-Vitis* group) of Vitaceae and *Magnolia* of Magnoliaceae. Biogeographic inference and fossil evidence suggest that the ancestors of Viteae were widely distributed from North America to Europe during the Paleocene to the Eocene, followed by widespread extinction and survival of relicts in the tropical New World. During the climate warming in the early Miocene, an ancestor migrated northward from the refugia with subsequent diversification in North America and Eurasia. Similarly, *Magnolia* had once been widely distributed in the Northern Hemisphere in the early Palaeogene followed with extensive extinction and southward retreat to the low-latitude regions during the middle to late Eocene cooling climate. Magnoliaceae had been well diversified in the tropical regions with less development during the transition back to the temperate regions. Meanwhile, we found strong evidence for widespread incongruence and reticulate evolution among nuclear genes within both recent and an-
<figure><img src="image_310.png" /></figure>

cient lineages in the *Ampelocissus-Vitis* clade and *Magnolia* during their biome transitions and diversification in the Northern Hemisphere. The scenario we report here may represent a transition model between tropical and temperate regions in general for flowering plants that adapted to the global climate cooling and fluctuation in the Northern Hemisphere.

## S.132.4 The modernization of plant diversity on the Qinghai-Tibetan Plateau associated with the late Eocene climate transition

Tao Su¹,², Zhe-Kun Zhou², Jia Liu¹, Xin-Wen Zhang¹, Jiang Huang², Shu-Feng Li²

1 State Key Laboratory of Oil and Gas Reservoir Geology and Exploitation & Institute of Sedimentary Geology, Chengdu University of Technology, Chengdu, China. 2 CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, China

The modernization of plant diversity, which means the floristic assemblage in deep time being similar to nowadays, is a milestone for the evolution of global plant diversity; however, little is known about when did this process occur. Recent paleobotanical investigation on the Qinghai-Tibetan Plateau (QTP) sheds new lights on this topic. The late Paleocene Liuqu flora (~59 Ma) from southern QTP mainly consists of Annonaceae, Fabaceae, Moraceae and Palmae, suggesting a tropical forest. The middle Eocene Jianglang flora (~47 Ma) from central QTP is composed of Apocynaceae, Araceae, Menispermaceae, Sapindaceae and Simaroubaceae, there is no dominant species in this subtropic flora. The Dayu flora (~39 Ma) from central QTP consists of both woody plants and herbs, indicating the vegetation of open woodland. The floristic components of all these floras mentioned above are quite different from modern vegetations. It is not until the latest Eocene that the floristic components are similar to nowadays, which is evidenced by the Markam flora from eastern QTP with Fagaceae and Betulaceae as the most abundant taxa. This kind of evergreen deciduous broad-leaved mixed forest still survives in southeastern margin of the QTP nowadays. During the latest Eocene, the growth of the QTP and the associated climate change from dry to wet condition may contribute to the modernization of plant diversity on the QTP and regions nearby, and gradually shaped the rich plant diversity in this large area.

## S.132.5 Terrestrial plant evolution driven by the global cooling during the Eocene-Oligocene transition-examples from three tribes of Leguminosae

Lei Duan¹, Hong-Feng Chen¹, AJ Harris¹, Jun Wen²

1 South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China. 2 Department of Botany, Smithsonian Institution, Washington, USA

An abrupt, dramatic global cooling event occurred during the short period of the Eocene-Oligocene transition (EOT). This paleoclimatic event drove a large-scale aridification and increasingly seasonality, which drastically influenced the evolution of plants and distribution of global vegetation, especially, the great retreating of boreotropical flora. However so far, previous biogeographic studies have not well interpreted the molecular mechanism of the influence on terrestrial plants. The leguminous tribes Glycyrrhizeae (the liworice tribe), Adinobotryeae and Wisterieae (the Wisteria tribe) formed a clade, widely distributed in every continent except for Antarctica. Based on the data of orthologous nuclear genes and chloroplast genomes from deep next-generation sequencing, and using the interdisciplinary analyses of historical biogeography, molecular ecology and phylogenomics, our study revealed that the common ancestor of the three-tribe clade existed in boreotropical region during the Eocene, and experienced inter-tribal divergence, rapid niche evolution and biome shift from humid tropical forest to drier grassland or desert at the EOT. Subsequently, adaptive evolution occurred to many taxa of the clade, especially those of Glycyrrhizeae, to survive the changing habitats. On the other hand, the EOT global cooling differentiated the plant habits among the three tribes: herbs for Glycyrrhizeae, trees for Adinobotryeae and woody lianas for Wisterieae. To better investigate the noticeable point, with the annotated whole genomes obtained from the third-generation sequencing of representatives for each of the tribes, this study assessed the genome structure change and the evolution of the key gene families, whose function was to regulate the trait of habits by influencing the biosyntheses of cellulose and lignin. With the three tribes as a model, our results can provide a reference to the studies on plant evolution under the present background of global environmental changes.
<figure><img src="image_311.png" /></figure>

**S.132.6 Machine learning analyses of herbarium specimens identify morphological features characterizing tropical to temperate transitions**

Richard Hodel¹,², Alicia Talavera¹, Will Weaver³, Alex White², Mike Trizna², Rebecca Dikow⁴, Liz Zimmer¹, Ze-Long Nie⁵, Jun Wen¹

¹Department of Botany, National Museum of Natural History, MRC166, Smithsonian Institution, Washington, DC, USA. ²Data Science Lab, Office of the Chief Information Officer, Smithsonian Institution, Washington, DC, USA. ³Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI, USA. ⁴Computational Methods and Data, Yale University Library, Yale University, New Haven, CT, USA. ⁵Key Laboratory of Plant Resources Conservation and Utilization, College of Biology and Environmental Sciences, Jishou University, Jishou, Hunan, China

The phylogenomic age has revealed that the number of plant lineages that can successfully transition between temperate and tropical biomes is surprisingly low. One explanation for this trend invokes the difficulty of tropical lineages acquiring the necessary traits to tolerate the cooler, drier, and more seasonal temperate zone. Nevertheless, in rare cases, some lineages have been able to successfully migrate from the tropics to temperate zones during their evolutionary history. Presumably, these tran-

sitions were facilitated by the acquisition of morphological characters enabling adaptation to novel habitats (i.e., temperate environments). Synthesis of biogeographic studies of many plant lineages improved our understanding of the frequency of tropical-temperate transitions, but our understanding of associated trait evolution lags behind. Here, we use novel approaches for quantifying phenotype-machine learning algorithms that can classify and extract leaf, flower, and fruit features from digitized herbarium specimen images—to infer the morphological characters that were important for enabling transitions from tropical to temperate biomes. Specifically, we used the machine learning package LeafMachine2 to extract leaf, flower, and fruit traits from tens of thousands of specimens in targeted groups. We used the plum/cherry genus (Prunus, Rosaceae) and the grape genus (Vitis, Viaceae) because they represent rare lineages that successfully transitioned from tropical to temperate biomes. We used morphological data from tens of thousands of digitized specimens representing hundreds of species to trace continuous character evolution on time-calibrated phylogenies. This approach was selected so that the effects of both evolutionary history and past environmental conditions on observed phenotype could be considered. We measured leaf and reproductive characters and assessed phenology to identify features associated with biome shifts. Additionally, for comparative purposes, we investigated morphological variation within extant widespread species. The workflow developed for connecting morphological data with phylogenetic and environmental data is available as a user-customizable pipeline.

**S.133 THE SPECIAL AND ENDURING VALUE OF MODEL CLADES IN EVOLUTION AND ECOLOGY**

**S.133.1 What is a model clade? Viburnum as an example**

Erika J. Edwards¹, Michael J. Donoghue¹

¹Department of Ecology and Evolutionary Biology, Yale University, New Haven CT, USA

What makes a clade a model clade? In our view it is not because it provides a good example of some

particular phenomenon or process. This, we think, is true of every clade. Likewise, it is not because it is a clade that includes a model species such as *Arabidopsis thaliana*. Nor is it because it somehow represents a microcosm of the variety manifested by a more inclusive clade. Instead, we adopt the view that a model clade is simply one that has been studied intensively and from many different angles, leading naturally to a more holistic understanding. The expectation is that this concerted, integrative attention to a group of organisms will yield insights that would
<figure><img src="image_312.png" /></figure>

not otherwise be attainable, and this accumulation of context will invariably improve our interpretation of any one study in isolation. We view this context as critical in evaluating macro-evolutionary data and predict that studies of model clades will continue to provide the most compelling explanations of organismal evolution. Although there are no rules for the development of a model clade, there are some things that are foundational in providing a platform for diverse studies. One key element is the development of a firm understanding of phylogenetic relationships and of a phylogenetic classification system to support unambiguous communication across labs and generations. An additional consideration is clade size. There are distinct advantages to clades of moderate size – large enough to support convincing statistical analysis, but not so large that comprehensive surveys become too challenging. Viburnum (Adoxaceae) provides an example of a model clade in the sense that we have outlined here, and we provide concrete examples of novel insights that have emerged as a consequence of having carried out a diverse set of studies of Viburnum over multiple decades.

### S.133.2 The tribe Bignonieae (Bignoniaceae) as a model for understanding evolution, ecology, and biogeography in the Neotropics

Lúcia G. Lohmann¹, ²

University of California, Berkeley, California, USA.
2.Universidade de São Paulo, São Paulo, Brazil.

The tribe Bignonieae (Bignoniaceae) comprise ca. 20 genera and 400 species that are broadly distributed through the Neotropics, representing the largest and most successful clade of lianas in this part of the World. Species of Bignonieae show diverse morphology, ecology, and distribution, occurring in all major ecological zones (dry to wet) and providing an excellent opportunity to further understand habitat movements, niche transitions, and adaptations to dry/wet environments. The comprehensive phylogenetic, morphological, distribution, fossil, and ecological datasets gathered during the past three decades allow us to test hypotheses on the evolution of morphology, ecology, and biogeography of this important plant clade. Some of this work involved detailed studies of traits associated with

the climbing habit (e.g., wood anatomy, tendrils), insect-plant interactions (e.g., nectaries, mitedomatia, trichomes), pollination systems (e.g., traits associated with different flower whorls), and dispersal (i.e., seeds). While Bignoniaceae is a model system, our exhaustive exploration of the Neotropical Bignoniaceae established the basis for cross-taxonomic and multi-disciplinary studies that combine similar datasets for other organisms. The understanding of species' movements to dry environments across clades is allowing us to gain a deeper appreciation of clade-specific differences in the ability to adapt to climate change. Information of this nature is crucial to manage biodiversity, design sound policy, and establish conservation priorities.

### S.133.3 Hydnophytinae as a model clade

Guillaume Chomicki¹

1 Department of Biosciences, Durham University, Durham, UK.

Widespread in nature, mutualistic associations – cooperative interactions between unrelated species– are pivotal for the generation and maintenance of biodiversity. These partnerships vary from loose, facultative interactions between large guilds to tight, obligate interactions on which species depend for reproduction or survival. The Hydnophytinae is a clade of 105 species of epiphytic Rubiaceae, most of which form symbiotic mutualisms with ants. Despite its size, the Hydnophytinae have a higher rate of evolutionary transitions in mutualistic dependence magnitudes than mycorrhizal, pollination and seed dispersal mutualisms, making them an ideal model clade to study the evolution of mutualisms. I synthesize a decade of work and show how integrative, phylogenetically-informed studies of Hydnophytinae ant/plant symbioses have shed major insights on our understanding of mutualisms as a whole. Using phylogenetic comparative methods, field experiments, comparative genomics and transcriptomics, 3D imaging, microbiology and modeling, I show how mutualistic dependence is critical for determining the evolutionary stability of the partnership, its efficiency in trading resources, and show how it mediates trait evolution within and beyond those involved in the interaction. I also show how the Hydnophytinae can help us resolve the paradox of multi-species mutualisms and share new insights on mechanisms promoting obligate dependence between 'macro' partners.
<figure><img src="image_313.png" /></figure>

**S.133.4 The where, when, and why of pulsed trait evolution in an explosive plant radiation, Lupinus (Fabaceae)**

Livio Bätscher¹, Colin E. Hughes², Jurriaan M. de Vos¹

1 Department of Environmental Sciences, University of Basel, Basel, CH. 2 Department of Systematic and Evolutionary Botany, University of Zürich, Zürich, CH

We present an overview of why the genus *Lupinus* L. (Fabaceae) has emerged as a model clade for studying rapid recent plant radiations providing potent insights into the drives rapid ecological and phenotypic plant diversification. The Western New World *Lupinus* clade with more than 200 species, comprises a series of nested and parallel radiations, including the recent rapid Andean radiation. Previous work has shown that a combination of biome shifts and life form changes underpin accelerated species diversification, but detailed understanding of the interactions between environmental and phenotypic evolution is lacking. Moreover, while multiple drivers of high rates of species diversification have been proposed, the drivers of rapid ecological and morphological trait evolution remain unclear. Here we leverage taxonomic, morphological, biogeographic, and environmental data to ask what environmental factors drove the apparently differential rates of trait evolution among western New World *Lupinus* lineages. Specifically, we test which model classes, including models of incremental or pulsed change (i.e., Lévy processes), best describe evolution across eight traits, and whether life forms, climatic niches, and biographic realms affect the tempo and mode of trait evolution. We demonstrate a radically expanded trait space occupancy by the Andean clade. We reveal support for pulsed evolution in three of eight plant traits and identify five nested branches in the Andean clade that are congruent with “evolutionary jumps”, rather than a single shift to an overall higher rate of incremental evolution. We find that these shifts are associated with perennials occupying aseasonal temperature niches, but not all such lineages experience “jumps”, suggesting that longer life spans and longer growing seasons are prerequisites but, alone are insufficient to drive episodes of pulsed trait evolution.

Our findings demonstrate the existence of rapid bursts of trait evolution and suggest that pulsed evolution may be a common feature of plant radiations.

**S.133.5 Sarraceniaceae as a model clade**

David W. Armitage¹

1 Integrative Community Ecology Unit, Okinawa Institute of Science and Technology, Onna, Okinawa, Japan

Sarraceniaceae is the *Drosophila* of community ecology. The fluid-filled leaves of these carnivorous plants have, over the course of centuries, captivated professional and amateur botanists alike, and — owing to their unique phytotelmic habit — have been used by ecologists, entomologists, and microbiologists as miniature models for lakes, island chains, and animal digestive systems. The balance between tractability and realism in Sarraceniaceae microcosm studies has produced many key insights into the functioning and dynamics of communities and ecosystems that would otherwise be impossible in larger systems. Sarraceniaceae are similarly important from botanical and evolutionary perspectives. Captivating as these plants are, however, many aspects of their evolution and biology remain shrouded in mystery. While their phylogenetic placement is becoming clear, far less is known about their landscape genetic structuring, trait-environment relationships, and mechanisms of prey attraction, capture, digestion, and dispersal. Furthermore, major questions remain concerning the evolutionary history of the group, as well as mechanisms contributing to their local-scale diversification in the South-eastern United States and Guiana Highlands. In this talk, I discuss my research history with Sarraceniaceae and *Darlingtonia californica*, in particular. I show how an integration of ecological, genetic, and biomechanical studies focusing almost entirely on this single clade has, perhaps paradoxically, helped me become a more interdisciplinary scientist and more effective advisor and collaborator. I also introduce a brief history of the use of the Sarraceniaceae model system in community ecology — highlighting the groundwork of both established and early career researchers — and conclude by outlining areas which would further help enshrine the family as a model system that can effectively integrate many fields of science.
<figure><img src="image_314.png" /></figure>

S.133.6 Cuscuta (dodders; Convolvulaceae) as a model-system to study biology of heterotrophic plants

Saša Stefanović¹, Miguel A. García², Mihai Costea³

1 University of Toronto Mississauga, Mississauga, Canada. 2 Royal Botanical Gardens, Madrid, Spain. 3 Wilfrid Laurier University, Waterloo, Canada

Among several dozen independently evolved
groups of heterotrophs, *Cuscuta* is emerging as a
model clade for understanding different aspects
of heterotrophic plant biology due to the conver-
gence of several factors. First, *Cuscuta* is one of the
largest and most economically important lineages
of heterotrophic plants. Second, phylogenetic rela-
tionships within this species-rich group are well-re-
solved and strongly supported. Third, the shallow
phylogenetic level that encapsulates substantial
diversity observed in this genus makes the system
tractable. We are now interested to use this system
to explore in-depth three particular venues of het-
erotrophic plant biology: *Cuscuta* is one of only two
clades of haustorial parasites known to span the

trophic continuum from photosynthetic hemipara-
sites to non-photosynthetic, obligate holoparasites.
Most of its lineages are "cryptically photosynthetic"
and capable of limited and localized photosynthe-
sis. However, we were unable to recover any plas-
tome sequences from section *Subulatae*, suggest-
ing that plastomes of its species are likely entirely
lost, a condition encountered only once before in the
angiosperm evolution (Rafflesiaceae). Furthermore,
*Cuscuta* may represent a genus with the broadest
karyotypic and genome size diversity among all of
angiosperms. This group includes species not only
with regular monocentric but also with holocentric
chromosomes, a feature found in a very few phylo-
genetically scattered groups of plants. In addition,
the genus has diverse karyotypes, with enormous
variation reported for chromosome numbers, chro-
mosome size, and variation in nuclear DNA content
as well as presence of symmetrical and asymmetri-
cal karyotypes. Discordances between phylogenies
derived from different data sources suggest that
reticulation and polyploidy have likely played a sig-
nificant role in *Cuscuta* species diversification, with
potentially 10% of its species being of hybrid origin.
Some of these cases are consistent with more an-
cient hybridization events while represent relatively
recent reticulations. Our current progress in these
three areas of *Cuscuta* biology will be discussed.

S.134 TELLING PLANT SPECIES APART WITH
DNA. SESSION 1

S.134.1 Molecular barcoding
identification of invasive
species dispersed by wind
and in the soil seed bank
of South Georgia using
phylogen

Rosemary Newton¹, Calum Sweeney¹,²,
Rachel Day¹, Kaitalin White¹, Wayne
Dawson³,⁴, Bradley Myer⁵, Kelvin Floyd⁵,
Marcella Corcoran¹, Carey Kelting¹, Sally
Poncet⁵, Ken Passfield⁵, Felix Forest¹, Colin
Clubbe¹, Juan Viruel¹*

1 Royal Botanic Gardens, Kew, Richmond, UK, 2 Na-
tional Parks and Wildlife Services, Kenmare Road,
Killarney, Kerry, Ireland. 3 Department of Bioscienc-
es, Durham University, Durham, UK. 4 Department
of Evolution, Ecology and Behaviour, University of
Liverpool, Liverpool, UK. 5 Indigena Biosecurity Inter-
national, Nelson, New Zealand * Author for corre-
spondence: j.viruel@kew.org

Climate change and invasive species are major
causes of biodiversity loss. Invasive species are highly
competitive in ecosystems due to their ability to ger-
minate and grow in a wide range of conditions, their
capacity to produce high reproductive outputs and
their effective dispersal strategies. Climate change
has already severely impacted several ecosystems
across the planet, such as sub-Antarctic regions
with temperatures rising there fastest in recent de-
cades. The effects of climate change can exacerbate
<figure><img src="image_315.png" /></figure>

the impacts of invasive species by enhancing features that favour invasiveness, such as tolerance to a broad range of environments. Islands are particularly vulnerable to the effects of climate change due to reduced habitable space, small populations, and low functional redundancy. The remote sub-Antarctic island of South Georgia is 160 km in length with a vegetated area of approximately 30,000 ha predominantly confined to near coastal regions due to snow and ice being prominently present inland. South Georgia has a history of use as a land base for extraction of natural biotic resources from the surrounding coasts and ocean. Due to human occupation, non-native flora and fauna were inadvertently and/or intentionally introduced. Today, the introduced flora with 41 species outnumbers the native flora with 24 species. To mitigate this problem, the South Georgia Non-Native Plant Management Strategy was implemented during 2016-2020 to reduce the population size of non-native species on the island. We evaluate the efficacy of this eradication programme by quantifying the incidence of invasive species seeds in the soil seed bank. We designed a molecular barcoding approach to enable accurate identification of seedlings by evaluating the resolution of traditional barcoding markers (i.e., *rbcL* and *matK*) and optimising a high-throughput sequencing molecular barcoding method. We applied this method to identify seedlings that emerged from soil samples collected at key areas on the island.

## S.134.2 An evaluation of the taxonomic status of *Musa viridis*, *Musa splendida*, and *Musa paracoccinea* using high-throughput sequencing data

Yves Bawin¹,²,³, Arne Mertens¹, Sander de Backer¹, Dang Toan Vu⁴, Loan Thi Le⁴, Tuong Dang Vu⁴, Filip Vandelook¹,³, Sofie Meeus¹, Steven B. Janssens¹,³

1 Meise Botanic Garden, Meise, Belgium. 2 Hasselt University, Hasselt, Belgium. 3 KU Leuven, Leuven, Belgium. 4 Vietnamese Academy of Agricultural Sciences, Hanoi, Vietnam

The delimitation of species based on morphological traits can be challenging due to overlapping trait variation. The taxonomic identity of closely related sympatric species with individuals displaying inter-

mediate phenotypes is particularly questionable. However, thorough morphological assessments or traditional molecular analyses often lack resolution to clarify if two species are truly separate taxonomic entities or should be merged. Interestingly, advances in high-throughput sequencing methods and data processing during the last decades enable the use of high-resolution data to tackle taxonomic problems. Several species with an unclear taxonomic status are found in the banana family (Musaceae). In Vietnam, the *Musa* species *M. viridis*, *M. splendida*, and *M. paracoccinea* have very similar morphological trait characteristics. The main difference between *M. splendida* and the two other *Musa* species is the colour of the male flower bud: *M. splendida* has red flower buds, whereas *M. viridis* and *M. paracoccinea* have pink flower buds. All three species occur symmetrically, and individuals with red and pink male flower buds are found together in some populations. In this study, we used high-throughput (DArT) sequencing data from plants of nine populations to investigate the taxonomic status of these *Musa* species. Maximum likelihood phylogenetic trees were reconstructed for over 12,000 loci (< 120 base pairs) and one ASTRAL consensus tree was created from all locus trees. The individuals in the ASTRAL phylogenetic tree did not cluster by species identity, but by population, having individuals from different species in the same cluster. Consequently, we propose to merge the species *M. viridis*, *M. splendida*, and *M. paracoccinea* into one species named *M. splendida* according to nomenclatural conventions.

## S.134.3 Precision of molecular plant identification of plants in the coastal dunes in Thy National Park, Denmark

Conny B.A. Lange¹, Henrik Æ. Pedersen³, Else Ø. Andersen⁴ and Anders, S. Barfod²

1 Department of Plant and Environmental Sciences, University of Copenhagen, Denmark. 2 Department of Bioscience, University of Aarhus, Denmark. 3 Select Nature, Denmark 4 Thy National Park, Denmark

Coastal sand dunes are highly dynamic landscapes shaped by wind, water and salt. In Europe they are mainly distributed along the Atlantic coastlines. In Denmark sand dunes are widespread and are characterized by a rapid turnover of the constituent habitat types. In this project we address the need
for novel and efficient ways of monitoring dunes dynamics. Traditionally, a trained botanist identify plants either by recognizing them or using a scientific flora. However, this procedure is constricted to the flowering season and often requires complete plant specimens. Molecular plant identification is a tool that may prolong the monitoring season, can identify plants based on fragments, and can add to the quality of plant identification. To test the precision of molecular plant identification we collected four individuals from each of 100 plant species common to or characteristic of the sand dunes in Thy National Park. For each individual we sampled a herbarium voucher specimen and leaf material dried and preserved in silica gel. From the leaf material the DNA barcode genes *rbcL* and *matK* were sequenced. We used *rbcL* sequences to probe into the ability of the international databases, Genbank and BOLD to retrieve the correct identifications. Less than 50% of the individuals were unambiguously identified to the correct species. However, a barcode gap existed between all species, except for a few hybrids of *Juncus*. A Neighbour Joining analysis clustered individuals of each species and was congruent with the current APGIV classification. We conclude that caution should be exercised when basing plant identification purely on BOLD and GenBank. The most important sources of error in the online databases were mismatch between the sequence uploaded and the underlying herbarium voucher specimen, wrong identification of herbarium specimens, and mismatch between the length of the sequence in the database and the sequence submitted.

### S.134.4 Next-generation DNA barcoding using phylogenomic data

Benedikt G. Kuhnhäuser¹*, Jason Stevenson¹, Rowan Schley², László Csiba¹, Sidonie Bellot¹, William J. Baker¹

1 Royal Botanic Gardens Kew, UK. 2 University of Exeter, UK. * b.kuhnhaeuser@kew.org

DNA barcoding has the potential to enable and speed up plant identification, but traditional barcoding using single to few genes is limited by insufficient discriminatory power in plants. Novel sequencing technologies can routinely generate hundreds of genes even from degraded plant material, promising to overcome the current limitations and unlock a new era of next-generation DNA barcoding. Despite this potential, no bar-

coding tools exist that can handle dozens or hundreds of genes. To address this methodological gap, we have developed a novel approach to DNA barcoding that can take the information content of hundreds of genes into account. Here, we present our DNA identification toolkit and illustrate its potential to revolutionise plant identification using a case study of rattan furniture authentication. By comparison with traditional barcoding markers, we demonstrate a substantial increase in both accuracy and interpretability of our approach compared to existing methods. The toolkit is publicly and freely available (https://github.com/BenKuhnhaeuser/RattanID).

### S.134.5 Towards a nuclear DNA barcode for land plants

Peter M. Hollingsworth¹, Wu Huang¹, Cornelia Simon-Nutbrown¹, Flavia Pezzini¹, Li De-Zhu², Alex D. Twyford³

1 Royal Botanic Garden Edinburgh, Edinburgh, UK. 2 Kunming Institute of Botany Chinese Academy of Sciences, Kunming, China. 3 University of Edinburgh, Edinburgh, UK

Accurate and rapid identification of plant species is of fundamental importance for understanding global biodiversity, monitoring change, and managing biodiversity. Yet plant species identification can be challenging in many situations. To address this challenge, a substantial body of data has been generated using standard plant barcodes such as *rbcL*, *matK*, *trnH-psbA* and ITS for telling plant species apart. These barcode markers are useful for many applications, but do not provide universal species-level discrimination. There is thus considerable interest in developing new standardised plant barcoding methods. In this presentation we (1) review the strengths and limitations of the standard barcoding approach; (2) explore the potential for gains in discriminatory power by using complete plastid genomes as plant barcodes; and (3) explore recent work considering options for utilisation of nuclear loci for plant DNA barcoding. In the exploration of nuclear loci for species identification we cover the frequency with which plant species resolve as monophyletic based on studies to-date, the number of randomly selected nuclear loci required to give maximal species discrimination, and whether there are individual loci that show any exceptional species discrimination performance. We conclude by outlining a roadmap for further developing nuclear DNA barcoding approaches for plants.
## S.134.6 Plant DNA barcode library for native flowering plants in the arid region of northwestern China

Feng Song¹, Ting Li¹, Hai-Fei Yan¹, Ying Feng², Lu Jin¹, Kevin S. Burgess³, Xue-Jun Ge¹

1 Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China. 2. State Key Laboratory of Desert and Oasis Ecology, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi, China. 3. Department of Biology, Columbus State University, University System of Georgia, Columbus, USA

DNA barcoding is a well-established tool for rapid species identification and biodiversity monitoring. A reliable and traceable DNA barcode reference library with extensive coverage is necessary but unavailable for many geographical regions. The arid region in northwestern China, with a vast area of about 2.5 million km², is ecologically fragile and often overlooked in biodiversity studies. In particular, DNA barcode data from the arid region of China is lacking. We develop and evaluate the efficacy of an

extensive DNA barcode library for native flowering plants in the arid region of northwestern China. Plant specimens were collected, identified and vouchered for this purpose. The database utilized four DNA barcode markers, namely *rbcL*, *matK*, ITS and ITS2, for 1816 accessions (representing 890 species from 385 genera and 72 families), and consisted of 5196 barcode sequences. Individual barcodes varied in resolution rates: species- and genus-level rates for *rbcL*, *matK*, ITS and ITS2 were 79.9%–51.1%/76.1%, 79.9%–67.2%/88.9%, 85.0%–72.0%/88.2% and 81.0%–67.4%/84.9%, respectively. The three-barcode combination of *rbcL*+*matK*+ITS (RMI) revealed a higher species- and genus-level resolution (75.5%/92.1%, respectively). A total of 110 plastomes were newly generated as super-barcodes to increase species resolution for seven species-rich genera, namely *Astragalus*, *Caragana*, *Lactuca*, *Lappula*, *Lepidium*, *Silene* and *Zygophyllum*. Plastomes revealed higher species resolution compared to standard DNA barcodes and their combination. We suggest future databases include super-barcodes, especially for species-rich and complex genera. The plant DNA barcode library in the current study provides a valuable resource for future biological investigations in the arid regions of China.

## S.135 EVOLUTION AND FUNCTION OF CONVERGENT FLORAL PHENOTYPES. SESSION 1

### S.135.1 Trade-off mitigation: a key concept to understand the evolution and function of convergent floral phenotypes

Kazuharu Ohashi¹, Ayumu Tanaka¹, Andreas Jürgens², James D. Thomson³

1 University of Tsukuba, Tsukuba, Japan. 2 Technische Universität Darmstadt, Darmstadt, Germany. 3 University of Toronto, Toronto, Canada

Flowers are often thought to have adapted to pollination by specific animals due to the trade-off that one flower cannot adapt to diverse pollinators. If this is true, why do most flowers harbor various visitors, while maintaining distinct phenotypes among ecotypes, subspecies, and congeners? Because strong phenotypic trade-offs leading to disruptive selection have only been observed in highly specialized systems, we proposed a hypothesis that flowers can adapt to diverse pollinators simultaneously by evolutionarily mitigating trade-offs (Ohashi et al. 2021). In this talk, we explore three ideas about how floral phenotypic convergence evolves and functions from the perspective of trade-off mitigation. First, we demonstrate that certain phenotypic convergence in flowers—floral color change, compact inflorescences, and nocturnal anthesis—could have resulted from adaptive generalization for particular pollinator communities rather than adaptive specialization for specific pollinators. Next, based on the fact that even specialist flowers are often visited by minor pollinators, we discuss that any pollination system can be positioned along a continuum of “generalized systems for specific sets of pollinator groups.” With this in mind, we propose three types of functional traits that constitute a floral syndrome: those that improve pollination by primary visitation.
<figure><img src="image_316.png" /></figure>

tors, those that exclude flower visitors that act as con-
ditional parasites, and those that modify trait-fitness
relationships to mitigate trade-offs among pollinators.
Finally, we suggest that the concept of trade-off miti-
gation may provide an explanation for the uniformity
of floral phenotypes in some clades. Flowers attracting
a wide range of pollinators, such as compact inflores-
cences in Apiaceae, may have achieved "super-mit-
igating" phenotypes that could eliminate fitness val-
leys among all types of pollinators. In such flowers, any
local shift in pollinator composition will not exert dis-
ruptive selection pressure but rather result in a slight
change of the adaptive peak.

S.135.2 Andromonoecy as
a convergent resolution to
intra-locus sexual conflict in
bisexual flowers

Kai-Hsiu Chen¹, John R. Pannell¹

1 University of Lausanne, Department of Ecology and Evolution, Lausanne, Switzerland

Andromonoecy, the production of male and bisexu-
al flowers by the same hermaphroditic individual, has
evolved numerous times independently across flow-
ering plants and is found in 7% of species. Despite its
repeated evolution, its function remains poorly studied.
Here, we consider the convergent evolution of andro-
monoecy as a resolution to sexual antagonism, draw-
ing on data from the alpine plant *Pulsatilla alpina* that
shares common features with many andromonoe-
cious species. *Hypothesis*: We expected to detect ev-
idence for strong sexual conflict on female allocation
but not male allocation traits in bisexual flowers, i.e.,
female and male functions have different fitness opti-
ma only in the female allocation trait, thus favoring the
production of unisexual male and bisexual flowers. We
analyzed and compared selection gradients via fe-
male and male functions for five floral traits, including
sex allocation, phenology, and display traits, in bisexual
flowers of *P. alpina* using a paternity analysis. Our results
indicate strong sexual conflict in the number of pistils
and flowering date but not in the number of stamens
in bisexual flowers. Specifically, we found evidence for
disruptive selection on pistil number within flowers, such
that flowers with many or no pistils contributed more
to fitness than those with an intermediate number of
pistils. By abandoning the female function, unisexual
male flowers possess phenotypes that approach the
male optimum of different floral traits. We show that the
unisexual male flowers of andromonoecious species

may allow hermaphrodites to resolve sexual conflict
because modules with different genders can promote
fitness through their specialized sexual functions. We
discuss the extent to which modular variation in sex al-
location in species with other sexual systems involving
unisexuality and bisexuality (e.g., monoecy and gyno-
monoecy) may be explained similarly.

S.135.3 Plasticity-mediated
floral convergence triggers
the exploration of new
generalist pollinators niches

José M. Gómez¹,², Adela González-Megías²,³,
Eduardo Narbona⁴, Luis Navarro⁵, Francisco
Perfectti²,⁶, Cristina Armas¹

1 Estación Experimental de Zonas Áridas (CSIC), Al-
mería, Spain. 2 Research Unit Modeling Nature, Uni-
versidad de Granada, Granada, Spain. 3 Dpto. de
Zoología, Universidad de Granada, Granada, Spain.
4 Dpto. de Biología Molecular e Ingeniería Bioquími-
ca, Universidad Pablo de Olavide, Sevilla, Spain. 5
Dpto. de Biología Vegetal y Ciencias del Suelo, Uni-
versidad de Vigo, Vigo, Spain. 6 Dpto. de Genética,
Universidad de Granada, Granada, Spain.

Phenotypic convergence is a ubiquitous phenom-
enon that reveals the multiple evolutionary con-
straints that face the organisms and illustrates
the ability of natural selection to find similar solu-
tions to similar ecological problems. Uncovering
the possible mechanisms behind the origin of
convergence remains a fundamental goal in bi-
ology. Evolutionary theory shows that convergent
phenotypes emerge through independent genetic
changes selected over long periods of time. Here
we show that convergence can also arise through
phenotypic plasticity. By compiling the multidim-
mensional floral phenotype, the phylogenetic
relationships, and the pollination niche of more
than 3000 Brassicaceae species, we demonstrat-
ed that the mustard species *Moricandia arvensis*
exhibits a plastic-mediated intra-individual floral
disparity greater than that found not only between
species but also between higher taxonomical lev-
els such as genera and tribes. As a consequence
of this floral divergence, *M. arvensis* moves outside
the morphospace region occupied by its recent
ancestors and close relatives, crosses into a new
region where it encounters a different pollination
niche and converges phenotypically with distant
Brassicaceae lineages. We suggests that, by in-
ducing several phenotypes that explore simul-
<figure><img src="image_317.png" /></figure>

taneously different regions of the morphological space, plasticity may not only promote the evolution of novelties but also trigger rapid phenotypic convergence.

**S.135.4 Convergent evolution of sexual deception in Gorteria diffusa (Asteraceae)**

Beverley J. Glover¹, Boris Delahaie¹, Allan G. Ellis², Roisin Fattorini¹, Roman Kellenberger¹, Farahnoz Khojayori¹, Greg Mellers¹, Udhaya Ponraj¹, Xiao Wang¹

1 Department of Plant Sciences, University of Cambridge, UK. 2 Department of Botany and Zoology, Stellenbosch University, South Africa

Gorteria diffusa is endemic to South Africa, but its floral morphology varies extensively across its range. Geographically distinct populations produce capitula with no petal spots, with simple petal spots and with highly complex petal spots that mimic female bee flies. These last forms attract pollinators through sexual deception. We have studied the development of these complex petal spots and their evolution within the species, and find that sexual deception has emerged at least twice in the system. I describe our work to understand the genetic and genomic changes that underlie the repeated evolution of this novel trait.

**S.135.5 Convergent evolution of hummingbird pollination in the Neotropical spiral gingers: traits, genetics, and the selective environment**

Kathleen M. Kay¹, Kathleen Darragh², Dena L. Grossenbacher³, Pedro Juárez¹, Santiago Ramírez⁴, Kathryn Uckele¹, Oscar M. Vargas⁵

1 Department of Ecology and Evolutionary Biology, University of California, Santa Cruz; Santa Cruz, CA, USA. 2 Department of Biology, Indiana University, Bloomington, IN, USA. 3 Department of Biology, California Polytechnic State University, San Luis Obispo, CA, USA. 4 Center for Population Biology, University of California, Davis; Davis, CA, USA. 5 Department

of Biological Sciences, California State Polytechnic University, Humboldt; Arcata, CA, USA

The evolution of hummingbird pollination is common across angiosperms throughout the Americas, presenting an opportunity to examine convergence in traits, genetics, and selective environments to better understand how complex phenotypes arise. We examine many independent shifts from bee to hummingbird pollination in the Neotropical spiral gingers (Costus; Costaceae) and address common explanations for the prevalence of these transitions. We reconstruct ancestral pollination states on a well-resolved phylogeny and examine variation in traits and environmental correlates across independent shifts. Using a phylogenomic approach, we assess historical introgression events to determine if convergence could involve the sharing of alleles across species. We further dissect the genetic basis of a recent bee-to-bird shift and compare those findings to genetic changes underlying floral traits in other Costus species. We also test the hypothesis that hummingbird pollination is adaptive in high-elevation pollination environments relative to bee pollination by translocating flowering plants of both types across an elevational gradient in Costa Rica. A consistent set of traits predict hummingbird pollination, but not the stereotypical 'hummingbird' traits of long, red flowers. We find many shifts to hummingbird pollination, no reversals, a single shared phenotypic optimum across hummingbird flowers, and little evidence of introgression. Our genetic investigations find large effect color and scent loci, small effect morphology loci, and some overlap in loci across independent floral transitions. In contrast to other clades, we find no association between pollination and climate, and mixed evidence that hummingbird pollination is adaptive in montane environments. Evolutionary shifts to hummingbird pollination in Costus are highly convergent and directional, involve a surprising set of traits when compared with other plants with analogous transitions, and refute the generality of several common explanations for the prevalence of transitions from bee to hummingbird pollination.

**S.135.6 Convergence and divergence: specialization and generalization in the most diverse poricidal flowered plant lineage**

Vinícius L. G. Brito¹

1 Instituto de Biologia, Universidade Federal de Uberlândia, Uberlândia, Brasil
<figure><img src="image_318.png" /></figure>

Buzz-pollinated flowers represent some of the most iconic examples of evolutionary convergence in angiosperms. Typically, buzz-pollinated flowers exhibit a syndrome of convergently evolved traits including poricidal anthers, dimorphic stamens, and pollen as the sole floral reward. This suite of floral traits is thought to have evolved together in ca. 25 angiosperm families. However, at finer taxonomic levels, these traits are not always associated with one another. For example, stamen dimorphism appears to be context-dependent within the Melastomataceae (the most diverse plant lineage with poricidal flowers): dimorphic stamens tend to be associated with pollinator dependence, whereas pollinator-independent species are more likely to have monomorphic

stamens. Furthermore, some melastomes have acquired divergent traits which are associated with increased pollinator specialization. For example, independent evolutionary origins of oils by the stamens appear to have made melastome pollen unpalatable to many bees, so that unrelated oil-producing flowers are almost entirely visited by a small subset of buzzing bees – those that also collect oil. On the other side of the specialization spectrum, some nectar-producing melastomes have increased pore size, allowing them to be visited by a more diverse range of insects. Together, the shifts towards more specialized or generalized systems suggest a complex pattern of both diverging and converging traits within paraphyletic melastome lineages.

# S.136 BRYOPHYTE DIVERSITY, ENVIRONMENT AND CONSERVATION IN MEDITERRANEAN CLIMATE AREAS

## S.136.1 California bryophytes: evolution and conservation in a biodiversity hotspot

Brent D. Mishler¹, Ixchel González-Ramírez¹, John Mclaughlin², Benjamin Carter², Israel Borokini³

1 University and Jepson Herbaria, University of California, Berkeley, USA. 2 Sharsmith Herbarium, San Jose State University, USA. 3 Department of Ecology, Montana State University, Bozeman, USA

California is one of the world's biodiversity hotspots. It is also one of the places where biodiversity is most threatened, because of habitat loss, invasive species, and climate change. There is an urgent need to understand conservation priorities on the landscape of California for all groups of organisms, yet no such study has been made for mosses or liverworts. Bryophytes are indicators of ecosystem health, and contribute significantly to nutrient and water cycles. Therefore, we carried out spatial phylogenetic analyses for both groups. For mosses we pruned a larger spatial dataset of geo-referenced herbarium records for North America, plus matching phylogenetic data matrix, from Carter et al.

(2022, Journal of Biogeography, 10.1111/jbi.14385) to make a California subset. For liverworts we assembled a new spatial data set from GBIF and mined molecular data from Genbank. A maximum likelihood phylogeny was constructed for both groups and used along with newly produced species niche models to find regions of significant phylogenetic diversity and phylogenetic endemism within California, employing a spatial randomization. We also carried out phylogenetic complementarity analyses to identify conservation priorities on the landscape that optimally increase protected biodiversity, using an algorithm that considers current land protection status and data on landscape intactness to identify priority sites containing concentrations of lineages that are evolutionarily unique, vulnerable due to small range size, and poorly protected across their ranges. We evaluated priorities using three different dimensions of phylodiversity): a phylogram (i.e., genetic divergence), a chronogram (i.e., evolutionary time), or a cladogram (i.e., speciation events). These metrics yielded conservation priorities that agreed in many places but differed in others. We compare these results to those previously published for vascular plants of California (Kling et al. 2018, Phil. Trans. Roy. Soc. B, 374: 20170397). Several top priority regions of the state emerged from these comparisons.
<figure><img src="image_319.png" /></figure>

## S.136.2 Orthotrichaceae (Bryophyta) in Mediterranean climate areas: unveiling an unexpected diversity in Central Chile

Francisco Lara¹,², Juan Larraín³, Ricardo Garilleti⁴, Isabel Draper¹,²

1 Departamento de Biología, Universidad Autónoma de Madrid, Spain. 2 Centro de Investigación en Biodiversidad y Cambio Global, Universidad Autónoma de Madrid, Spain. 3 Centro de Investigación en Recursos Naturales y Sustentabilidad Santiago, RM, Chile. 4 Departamento de Botánica y Geología, Universidad de Valencia, Spain.

The five global regions with Mediterranean climates harbor high and unique biodiversity, hence all being part of recognized hotspots. A climate characterized by a prolonged period of drought coinciding with the hottest time of the year may not seem, at first glance, favorable for bryophytes. However, several groups of mosses and liverworts are widely diversified in this environment. The representation of Orthotrichaceae in the different Mediterranean floristic regions is highly unequal, both in terms of the genera present, and in the overall specific diversity. The Mediterranean Basin and the California Floristic Province boast a notable diversity of species distributed among few genera, all belonging to the subfamily Orthotrichoideae. Conversely, in the Cape Floristic Province and Southwest Australia, there are relatively few species, but there is a significative presence of Macromitrioideae genera. In contrast to this, in Central Chile the diversity reported so far is very low, both in number of genera and species, with all taxa belonging to Orthotrichoideae. But, to what extent is this diversity scenery realistic for the southern hemisphere Mediterranean areas? Our studies in Central Chile indicate that there is indeed a considerable hidden diversity of Orthotrichaceae there. Specifically, our results unveil that there is a high number of previously unknown species belonging to the genera *Orthotrichum* and *Lewinskya*. Most are species restricted to the Chilean Mediterranean biome. However, two particularly diverse environments, sharing few species among them, can be distinguished: the fog oases of the coastal hills and the high elevation zones of the inland Mediterranean mountains. We provide

data on the adaptive morphological characteristics of the new species, their distribution patterns, and their phylogenetic relationships, considering a broad framework that integrates species from other South American areas and the remaining Mediterranean regions worldwide.

## S.136.3 Achievements, failures, and challenges in bryophyte conservation in Spain

Belén Albertos¹, Elena Ruiz², Patxi Heras³, Ricardo Garilleti¹, Marta Infante³

1 Universitat de València, Spain. 2 Universitat Autònoma de Barcelona, Spain. 3 Museo de Ciencias Naturales de Álava, Spain

The conservation of bryophytes in Spain is at a considerable standstill. Despite notable initiatives in the field of regulation, such as the elaboration of the standard list, the improvement of some regional protection catalogues and the process initiated to extend national catalogues, the number of protected species is insufficient and centered on species protected at the European level. Improvements in the legislative field do not necessarily mean an improvement in the protection of bryophyte species because the legally established management mechanisms are not developed. The decentralisation of the Spanish state, while improving the distribution of funds, limits the control mechanisms over the autonomous communities. The level of non-compliance, generally high in our administrations, is aggravated by the fact that it is a small group of species that does not justify the administrations' investment in specialised training for their technicians. The improvement of the administrative situation will hopefully lead to an improvement in investment in research and monitoring projects, as the current situation means that there is a lack of demand for such studies. Generating a need in the administration may increase the number of research projects, both requested and granted, on bryophyte conservation and stimulate interest in training staff in charge of the management of natural areas and protected species. A review of the current situation is presented, with its achievements and shortcomings, and the most relevant scientific, technical, and administrative needs are analysed.
<figure><img src="image_320.png" /></figure>

## S.136.4 Resilient nature of cosmopolitan mosses in the Mediterranean Basin: insights into climate-driven molecular adaptation

Mahmoud Magdy¹,², Olaf Werner¹, Rosa M. Ros¹

1 Department of Plant Biology, Faculty of Biology, Murcia University, Murcia, Spain. 2 Department of Genetics, Faculty of Agriculture, Ain Shams University, Cairo, Egypt

Across Mediterranean mountainous regions and surrounding lowlands, the ecological resilience of cosmopolitan mosses, *Bryum argenteum* Hedw., *Ceratodon purpureus* (Hedw.) Brid, and *Funaria hygrometrica* Hedw. delineated distinct clades. Our research story provides insights into the interaction between landscape diversity, climate change, and molecular adaptation in mosses. The genetic analyses of *B. argenteum* along the gradient surface of the Sierra Nevada Mountains compared to Tenerife Island unveiled a story of recurrent colonization spanning over millennia. Despite indications of native status, shared haplotypes with continental areas suggest recent introductions. Clade distinctions among genotypes mirrored enduring climatic barriers between the highlands and lowlands of the Sierra Nevada Mountains, influencing migration dynamics. In *C. purpureus*, morphological variations posed challenges in taxonomic delineation. The evolutionary trajectory and the emergence of a new *Ceratodon* species through peripatric speciation, genome expansion, and skewed sex ratios within the Mediterranean landscape. These genetic insights led to the identification of *Ceratodon amazonum* Nieto-Lugilde, O. Werner, S.F. McDaniel & Ros and the consideration of *Ceratodon conicus* (Hampe ex Mull. Hal.) Lindb., as a nothospecies, emphasizes the delicate impact of environmental factors on moss taxonomy amidst changing landscapes. Exploration of *F. hygrometrica* across the Sierra Nevada Mountains highlighted the correlation between landscape diversity and genetic makeup. Diverse landscapes harbored higher genetic diversity, possibly linked to spore establishment challenges in unsuitable environments. Altitudinal gradients influenced genetic structures, while precipitation and temperature shaped specific genetic markers, underscoring the pivotal role of environmental heterogeneity in shaping genetic diversity. Our ongoing investigation into the Heat-Shock Factor gene family aims to unravel mosses' molecular

responses to climatic shifts, using *F. hygrometrica* as a model. Deciphering genetic variations and expression dynamics among genotypes will illuminate how these molecular mechanisms orchestrate adaptive responses to high temperatures and dehydration, offering insights into mosses' adaptive strategies amidst climate change challenges.

## S.136.5 Bryophytes of Australian Mediterranean climate areas: Species diversity, adaptation, distribution and conservation

D. Christine Cargill¹, Josephine Milne²

1 Australian National Herbarium. Centre for Australian National Biodiversity Research (a joint venture between Parks Australia and CSIRO) Canberra. Australia. 2 Honorary Associate, National Herbarium of Victoria, Royal Botanic Gardens Victoria, South Yarra. Australia

Bryophytes that occur in the Australian Mediterranean biome have been the subject of many studies: regional checklists, taxonomic revisions, ecological studies of biocrust species and flora surveys of various locations across the biome. However, no cohesive study of the bryophytes which occur there has ever been attempted. We are presenting our observations and studies of targeted groups of bryophytes which are particularly diverse and/or particularly adapted to this biome as a first step to address this broader knowledge gap. Complex thalloid liverworts are well suited to this habitat type and the genus Riccia is the largest and most diversified group. South Africa and Australia both have over 50 species, half of the known diversity for the genus. Other families that have been successful in this biome include the Aytoniaceae, comprised of cosmopolitan species such as Reboulia, Plagiochasma and Asterella. The highly diverse genus Fossombronia has also developed adaptations to survive in extreme conditions, allowing it to thrive in Mediterranean conditions. However, it is the discovery of elusive and rare members of the Sphaerocarpales that has opened up the habitat of salt marshes, rarely visited by bryologists leading to the discovery and hitherto unknown diversity and range for these taxa. Many moss taxa are also well adapted to the extremes of this area, in particular tiny mosses such as Phascum, Stonea and Phascopsis, which thrive in biocrusts. We present the varied ways in which these bryophytes successful-