# Assessing the ecological impacts of invasive alien plants of France using adapted protocols of EICAT and EICAT+

Thomas de Solan, Aurélien Caillon, Guillaume Fried, Yohann Soubeyran, Arnaud Albert

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To cite this version:

Thomas de Solan, Aurélien Caillon, Guillaume Fried, Yohann Soubeyran, Arnaud Albert. Assessing the ecological impacts of invasive alien plants of France using adapted protocols of EICAT and EICAT+. XX International Botanical Congress, Jul 2024, Madrid, Spain. hal-04908563

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# IBC 2024
XX International Botanical Congress
**Madrid Spain**

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Book of Abstracts.
Oral presentations

July, 21st - 27th, 2024

ibcmadrid2024.com • info@ibcmadrid2024.com

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\#IBC2024
Fase20
congresos

**Title of the Work:** Title of the Work: XX International Botanical Congress IBC 2024, Spain.
Book of Abstracts. Oral presentations.

Publisher: Fase 20 Ediciones

C/ Narváez, 15- 1º Izqda.- 28009 MADRID

www.fase20.com

ISBN: ISBN: 978-84-09-63655-6

© Copyright 2024. All rights reserved.

The content of this publication may not be reproduced or transmitted by any electronic or mechanical means, including photocopying, magnetic recording, or by any information storage and retrieval system, in any form, or by any means, without prior written permission from the copyright holders.

Total or partial reproduction of this material, including images and tables, in any form—whether electronic, mechanical, by photocopy, or any other means—is strictly prohibited without express written authorization from Fase 20 Ediciones.

The editors do not accept any responsibility or legal liability for errors or omissions regarding the accuracy of the information obtained in this work. It is also assumed that the reader possesses the necessary knowledge to interpret the information provided in this text. In any case, the use of this manual cannot replace the professional judgment of the physician, who will be solely responsible for their clinical decisions.

The sale or exchange of this book for profit is strictly prohibited without the express written authorization of the Publisher.
INDEX BY SYMPOSIA

Invited Speakers 4

Symposia Session 1 Monday 20

Symposia Session 2 73

Symposia Session 3 127

Symposia Session 4 Tuesday 180

Symposia Session 5 236

Symposia Session 6 290

Symposia Session 7 Wednesday 341

Symposia Session 8 Thursday 394

Symposia Session 9 448

Symposia Session 10 502

Symposia Session 11 Friday 556

Symposia Session 12 612

Symposia Session 13 665

Symposia Session 14 Saturday 717

Symposia Session 15 773

AUTHOR INDEX BY FIRST NAME 822

SPONSORS 882
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## Invited Speakers

### Why botany? Why now?

Sandra Knapp.

Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom

### Abstract

Botany – broadly speaking, the study of algae, fungi, and plants – is a global enterprise. We depend upon the organisms for the very air we breathe, yet their survival, along with our own, is ever more in question. How should we come together to tackle this planetary emergency – the mutually reinforcing threats of climate change and biodiversity loss? Individual botanists, or individual botanical institutions, cannot tackle this alone. At this critical time in Earth's history, it is more important than ever to work together – across boundaries, institutions, and career stages – to effect change and affect our planetary future. The time for concentrating the power and influence in a few institutions is gone, and our strength lies in our diverse skills and complementary understandings of the natural world. Our goal must be to create a level playing field where all those studying algae, fungi, and plants can come together to share ideas and exchange knowledge, so we can together develop solutions both from nature but most importantly for nature. I will share some learnings from projects I have been fortunate enough to have been involved in and will bring new ideas for discussion. Today, at almost the quarter-point of this century, it is time for us to join efforts in better, more diverse, and more distributed ways to find solutions for Earth's problems so the algae, fungi, and plants upon which we all depend and that we all so love, have a secure future.

### Unraveling diversity and evolution of lichens in the genomic era

H. Thorsten Lumbsch, Todd J. Widhelm, Felix Grewe

Collections, Conservation, and Research and Grainger Bioinformatics Center, The Field Museum, Chicago, USA

### Abstract

Our understanding of the biology, evolution, and diversity of lichens has dramatically changed over the last decade. This symbiotic system has been shown to be complex. Several fungi can be involved in the symbiosis and recent studies suggest a diversity of photosynthetic partners shaped by phylogeny and ecology. In addition, bacteria and various groups of other fungi have been found to form part of the lichen holobiome. Whereas lichens were thought to be ancient organisms, molecular data suggest that the evolution of these organisms happened in parallel to land plants and there is no indication from molecular or well-preserved fossils that would support an ancient origin of the lichen symbiosis. The knowledge of the phylogeny of lichenized fungi has been revolutionized with genetic and genomic data that resulted in a classification that is dramatically different from the classifications inferred from phenotypical data. Potential reasons for the bewildering complexity of phenotypical data in lichens are discussed in the presentation.

Genomic data have also fundamentally changed our understanding of species delimitation and biogeography of lichens. In the pre-molecular era, it was assumed that ecology was the main driver shaping distri-
butional ranges with only very limited attempts to interpret observed patterns in a historical biogeographical framework. Molecular data showed that numerous species that were interpreted as having wide, inter-continental distributions belong to distinct clades, often not closely related to each other. As a consequence, historical biogeography has seen a revival in lichenology over the last decade. The genomic era is in its infancy in lichenology and some open questions will be highlighted that will be addressed with the growing number of genomic data becoming available

# Tropical tree adaptation to fire regimes and infertile soils: key for ecosystem conservation

Kaoru Kitajima¹, Hiroki Sato2, Joseph Percival1, Tomohiro Nishigaki 3, Kaoru Oshima1, Ando Rakotomamonjy1, Zo Lalaina Razafiarison4

1 Graduate School of Agriculture, Kyoto University, Kyoto, Japan

2 Graduate School of Asian and African Area Studies, Kyoto University, Kyoto, Japan

3 Japan International Research Center for Agricultural Sciences, Tsukuba, Japan

4 Faculty of Sciences, University of Antananarivo, Antananarivo, Madagascar

## Abstract

Fire is a key threat to the conservation of natural forests worldwide. In many tropical countries, especially in Africa, population increase and expansion of agricultural lands encroach the remaining frontier of pristine forests, many of which are on marginal lands that do not support sustainable agricultural production. This talk highlights the case in NW Madagascar, where we have established a 15-ha forest monitoring plot in a seasonally dry forest on infertile white sand within Ankarafantsika National Park. This forest supports a high diversity of plants and animals, including eight species of lemurs in mutualistic relationships with many tree species via frugivory-seed dispersal. The woody plant community of this forest has a high level of endemism, with dominance of evergreen species despite the prolonged dry season (April November). Fire has been a main driver of forest loss in the region, resulting in the highest rate of forest loss in Madagascar during the last three decades. During the last five years, the number and area burned increased rapidly within the national park border. We have conducted intensive analyses of ca. 3000 ha near the southern border of the park, where there were no fires until 2014. Large areas of primary forest burned in 2017, 2019, and 2021, partly due to a lack of fire adaptation in the tree community. The first burn makes a site susceptible to repeated burns, as dead woody materials above and below ground serve as fuel to support intense burns. After a few fire episodes, the dense closed-canopy forest turns into a barren desert.⁶⁾ The fire vulnerability of the primary forest in Ankarafantsika is a testimony to the importance of paying attention to variations in fire adaptation within the tropical dry forest biome to design effective conservation strategies at the landscape level.

# The science of plant conservation in a changing Mediterranean world

Thompson, John D

CEFE, Univ Montpellier, CNRS, EPHE, IRD, Montpellier, France
## Abstract

The Mediterranean region and its climate are changing, perhaps faster than we realize. As scientists, a modern challenge is to identify how species, communities, and ecosystems respond to global change and to integrate information on the underlying processes into conservation management and recommendations for political decision. This knowledge is necessary to fulfill conservation objectives. This talk will focus on different elements of contemporary ecological and evolutionary dynamics of West Mediterranean endemic plant species whose distribution patterns cover all, or a small part of, a narrow belt from northern Morocco, though eastern Spain, to mediterranean France. This distribution is a recurrent pattern in the flora of the West Mediterranean. The often narrow latitudinal range of species with this distribution pattern contrasts with a wide longitudinal range. As a result, many species occur in areas subject to different representations of the Mediterranean climate, and very often in peripherally isolated populations at the limits of their distribution. Using examples of several species with this distribution pattern I will illustrate ongoing processes of adaptation in a changing Mediterranean climate and the issues associated with conserving species that show marked variability in the their ecological niche on different spatial scales, their spatial genetic structure, and variation in their floral biology at their distribution limits. I conclude with a message concerning how our knowledge of the history of the Mediterranean region improves not only our understanding of this variability but also the reasons and values for local efforts to conserve such species, often in areas where they are rare.

# Cryptic disruptions of plant-animal mutualisms in the Anthropocene

Jordano, Pedro

Estación Biológica de Doñana, CSIC, and Dept. Plant Biology and Ecology, Univ. Sevilla, Spain

## Abstract

Pairwise plant-animal mutualistic interactions build up into mega-diverse networks involving dozens of interacting species. These mutualisms among free-living species fundamentally consist of food provisioning by plants and, their counterpart, plant propagule (pollen, seeds) movement by the animals. They are based on interaction outcomes serviced by multiple species, so that studies focusing on pairwise interactions in isolation underestimate the levels of biodiversity required to maintain multifunctional networks. Loss of biodiversity in these assemblages entails losses of key functional services that may remain cryptic, i.e., their consequences undetected well after the loss. I explore study cases documenting the extinction of mutualistic interactions, and the loss of associated services that may occur well before the partner species become extinct. In addition, also how addition of interactions (e.g., by invading species) may deconstruct native interaction networks. The combination of direct and passive interaction sampling (e.g., direct census, camera traps) and last-generation sequencing (DNA barcoding) allow to identify species-specific contributions to these mutualisms and estimate both demographic and genetic outcomes of interactions. The functionality of most generalized mutualisms among free-living species relies on complementarity of effects across a high diversity of partners. Disruptions of these mutualisms by anthropogenic drivers may remain subtle and undetected despite being pervasive in many real world ecosystems. The loss of these interactions, frequently occurring well before their plant and animal partners go extinct, generates a type of extinction debt not adequately addressed when biodiversity loss is considered.
# Evolution of floral disparity through integration of fossil and extant morphological diversity

Susana Magallón, Andrea M. López-Martínez, Maria von Balthazar, Jürg Schönenberger, Hervé Sauquet, Marion Chartier.

## Abstract

Integration of molecular data, to provide a general phylogenetic framework, and morphological data, to allow incorporation of fossils, represents a cardinal approach to investigate evolution in deep time. We assembled a morphological matrix for 1201 extant species representing all angiosperm families, and 121 well-preserved fossil flowers, and in combination with a molecular database for extant species based on exemplar representation, used it to investigate methodological issues relating to integration of extant and fossil taxa in phylogenetic estimation; divergence time estimation in a full Total Evidence approach; and estimation of the theoretical floral morphospace. Phylogenetic analyses used different optimization criteria and kinds of data to estimate relationships, as well as uncertainty in fossil placements. We found that the joint use of molecular and morphological data in a parametric context allows to recover a phylogenetic framework in agreement with molecular estimates, and fossils associated to branches in agreement with assessments based on detailed morphological comparisons. Nevertheless, uncertainty associated to fossil placements is usually high. An attempt to estimate divergence times using morphological, molecular and temporal information indicates that, while available models to integrate extant and fossil species in the same diversification process represent significant advances, there are practical difficulties with fossils for which few characters can be scored, and in the free estimation of model parameters. The theoretical morphospace of floral structure was estimated through the construction of a pairwise distance matrix among extant and fossil species, estimation of disparity, and ordination techniques to reduce dimensionality. The area of the theoretical morphospace occupied by extant and fossil species was identified, as well as how morphospace occupation has changed through time and among groups. A decrease in morphospace occupation towards the present and canalization in the of morphospace occupation among derived clades are documented, in agreement with previous independent observations.

# Integrating Plant Hydraulics and Ecological Processes Across Scales

Rafael Oliveira

Ph.D in Integrative Biology at the University of California, Berkeley. Professor at Plant Biology Department at University of Campinas, Campinas, Brazil

## Abstract

Plant hydraulics is increasingly recognized as a central link between plant biology and fields such as community and ecosystem ecology, climatology, and earth-system science. Understanding the intricate interactions between plant hydraulics and ecological processes is essential for predicting plant responses to environmental changes and managing ecosystems effectively. In this talk, I will present an overview of recent findings that demonstrate how hydraulic traits can explain species distributions across various spatial scales and influence ecological processes such as growth, demography, community assembly, and carbon dynamics,
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particularly in tropical ecosystems, which are crucial for global water and carbon cycles. Furthermore, I will discuss how plant hydraulics can offer a mechanistic explanation for vegetation resistance or vulnerability to predicted climate changes, and how this knowledge can be used to forecast future dynamics of species and communities in changing climatic conditions.

# Evolution of woody plants on the tropical coasts inferred from a full set of mangrove genomes

Shi, Suhua

School of Life Sciences, Sun Yat-sen University, Guangzhou, China.

## Abstract

Genomic studies are now poised to explore whole communities of species. The ~70 species of woody plants that anchor the coastal ecosystems of the tropics, collectively referred to as mangroves, are particularly suited to this exploration. In this study, we de novo sequenced the whole genomes of 32 mangroves, which we combined with other sequences of 30 additional species, comprising almost all mangroves globally. These community-wide genomic data will be valuable for ecology, evolution and biodiversity research. While the data revealed 27 independent origins of mangroves, the total phylogeny shows only modest increases in species number, even in coastal areas of active speciation, suggesting that mangrove extinction is common. A possible explanation for common extinction is the frequent sea-level rises and falls documented in the geological record. Indeed, near-extinctions of species with extremely small population size (N) often happened during periods of rapid SLR, as revealed by the genome-wide heterozygosity of almost all mangroves. Reduction in N has possibly been further compounded by population fragmentation and the subsequent accumulation of deleterious mutations, thus pushing mangroves even closer to extinction. The impact of the next SLR will be exacerbated by human encroachment into these mangrove habitats. Finally, in addition to possible extinctions, this talk will also cover the opposite phenomenon, namely, the patterns speciation along the tropical coasts.

# The phylogenetics of plant geography: past, present, and future

Donoghue, Michael J.

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

## Abstract

In decades past, vicariance/cladistic biogeographers emphasized the importance of discovering general biogeographic patterns and their underlying causes. Since then, however, most of the focus has been on inferring geographic movements using phylogenetic trees of individual clades. Another development has been the application of phylogenetic approaches to major biogeographic patterns of interest especially to ecologists. With the hope of revealing general patterns I have revisited and updated two papers that I published with colleagues in 2004 one on the latitudinal gradient in species richness and one on major disjunctions around the Northern Hemisphere. In both cases, I have carried out a systematic review of progress based on plant studies published
over the past twenty years. Although a number of hypotheses have been well supported over this time period, and others can now safely be set aside, there are still substantial gaps in our understanding, and a need for further coordination of efforts and methodological improvements. I will present the results of these studies while highlighting the value of syntheses that draw upon multiple primary analyses. These observations also highlight improvements that can be made in studying individual clades. To illustrate many of the central points I will make special reference to our ongoing studies of Viburnum biogeography

# Delving into South African medicinal plant treasures

Nokwanda Makunga

Department of Botany and Zoology, Faculty of Science, Stellenbosch University, 7600

## Abstract

South Africa has an incredible floral heritage that has a strong history of being important to local people for diverse range of ethno-cultural uses and those plants that are exploited for medicinal purposes have also encouraged entry of new natural products into global markets. There are thus unique opportunities that are presented by interfacing various biotechnologies to study medicinal plants of South Africa, more especially with the intention of generating socioeconomic benefits. This talk will touch on historical aspects linked to the exploitation of medicinal plant resources and define the importance of indigenous knowledge and medicinal plants when interfaced with biotechnology in relation to the bioeconomy strategy of South Africa. Using specific examples from medicinal plants that occur in the greater Cape Floristic Region, various studies are thus presented and summarized. The application of LC-MS-based metabolomics technologies has been useful as a qualitative tool, enabling fast detection and a reliable method of studying mesembrine alkaloids produced by Sceletium tortuosum (Aizoaceae). Microplant and callus cultures also accumulate a wide array of these alkaloids. By applying both non-targeted and targeted metabolomics, influence of varied micro environments on the phytochemistry and cytotoxicity of Sutherlandia frutescens (Fabaceae) plants is evident. Proteomic analyses under stressed conditions provided new insights into the impacts of salinity and drought stress with respect to both central and specialized metabolism of S. frutescens. Combining microsatellite and chemometric analyses confirmed biogeographical-based genetic structure amongst populations of Dodonaea viscosa (Sapindaceae) and Aspalathus linearis (Fabaceae), pinpointing chemotype(s) that may fit domestication and industrialization for the production of phytopharmaceutics. These studies illustrate the exciting potential of medical plants of South Africa that still remain chemically under explored.

# Distyly as a model for studying convergent evolution – insights from genomic studies

Tanja Slotte

Professor in Ecological Genomics, Department of Ecology, Environment and Plant Sciences, Stockholm University, Stockholm, Sweden

## Abstract

Recent advances in sequencing technologies, in combination with powerful new simulation and analysis tools, have opened up a wealth of new possibilities to address classic evolutionary questions in non-model
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systems. In this talk, I will highlight how we are harnessing this development to study the evolution of plant mating systems, with a focus on the evolution of a balanced floral polymorphism termed distyly.

Distyly is a floral polymorphism governed by a supergene, that has evolved convergently in multiple angiosperm lineages. Distyly promotes outcrossing and efficient pollen transfer via the presence of two floral morphs that differ reciprocally in the positions of their male and female sexual organs. These morph differences are usually coupled with a heteromorphic self-incompatibility system that reduces self- and intra-morph fertilization. Loss of distyly has also occurred very frequently, often in association with convergent evolution of floral selfing syndromes. Studying the evolution and loss of distyly therefore offers an opportunity to address classic questions on the genetic basis of convergent phenotypic evolution, supergene breakdown and the genomic consequences of selfing.

Here, I will present our work on these topics in Linum, a non-model plant system that is exceptionally well suited for this purpose. Harnessing new high-quality genome assemblies, we have identified and characterized the distyly supergene in Linum at the molecular level, tested theory on genetic causes of loss of distyly, and documented pervasive population genomic consequences of this shift. Genomic studies of distyly in Linum and other systems have revealed striking similarities in hemizygous genetic architectures and molecular evolution of independently evolved distyly supergenes and provided a foundation for further characterization of the genetic underpinnings and genomic consequences of convergent evolution and loss of distyly. This body of work demonstrates the power of genomic analyses in non-model systems to shed new light on classic evolutionary questions.

# Biodiversity and ecosystem function in
subtropical forest

Ma, Keping

Institute of Botany, Chinese Academy of Sciences

## Abstract

In order to explore the relationship between biodiversity and ecosystem functioning such as primary productivity in forest ecosystem, a large manipulation experiment was established in subtropical China in 2009 and 2010. Based on the experiment platform, German, Swiss and Chinese scientists worked together and published more than 200 papers. The key findings are as follows. 1. After 8 years, 16-species mixtures had accumulated over twice the amount of carbon found in average monocultures and similar amounts as those of two commercial monocultures. Species richness effects were strongly associated with functional and phylogenetic diversity. 2. Effects of functional diversity (FD) values on productivity became larger than effects of community-weighted mean (CWM) values after 7 years of forest development and that the FD values also became more reliable predictors of productivity than the CWM values. In contrast to CWM, FD values consistently increased productivity across ten different species-pool subsets. 3. Tree species richness improved community stability by increasing asynchrony. That is, at higher species richness, interannual variation in productivity among tree species buffered the community against stress-related productivity declines. This effect was mediated by the diversity of species hydraulic traits regarding drought tolerance and stomatal control. 4. Effects of experimentally increased tree species richness were consistently positive for species richness and abundance of herbivores, predators and parasitoids. Richness effects decreased as trophic levels increased for species richness and abundance of all trophic groups. Multitrophic species richness and abundance of arthropods were important mediators of plant diversity effects on tree productivity. These long-term studies emphasize the importance of planting functionally diverse, mixed-species for the restoration and reforestation projects to improve ecosystem functioning, maintain multitrophic diversity and mitigate climate change.
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# Pollination ecotypes: niche-driven floral evolution below the species level

Steven D. Johnson

DST-NRF SARChI chair in Evolutionary Biology. Associate editor of Proceedings of the Royal Society B and Biological Journal of the Linnean Society.

## Abstract:

Ecological niche shifts are a key driver of phenotypic divergence among populations and can also contribute to reproductive isolation. While patterns of historical ecological shifts and the associated trait-environment correlations are well-documented at the macroevolutionary level, the process is initiated below the species level, as shown originally by Turesson, Clausen and others in their seminal studies of edaphic and climate ecotypes in plants. Pollinators are critical for the establishment and persistence of populations of the majority of plant species, thereby forming part of their fundamental niche. Novel floral trait combinations allow plants to exploit particular pollinator niches in local habitats and evolve primarily through sexual selection due to their effect on plant mating success. Shifts in pollinator niches among plant populations lead to the formation of "pollination ecotypes". Work on pollination ecotypes has revealed that their origin is best explained by contemporaneous ecological gradients arising from the mosaic-like spatio-temporal distribution of pollinators. I discuss case studies on the evolution of pollination ecotypes and use these to identify a number of challenging practical and conceptual issues. These include the paucity of reliable natural history data, difficulty of knowing when ecotypes should be considered species, reconstructing the number and direction of intraspecific shifts, and distinguishing between extinctions and isolating barriers as explanations for phenotypic and genetic discontinuities among population clusters.

# The Changing Face of Herbarium Collections

Pamela S. Soltis

## Abstract

Herbaria have long provided the foundation for our understanding of plant diversity and distributions. Beyond their fundamental role in taxonomy, herbarium specimens, like other natural history collections, reflect a history of unanticipated uses. Although collected to document plant occurrences, the world's 400,000,000 herbarium specimens, spanning space and time, provide unparalleled resources for plant science and are indeed undergoing a renaissance in their use. Preserved plant tissues are reservoirs of genetic information that can be used to reconstruct the evolutionary history of plants. The resulting family tree provides a foundation for basic biology and for solving many societal problems, such as food security and the search for new medicines. Digitized herbarium specimens also offer new research opportunities, particularly in relation to climate change and conservation. These herbarium resources provide novel opportunities for mobilizing and integrating massive amounts of information from organismal biology, ecology, genetics, climatology, and other disciplines. As the number of online records continues to grow, these data provide excellent information on species distributions, changes in distributions over time, phenology, and a host of traits. Integration of information from specimen records with phylogenies, climate data, and other resources enables new questions to be addressed while also providing new perspectives on longstanding questions in ecology and evolutionary biology. Through a series of case studies, I will illustrate some of the many uses to which herbarium specimens and their digital data are currently being applied. These case studies link specimen data and related heterogeneous data sources and show how high-throughput molecular analysis and artificial intelligence can be applied to centuries-old herbarium specimens to address a range of evolutionary, ecological, and societal problems.
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# Exploring the underground: unraveling the complex interplay of biodiversity and disease in plant communities

Liesje Mommer

Wageningen University, Wageningen, the Netherlands

## Abstract

Biodiversity loss is a significant driver of disease risk in ecosystems, yet the intricate relationships between diversity and disease remain poorly understood. This keynote lecture will delve into the knowledge frontiers of this field.

Traditionally, plant ecologists have explained diversity-disease relationships by distinguishing between host and non-host species. When biodiversity loss increases the density of hosts compared to non-hosts, disease risk increases, and vice versa. However, focusing on this host-non-host dichotomy overlooks the substantial variation in 'host quality'—a measure of a host's contribution to parasite fitness.

To illustrate this concept, I will present findings from a temperate grassland biodiversity experiment involving 16 plant species (forbs and grasses). Using Illumina MiSeq amplicon sequencing, we assessed the abundance of root-associated fungal pathogens in plant monocultures and four-species mixtures. Our study identified three major fungal pathogens with differing host ranges: Paraphoma chrysanthemicola, associated with forb species of the Asteraceae family; Slopeiomyces cylindrosporus, associated with grass species; and Rhizoctonia solani, affecting both forb and grass species. We discovered that plant community composition rather than plant species richness per se significantly influences pathogen relative abundance. The presence of specific high quality host species appeared key.

Another concept that may modify diversity-disease relationships belowground in the presence of arbuscular mycorrhizal fungi. These organisms are form symbiotic relationships with 80% of plant species and are known to enhance nutrient uptake in plants. I will present preliminary evidence that these mycorrhizal fungi modify plant-pathogen interactions in grassland systems.

Our findings highlight the intricate interplay between plant community properties, host-pathogen and host-symbiont interactions and their vital role for understanding the impacts on overall grassland productivity and ecosystem health. Be invited to journey underground to uncover the hidden dynamics of biodiversity and disease in plant communities.

# Into the wild: addressing plant extinction through population reintroductions

Gallagher, Rachael V.

1 Hawkesbury Institute for the Environment, Western Sydney University, Richmond, NSW, Australia

## Abstract

Over the last three decades, vast international resources have been committed to conservation seed banking, to great success. Seed bank collections have grown rapidly since the millennium and seeds and plant germplasm are now routinely stored to safeguard species against extinction. Yet seed banks were never intended to be a final stop along the road to recovery for plant species. Currently, when we do reintroduce or translocate plant populations, failure rates associated with their establishment can be unacceptably high. We also often do not know
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where to situate new plant populations relative to their historical distributions, given immense changes in climate and land use. New science is therefore needed to improve and intensify the strategic reintroduction of plant populations to the landscape. In this talk, I will outline gaps in ecological and conservation knowledge that hamper current reintroduction success, explore solutions, and demonstrate how we might use emerging financial market mechanisms to fund initiatives that reduce plant extinction risk. I hope to inspire new thinking about how we can thoughtfully design initiatives that get more plant species out of the fridge and into the wild across the world.

# Bringing ecophysiology closer to plants from extreme environments and diverse phylogenetic groups

Flexas Sans, Jaume

INAGEA - Universitat de les Illes Balears, Palma, Spain

## Abstract

Plant (eco) physiology deals with the physiological responses of plants to environmental conditions. Within this definition, it should deal with any kind of plant and environment. However, perhaps because this discipline often requires the use of electronic devices and heavy equipment, most studies deal with a limited number of Angiosperms, especially crops. Studies are most often conducted under tropical, temperate and Mediterranean conditions, with scarce studies in more extreme environments such as high elevation mountains, hot and cold deserts.

Here I will show examples on how my research group has recently expanded along the land plant phylogeny and towards climatically extreme ecosystems. First, I will show how photosynthetic capacity changes along the land plant phylogeny, and its underlying physiological, anatomical and biochemical causes. Second, I will present some home-made, relatively portable techniques, for assessing the short-term tolerance of plant photosynthetic organs to dehydration, low and high temperature, and excess visible and UV-B radiation, and how the use of these devices is allowing us to test for the trade-off between photosynthetic capacity and abiotic stress tolerance across the entire land plant phylogeny and along the entire spectrum of Earth biomes.

Finally, I will focus on two specific studies in some extreme environments. In the first, I will show how well standardized, portable and commercially available ecophysiological tools allowed for a diagnose of different species physiological status without the need for a previous knowledge of each species optimal status along a Chinese transect, from the Xinjiang steppes through the Taklamakan desert to the Tibet Plateau. In the second, I will deal with the particular case of giant cushion Azorella species along the Chilean Andes Range, and discuss how its particular anatomy generates a very contrasted microclimate that may allow these species to establish and, perhaps, help them to facilitate other species.

# The evolution of extant South American tropical Biomes

Jaramillo, Carlos

Smithsonian Tropical Research Institute, Panama

## Abstract

When and why South American tropical biomes developed? Tropical vegetation had a complete turnover from non-angiosperm dominance at the onset of the Cretaceous to angiosperm dominance nowadays. Cretaceous tropical biomes do not have extant equivalents; lowland forests were dominated mainly by gym-
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nosperms and ferns lacking a closed canopy. This structure was transformed following the Cretaceous Pa-
leogene meteorite event. The extant lowland tropical rainforests first developed at the onset of the Cenozoic
with a multistratified forest, an angiosperm-dominated closed canopy, and abundant legumes. Cenozo-
ic rainforest diversity has increased during warmings and decreased during coolings. Tropical dry forests
emerged at least by the late Eocene, whereas other drier Neotropical biomes are much younger.

Reimagining arid zone ecology in a
changing world: is rainfall all that matters?

Austin, A.

IFEVA-CONICET, University of Buenos Aires, Buenos Aires, Argentina

Abstract

While forests have received intense attention regarding their role in the global carbon cycle, the drier parts of the world, constituting over 40% of the terrestrial land surface, are often overlooked as unproductive and unresponsive to human-induced global change. In contrast to forest ecosystems, these arid regions present unique conditions, encompassing a matrix of climate, land use, and plant-soil interactions that suggest complex responses to changing environmental conditions, which we are only beginning to comprehend. Our research specifically focuses on understanding how alternative climatic controls, such as exposure to solar radiation, temperature, and the seasonality of rainfall, exert substantial impacts on carbon turnover and storage in arid land ecosystems. In contrast to a singular focus on rainfall as the primary determinant of ecosystem functioning in arid lands, our findings highlight the significant, positive effect of solar radiation exposure on carbon emissions and subsequent microbial turnover. By shedding light on these often-neglected arid land ecosystems and their specific controls, we may provide more predictive power for effective management and conservation amidst ongoing climate and land use changes.

Machine learning and new inference
algorithms: expanding what is possible
in evolutionary biology and phylogenetic
analysis

Sanmartín, Isabel

Real Jardín Botánico, CSIC, Madrid, Spain

Abstract

Investigators across different fields of evolutionary biology are increasingly turning their attention to more integrative
and complex inference models. This trend is fueled partly by the emergence of next generation sequencing technol-
ogies, scaling up the number of genes, individuals, and species sampled, and partly by the rapid expansion of bio-
diversity, climatic and biogeographic databases at the local, regional and community scales. This massive increase
in the quantity of data available has opened up a range of new questions but also unexpected challenges that
require the development of more sophisticated approaches to data modeling. For example, a standing challenge is
how to hybridize lineage-based, individual-based, and gene-based models to adress all aspects of biodiversity loss
(richness, abundance, genetic diversity) in order to explain the biotic impacts of climate change. The COVID-19 pan-
<figure><img src="image_16.png" /></figure>

demic introduced the need for models that are computationally efficient and can analyze in a short time thousands of sequences differing only in a few nucleotides. Statistical models in evolutionary biology are typically built around stochastic parameters interconnected by rigid dependencies, but there is a growing demand for models that allow hidden, unobserved components. Moreover, some commonly employed models for evolutionary inference face challenges related to non-identifiability, where distinct model parameters induce the same probability distributions and are structurally or practically indistinguishable. Phylogenetic inference itself is classified as a non-polynomial time problem, meaning that the number of steps needed to solve a problem grows exponentially with the size of the problem. Here, I review some possible solutions and paths forward in relation to these challenges, including the advent of online Bayesian phylogenetics, the popularization of (deep) machine learning approaches, and the introduction of probabilistic programming languages that promise to speed up analyses and tackle more complex biological questions in fields as diverse as phylogenetics, epidemiology, community ecology and phylogeography.

# What do we know about mutualism? A thirty-year retrospective on a very young field

Judith L. Bronstein

University of Arizona, Tucson, USA

## Abstract

The study of mutualism (interspecific, mutually beneficial interactions) is a surprisingly young subfield of ecology and evolutionary biology. Building on centuries of natural history knowledge, it began to coalesce in the 1970s; key patterns and processes only started to attract intensive attention in the 1990s. I will start by briefly summarizing the best-understood mutualisms in which plants are involved, interactions that have fundamentally shaped plant biology: nutritional mutualisms (including symbioses with mycorrhizal fungi and Rhizobium bacteria), transportation mutualisms (biotic pollination and seed dispersal) and protection mutualisms (involving ants that attack herbivores). I will then offer a capsule history of the study of mutualism as a whole, focusing on its origins and subsequent development. I will highlight key research questions that have emerged in recent years, as well as some main findings to date from each of them. Finally, I will lay out questions likely to drive mutualism research in the coming years. My approach throughout is to highlight features uniting mutualisms that differ radically in natural history, as well as features that make mutualisms similar to and different from other types of interspecific interaction. We are only now beginning to recognize what we need to know about these critically important interactions to help secure the future of plants and the larger systems in which they occur. As a whole, it is an exciting time for this very young field.

# Diploidization, Polyploidy, and the Evolution of Plant Diversity

Barker, Michael S.

Department of Ecology & Evolutionary Biology University of Arizona

## Abstract

Polyploidy, or whole-genome duplication, is widely recognized as an important force in the evolution of plants. Ancient genome duplications have driven the re-organization of plant genomes and nearly 1/3 of contemporary flowering plants are of recent polyploid origin. Despite the recognition that polyploidy has been important
<figure><img src="image_17.png" /></figure>

across plant evolution, there are still many unresolved issues surrounding polyploidy and its role in evolution. Among these issues is reconciling the apparent success of ancient polyploid species with observations that contemporary polyploids have much lower net diversification rates than their diploid relatives. Here, I present new analyses of polyploidy across the plant phylogeny and explore the impact of species tree topology on WGD inferences. Using these new inferences, I will present phylogenetic simulations that explore the legacy of ancient WGDs with different diversification dynamics of polyploid and diploid species. We find that multiple rounds of polyploidy occur in the ancestry of lineages when diploid species have higher net diversification rates than polyploid species. This occurs in simulations using arbitrary rates as well as those using parameters derived from empirical analyses. Using empirical estimates of diploid and polyploid diversification rates from plants and estimates of the rate of polyploid speciation from diploids, our simulations indicate an average of 3.5 rounds of WGD in flowering plant species. This is consistent with observations from phylogenomic studies and suggests that the results of many independent lines of research, from empirical estimates of diversification rates to inferences of WGDs, broadly agree with each other and are likely generally accurate. Our simulations also indicate that the rate of diploidization is an important driver of the evolutionary dynamics of polyploidy and may explain variation in the numbers of ancient WGDs inferred in different lineages of eukaryotes. Overall, our simulations reconcile the seemingly contradictory observations of low polyploid diversification rates with the repeated rounds of WGDs that have been inferred throughout the evolutionary history of flowering plants.

# One third backwards! The varied ways plants have already changed in response to climate change and the implications for plant communities and conservation.

Moles, Angela

Evolution & Ecology Research Centre, UNSW Sydney, NSW 2052, AUSTRALIA

## Abstract

Plant species are already responding to climate change by undergoing rapid evolution, altering their phe-nology, and undergoing remarkably rapid shifts in distribution. I will review current understanding of these changes, highlighting the fact that different species are changing in different ways and at different rates, and that not all of the responses are as we would expect. Most strikingly, a third of species seem to be shifting their distributions downhill or toward the equator rather than shifting to cooler areas. This variation in species responses to climate change means that broad change in plant communities seems inevitable. I will finish by discussing implications of all this change for conservation.

# State of the World's Plants and Fungi

Alexandre Antonelli

Director of Science at the Royal Botanic Gardens, Kew, UK. Professor of Biodiversity and Systematic at the University of Gothenburg, Sweden, and Visiting Professor at the University of Oxford.

## Abstract

The race is on to scientifically describe the world's plants and fungi, so we can better understand, protect, and sustainably use biodiversity. But how do we accelerate scientific research and conservation, while strength-
ening local capacity and equity? These are questions at the heart of Kew's State of the World's Plants and Fungi reports, which bring together global scientific expertise, large data sets and practical experiences to synthesise current knowledge on plants and fungi. In this talk I will provide a short overview of the themes published so far and then take a deep dive into the latest report focused on how our communities could collectively work to fill in the most critical knowledge gaps - through innovative and sometimes unexpected approaches.

# Save the male! Reflections on sex and gender in flowers and flowering plants

Pannell, John R.¹

Department of Ecology and Evolution, University of Lausanne, 1015 Lausanne, Switzerland

## Abstract

Plant sexual reproduction involves strategic decisions in the context of variation in sex and gender. These decisions include who to mate with and how often (the mating system), whether and when to mate as a male or a female partner, and how much resource to allocate to each sexual function (the sex allocation). All these decisions may be subject to tradeoffs and will often reflect only a partial resolution to genetic and sexual conflicts. The wonderful diversity of flowers illustrates the numerous ways in which conflict resolution has been achieved. In this presentation, I will address the complexity of mating systems and sex allocation, especially with a view to the benefits and costs of being male. How does selection on the mating system interact with selection on sex allocation? How well do we understand the costs and benefits of being both male and female? Can we satisfactorily address Charles Darwin's difficulty in understanding why hermaphrodite plants should ever have been rendered dioecious? To what extent can current theory explain the distribution of combined versus separate sexes among individuals in a population? Can the same theory also help to explain why female flowers are so rarer than male flowers on plants with bisexual flowers, whereas females are so much more likely than males to co-occur with hermaphrodites in a population? To address these questions, I will reflect on variation in sex and gender in plants, drawing upon: insights gained from observations of plant and floral sex ratios; evidence of sexual selection in wild and experimental populations; observations of rapid transitions between combined and separate sexes under experimental evolution; the association between mating pair identity and sex allocation strategies; and the phylogenetic distributions of reproductive and other traits.

# Morphological and genomic consistency for 20 million years

Elizabeth A. Kellogg

Robert E. King Distinguished Investigator at the Donald Danforth Plant Science Center in St. Louis, Missouri. Member of the National Academy of Sciences of the U.S. and is an Académica (Fellow) of the Academia Nacional de Ciencias, Argentina.

## Abstract

Nearly one fifth of the world's land area is inhabited by wild grasses of a single clade, the tribe Andropogoneae, which originated about 20 million years ago in the early Miocene. A cultivated member of this same tribe, maize, now replaces wild grasses in many areas, dominating the landscape and the world's economy. The
<figure><img src="image_18.png" /></figure>

tribe is characterized by spikelets borne in pairs, with the spikelet pair as the dispersal unit. Thanks to this type of dispersal unit, the tribe spread throughout the Old World, diversifying into more than 1000 species. Much later and coincident with the well-documented global grassland expansion, members of the tribe arrived in the Americas as the result of several independent dispersal events. Aerodynamic properties of the spikelet pairs are consistent with long-distance dispersal by wind for most, but not all, species. We have generated de novo genome sequences for about 40% of the 1200 species in the tribe using a combination of long-read and short-read sequencing. The 33 new long-read genomes are high quality as shown by several measures of completeness and contiguity. Genome alignments indicate collinearity of genomes, with the ancestral haploid number and architecture of 10 chromosomes being preserved even through polyploidization events. We have developed data and tools to improve annotation of the maize genome, inferred the chromosomal evolution of maize, investigated amplification of transposable elements, and begun to identify conserved regulatory sequences. Together these resources can enhance breeding for environmental change in the crops and illuminate the history of wild species.

# Beyond regional flora's: what do we know about the ecology and evolution of the African flora?

Muasya, Muthama

University of Cape Town, Cape Town, South Africa

## Abstract

Studies on the diversity of African plants have lagged. Efforts to document the floral wealth have spanned nearly two centuries, with oldest treatments in Flora Capensis dating back to 1860s, but the tropical regions of the continent have only been more recently studied, and knowledge gaps remain. Overwhelmingly, there is good coverage in documenting species occurrence and taxonomy, and indepth studies have focused on crops and other economically important taxa. Ecological and evolutionary studies have sought to explain the origin and processes maintaining the distinctive assemblages, in part driven by questions on the origin of hominins and other iconic fauna. The continent has a rich diversity of vascular plants, with over 65,000 species, distributed in five major vegetation types (lowland forests, savannas, temperate, montane and alpine, and desert). Africa was covered in lowland forests at about 60 million years ago, gradually opening into savannas since the Eocene, and fragmented further by the uplifting and volcanism during the mid Miocene. Africa position, straddling the equator and its earlier emergence of open habitats, drove origin of earliest savanna lineages, in particular C4 graminoids and associated fauna, which dispersed into other continents as open habitats evolved. The temperate (Cape) flora, together with the montane and alpine flora, have assembled more recently, seeded from the savanna and other temperate floras, though the directionality of dispersal between oceanic and sky islands is contentious. Most intriguing is the assembly of the arid flora, where there is evidence for both antiquity and recent origin, and hypotheses on an African arid corridor persists. Heterogeneity in topography, soils and climate, together with fire and biotic interactions, drives the wide species riches and functional diversity. With a focus on the Cenozoic, the current knowledge and gaps on the ecology and evolution of African flora will be interrogated.
<figure><img src="image_19.png" /></figure>

# Evolution and development of iridescent petals

Glover, Beverly J.

University of Cambridge, UK

## Abstract

The enormous diversity of flower colour is largely the result of modifications to and combinations of chemical pigments. But some plants also use physics to modify the colour of their flowers. In this talk I will describe the way that diffractions gratings, formed from folds of the epidermal cuticle, can produce structural colour in flowers. I will summarise our recent work analysing the distribution and diversity of this trait in the angiosperms, its function in pollinator attraction and the developmental programmes underlying the formation of floral iridescence.
**S.1 MERGING EVOLUTIONARY APPROACHES WITH ETHNOBOTANICAL KNOWLEDGE. SESSION 1**

**S.1.1 The potential of phylogenies to safeguard human well-being**

Rafael Molina-Venegas¹,²

1 Department of Ecology, Faculty of Science, Universidad Autónoma de Madrid, Spain. 2 Biodiversity and Global Change Research Center, Universidad Autónoma de Madrid, Spain.

The idea that conserving evolutionary history is more efficient than species-based approaches for capturing biodiversity benefits to people has remained largely theoretical since the early 1990s. These benefits may not necessarily be currently known or realized benefits but 'option values', that is, benefits that are yet to be discovered or demanded by future generations, which hampers assessing the long-standing hypothesis that preserving the main branches of the Tree of Life (i.e., high levels of phylogenetic diversity) would help to safeguard human well-being. However, it is reasonable to assert that known benefits today were at one point in human history unknown future options. Hence, proving that current benefits of biodiversity (i.e., the 'option values' of past generations) are to a great extent provided by subsets of taxa that encapsulate high levels of phylogenetic diversity could serve as a proof of concept on the relationship between evolutionary history and future options. Adhering to this reasoning, some recent studies have begun to yield empirical evidence supporting the theoretical background, thus opening an exciting avenue of research to further investigate the complex relationship between biodiversity and human well-being.

**S.1.2 The hot node analysis incompletely explains the position of phylogenetic structure in taxon samples**

Darío Atienza-Barthelemy¹, Manuel J.

Macía¹,², Rafael Molina-Venegas²,³

1 Department of Biology (Botany), Faculty of Science, Universidad Autónoma de Madrid, Spain. 2 Biodiversity and Global Change Research Center, Universidad Autónoma de Madrid, Spain. 3 Department of Ecology, Faculty of Science, Universidad Autónoma de Madrid, Spain.

Advances in phylogenetics have provided a revolutionary way to address ethnobotanical data, generally based on calculating the phylogenetic structure of a set of medicinal plants, and identifying the clades showing significant overabundance of such plants (i.e., hot nodes). However, this emerging discipline is in need for more quantitative evidence, and the common assumption that hot nodes represent the main clades responsible for phylogenetic structure remains to be tested. Here, we use the most extended metrics on the field and a comprehensive database (1,803 medicinal species) to calculate the phylogenetic structure of the medicinal plants of Ecuador. Specifically, we (1) conduct the analyses on both species- and genus-level phylogenies to assess the influence of the phylogenetic grain, and (2) evaluate whether hot nodes can tackle the position of phylogenetic structure using a jack-knifing procedure to identify clades significantly influencing phylogenetic distance-based metrics. Phylogenetic structure was found to be strongly dependent on taxonomic level and the identified hot nodes incompletely matched those with a significant contribution to phylogenetic structure metrics. Significant information may be missed whenever the influence of the taxonomic level is not evaluated, and the position of phylogenetic structure cannot be fully tackled.
<figure><img src="image_20.png" /></figure>

led by the hot nodes. We suggest combining different phylogenetic grains and methods to identify highly influencing clades before drawing any conclusion that may misguide the search for plants with bioactive potential.

## S.1.3 A new phylogenetic analysis pipeline to support the identification and the prioritisation of plants with medicinal potential

Giovanni Zecca¹,², Elisa Toini¹,², Massimo Labra¹,², Fabrizio Grassi¹,²

1 Department of Biotechnology and Biosciences, University of Milano-Bicocca, Milan, Italy. 2 NBFC, National Biodiversity Future Center, Palermo, Italy.

Recent studies have proposed that phylogenetic methods can be effective tools for exploiting the untapped potential of medicinal plants. Under the assumption that medicinal properties tend to be phylogenetically clustered, phylogenies can be used to narrow down the relevant species for bioprospecting, increase the discovery rate and reduce the time and costs involved. Nonetheless, some restrictions exist. By integrating ethnobotanical knowledge with phylogenetic methods, it has often been necessary to group plants into macro-categories of therapeutic effects, limiting the potential of the approach. In many studies "hot nodes" corresponding to lineages overrepresented in medicinal taxa were identified. However, as larger phylogenies and databases become available, the possibility of huge lineages being identified by "hot nodes" is also likely to increase, once again posing a prioritization problem. To tackle these issues, we propose a new analysis pipeline, mixing existing and new developed methods. Briefly, our workflow is as follows: 1) using selected databases that integrate traditional medicine data with growing molecular information, species linked to specific diseases or specific biological effects are obtained; 2) different methods to estimate the phylogenetic clustering of taxa are applied using a reference phylogeny; 3) where a phylogenetic signal has emerged, "hot nodes" are repeatedly estimated to assess their robustness; 4) in case of large lineages subtended by stable "hot nodes", a (relative) probability of being linked to the disease or biological effect of interest is calculated for each taxa; 5) results are summarised and graphically displayed

to combine different information. To test our pipeline data related to 10 diseases and 10 biological effects were obtained from two different public databases and were analysed together with a published phylogeny including ~ 30,000 species. Our results show the effectiveness of the approach suggesting that it will be useful in the bioprospecting of new potential medicinal plants.

## S.1.4 Exploring ethnobotanical knowledge dynamics across time scales: the contribution of phylogenetic comparative methods

Irene Teixidor-Toneu¹,²*, Anneleen Kool², Simon J. Greenhill³,⁴, Karoline Kjesrud⁵, Jade J. Sandstedt², Fiona M. Jordan⁶

1 Mediterranean Institute for Biodiversity and Ecology, French National Research Institute for Sustainable Development, Marseille, France. 2 Natural History Museum, University of Oslo, Oslo, Norway. 3 Australian National University College of Arts and Social Sciences, ARC Centre of Excellence for the Dynamics of Language, Canberra, Australia. 4 Department of Linguistic and Cultural Evolution, Max-Planck-Institut fur Menschheitsgeschichte, Jena, Germany. 5 Museum of Cultural History, University of Oslo, Oslo, Norway. 6 Department of Anthropology and Archaeology, University of Bristol, Bristol, UK.

Knowledge of plants is dynamic and can change at rapid time scales. Grasping past human-plant relations is largely concomitant to having historical and archaeological evidence. Yet, apart from major crops, this is uncommon for most social-ecological systems and time periods. This presentation outlines a multi-disciplinary triangulation method that combines inferences of ancestral plant uses by a cultural evolutionary approach with historical, linguistic, and ethnobotanical evidence (Teixidor-Toneu et al 2021). Phylogenetic comparative methods that use linguistic phylogenies representing people's historical relations are applied to ethnobotanical data. Results fill gaps of evidence and link data sources for the study of past human-plant relations. The triangulation method is illustrated by taking the Nordic countries as a case study, as extensive historical collections of various plant use records are very limited until the 1700s and there is also
<figure><img src="image_21.png" /></figure>

limited direct evidence of plant use from archaeo-
logical contexts. A language tree models the
population history of the Nordic region and allows
to computationally infer ancestral plant uses
from recent ethnographic data. We evidence a
wealth of Viking-age plant uses, especially for
medicine and food. All plants for which ancestral
uses were inferred by phylogenetic compara-
tive methods had vernacular names with shared
North Germanic word roots, confirming the past
knowledge and importance of these species. We
inferred the ancestral use of a diversity of spe-
cies (such as *Angelica archangelica* or *Junipe-
rus communis*) and uses (wild foods, beer, and
mead brewing) whose cultural history had been
overlooked in botanical or archaeological stud-
ies. This multi-disciplinary triangulation method
evidences that while most plant use knowledge
may quickly arise and fade through time, some
human-plant relations persist through centuries
shaping both culture and the environment.

S.1.5 INDIGENOMICS: Linking
indigenous knowledge with
metabolomics to assess the
chemical traits underpinning
human-plant interactions

Rodrigo Cámara-Leret¹, Luiz L. Saldanha¹,
Kim Castro¹, Juan C. Copete¹, Lauren Raz²,
Uldarico Matapí³

1 Department of Systematic & Evolutionary Botany, University of Zurich. 2 Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Bogotá, Colombia. 3 Independent researcher, Bogotá, Colombia.

INDIGENOMICS is a Swiss National Science Founda-
tion Starting Grant (2023-2028) that aims to inte-
grate Indigenous ways of knowing with science to
address a long-standing question about the chem-
ical traits that underpin human-plant interactions.
Co-developed with the Indigenous Matapí of the
Colombian Amazon, we are testing hypotheses that
link Matapí and western ecological knowledge to
better understand how plant chemistry influences
the use, management, and conservation of Amaz-
zonian forests. Coupling metabolomics and Indig-
enous knowledge, we unravel the phytochemical
space used by Indigenous communities to predict
how much chemodiversity may vanish as global
change accelerates across Amazonia. Supporting

Indigenous elders and their apprentices, INDIG-
NOMICS boosts indigenous-led research to preserve
the endangered knowledge of the Matapí— thus
strengthening Indigenous contributions to science
while creating opportunities that reinforce cultural
heritage.

S.1.6 Botanicultural
macroevolution? Theory
and tools for investigating
cross-cultural uses of plants

Fiona M. Jordan¹, Julie Hawkins², Jamie
Thompson³

1. Anthropology & Archaeology, University of Bristol, UK. 2 Ecology & Evolutionary Biology, University of Reading, UK. 3 Life Sciences, University of Bath, UK.

Evolutionary approaches to human culture and
language are now commonplace in anthropoloy-
gy, including the use of comparative phylogenetic
methods to understand cultural macroevolution.
Using linguistic phylogenies to model human pop-
ulation history, it is possible to infer the dynamics
of change in ethnographic, archaeological, mate-
rial, linguistic and cognitive phenomena. Ancestral
features of cultural groups can be estimated, and
both transformational sequences of change as well
as coevolutionary hypotheses about culture and
environment can be robustly tested while account-
ing for historical autocorrelation between popula-
tions. Language families spanning the globe and
human lifeways are now represented by computa-
tionally inferred language phylogenies, and the ma-
turity of the field has accelerated with large-scale
cross-cultural databases and data collection sur-
veys. To further promote ethnobotanical research in
this arena, we present a selection of key studies that
investigate botanical resource use. Our recent study
of Polynesian medicinal plant traditions explores the
roles for adaptation and ancestral knowledge in
plant-based healthcare at pre-historic timescales.
We test the extent to which (a) the new floristic en-
vironments encountered, (b) cultural ancestry or (c)
geographic proximity predict the composition of
ethnopharmacopoeias in seven Oceanic ethnolin-
guistic groups. This small study highlights the need
for large-scale ethnobotanic databases in order to
progress future work, while highlighting the chal-
lenges of collating and synthesising traditional eco-
logical knowledge in ways that are sensitive to the
CARE principles of indigenous data management.
<figure><img src="image_22.png" /></figure>

**S.2 THE ANGIOSPERM REPRODUCTIVE**
**SPECTRUM FROM FLOWERS TO SEEDS.**
**SESSION 1**

**S.2.1 The eFLOWER project:**
**what have we done, where**
**are we, and where are we**
**heading now?**

Hervé Sauquet¹,²,³, Andrea M. López-Martínez⁴, Maria von Balthazar⁵, Marion Chartier⁵, Julian Herting¹, Santiago Ramírez-Barahona⁴, Susana Magallón⁴, Jürg Schönenberger⁵

1 Botanic Gardens of Sydney, Mount Annan, Australia.

2 University of New South Wales, Sydney, Australia. 3

University of Sydney, Sydney, Australia. 4 Universidad

Nacional Autónoma de México, Ciudad de México,

Mexico. 5 University of Vienna, Vienna, Austria.

The eFLOWER project started thirteen years ago as a long-term initiative to address key questions on the early evolution and diversification of angio-sperm flowers. A central part of eFLOWER has been the development and continuous curation and ex-pansion of a collaborative database named PROTE-US allowing to record floral traits at the species level while documenting the explicit source of each data point. Here we review the diverse contributions that eFLOWER has brought so far to our understanding of floral evolution, reflect upon the questions and chal-lenges encountered along the way, and outline fu-ture directions. An important milestone was reached seven years ago with the publication of a new mod-el for the ancestral flower of all angiosperms, based on ancestral state reconstructions using a dataset of 792 species sampled from most families. Despite uncertainty remaining for some traits, this new mod-el proved effective in questioning long-term as-sumptions and opening new research opportunities on floral evolution. Interestingly, this model appears to be robust to new analyses of a recently published, expanded dataset of 1201 species sampled from all families, but some results continue to be highly sen-sitive to methodological approaches and models of character evolution. Another significant milestone has been the development and evaluation of angio-sperm-wide 'phyloscan' approaches to assess the

phylogenetic position of fossil flowers using these datasets of floral traits. While these methods appear to provide a powerful new tool to systematically assess fossil flowers, they have also revealed high uncertainty in the placement of many taxa, reflecting the limits of the number of preserved, standardised characters that may be recorded from fossil flowers. Lastly, we present briefly some recent new avenues linked with this project, including new ideas on the origin of pollination and evidence on early angio-sperm diversification from morphospace analyses of floral disparity.

**S.2.2 Evolution of fruits and seeds of Solanaceae, from extinct to extant diversity**

Rocio Deanna¹,²

1 Instituto Multidisciplinario de Biología Vegetal (CONICET-UNC), Córdoba, Argentina. 2 Depart-ment of Ecology and Evolutionary Biology, Universi-ty of Colorado at Boulder, Boulder, USA. rociodean-na@gmail.com

The placement of extinct taxa on phylogenies relies on scoring informative morphological characters, which can be difficult given the fragmentary nature of the fossil record. Although detailed studies of Solanaceae fossils remain few, the family exhibits wide variation in seed and fruit traits to use in estimating fossil placement from morphological data. Therefore, we reviewed a total of 106 seed and seven fruit fossils of Solanaceae, including two recently published South American species that are the oldest records in the family. We scored 28 traits, including 11 continuous and 17 discrete, for all the fossils gathered and almost 400 extant taxa, sampling at least one species per genus. We then performed a cluster analysis to morphologically compare these fossil seeds to extant Solanaceae, along with a phylogenetic analysis including all the morphological traits and 10 DNA regions in order to infer the phylogenetic placement of the fossils. We also reconstructed the ancestral states of the most conspicuous morphological traits over a molecular phylogenetic tree
<figure><img src="image_23.png" /></figure>

to finally compare the morphological evolution in-
ferred with the placement of the fossils. The clusters
found in the statistical analysis were congruent with
the phylogenetic placement of the fossils in all the
species, except by some fossil seeds. According to
the morphological reconstructions, traits that have
been acquired only once in the family were the most
informative to place the fossils, like the finger-like
appendages in the fruiting calyx of the fruit fossil

*Lycianthoides calycina* placed in the chili pepper clade. Converged traits (e.g., inflated fruiting calyx) were less informative but the combination of fruiting calyx traits was sufficient to place the *Physalis* fruit fossils. Continuous traits were highly informative, especially in the placement of the fossil seeds. These results lay the foundation for new total evidence dating analysis for the entire family.

S.2.3 The evolution of embryo size in angiosperms seeds

Filip Vandelook¹², Angelino Carta³

1 Meise Botanic Garden, Meise, Belgium. 2 Katho-
lieke Universiteit Leuven, Leuven, Belgium. 3 Univer-
sity of Pisa, Pisa, Italy.

Seeds are the main dispersal and propagation units of angiosperms, and they are very diverse in their external and internal features. When dispersed, seeds consist of an embryo, the next generation of a plant, surrounded by the seed and/or fruit coat and by an alternative storage tissue (endosperm or perisperm) in case the nutritive reserves are not stored in the cotyledons. Examining the relative allocation of seed reserves by quantifying the relative embryo size at dispersal (i.e., size of the embryo relative to the seed, ES) across angiosperms, sets the basis to track the evolutionary history of this key reproductive trait related to germination timing and offspring survival. We used macroevolutionary approaches to estimate adaptive shifts of ES in the entire angiosperm radiation. We also evaluated how tempo (i.e., evolutionary rates) and mode (i.e., evolutionary process) of ES evolution changed through time and the role of paleo-temperatures on ES evolutionary rates. Early angiosperms typically had a low ES, which is still reflected in contemporary magnoliids and ANA-grade species. The analysis of major shifts in ES throughout angiosperm evolution revealed that these shifts occurred after 145 Myr and were predominantly associated with the formation of the main angiosperm lineages. Comparison of a series of paleo-inspired models showed that tempo and mode of ES evolution altered from a very dynamic ES evolution early in angiosperm history towards a more static evolution driven by stronger stabilizing selection. Finally, a consistent trend for a slowdown in rates of ES evolution during periods of climate warming was found. Fast evolutionary rates under cold climates are consistent with the macroecolog-

ical observation that rates of phenotypic evolution
are higher at high latitudes. This study provides cru-
cial new insights in seed trait evolution, which con-
tribute to understanding the diversification of repro-
ductive strategies in angiosperms.

S.2.4 Are flower and fruit colors associated with environment? Exploring global ecogeographic patterns in flower and fruit colors

Agnes S. Dellinger¹, Leah Meier², Stacey D. Smith², Miranda Sinnott-Armstrong²,³

1 Department of Botany and Biodiversity Research, University of Vienna, Vienna, Austria. 2 Department of Ecology and Evolutionary Biology, University of Colorado Boulder, Boulder, CO, USA. 3 Department of Chemistry, University of Cambridge, Cambridge, UK

Flowers and fruits are central to plant reproduction because they facilitate gene dispersal through pollen or seeds, respectively. In species engaging in biotic interactions during reproduction (i.e., with pollinators and dispersers), flower and fruit coloration are important as visually attractive cues. Different groups of pollinators (i.e., bees, birds, bats) and dispersers (i.e., birds, monkeys, lizards) differ in their sensory abilities, which might drive color evolution in flowers and fruits. The abiotic environment may also be an important selective factor on coloration. For instance, the synthesis of anthocyanin pigments (most reds, pinks, and blacks) is triggered by cold temperatures and UV-B radiation. Although flowers and fruits originate from the same organ and con-
<figure><img src="image_24.png" /></figure>

stitute a continuous developmental sequence, we
currently lack understanding of whether flower and
fruit traits are correlated in space. If flower and fruit
colors show similar spatial patterns, those patterns
may be influenced strongly by similar abiotic fac-
tors. If, however, flower and fruit color are decoupled,
then they likely result from different selective agents
(either abiotic or biotic). To address this gap, we col-
lated a dataset of 2800+ species from 51 clades with
animal-pollinated flowers and animal-dispersed
fruits and scored flower and fruit colors into eight
categories. We combined the color data with GBIF
occurrence records for these species and found
striking differences in the geographic distribution
of flower and fruit colors. Overall, fruit color diversity
is higher in the tropics, but flower color diversity is
not. Red and pink in both flowers and fruits are more
prevalent at high northern latitudes, which suggests
that a common selective agent may drive the evo-
lution of red/pink under those environmental con-
ditions. Aside from red/pink, flower and fruit colors
and color diversity are decoupled across the globe,
indicating different importance of abiotic and biotic
selective agents.

S.2.5 The emergence of a
floral economic spectrum
and its association with the
leaf economic spectrum

Lars Götzenberger¹,², Marketa Applova¹,²,
Anna E-Vojtko¹

1 Institute of Botany of the Czech Academy of Sciences, Trebon, Czech Republic. 2 Department of Botany, University of South Bohemia, Budweis, Czech Republic.

Understanding resource allocation strategies in
plants is crucial for unraveling ecological and evo-
lutionary dynamics. The traditional fast-slow con-
tinuum is a well-established framework that has
primarily been explored through vegetative above
and below-ground traits of leaves and roots. Re-
cent propositions suggest the existence of a floral
economic spectrum (FES) that parallels leaf traits,
in particular with regard to area and mass-related
traits, as well as longevity of individual organs. In a
survey of herbs across a wide range of plant fam-
ilies, we confirm the expected negative correlation
between specific leaf area (SLA) and leaf dry mat-
ter content (LDMC) but also demonstrate a similar
relationship in flowers, i.e., a negative correlation

between specific floral area (SFA) and floral dry
matter content (FDMC). Across organs, the anal-
ogous traits are positively associated, i.e., FDMC
correlates positively with LDMC, and SLA with SFA.
In addition, with data from a greenhouse experi-
ment of 8 species, we show that these relationships
also hold within species, and are influenced by wa-
ter availability. Our findings show that flowers and
leaves, though serving very different functions, are
constrained at least to some degree with respect
to their morphology and structure. The often-as-
sumed independent evolution of floral characters
depends much on what traits are considered, and
the same can be said about the previously report-
ed independent dimension of floral traits in the
space of plant form and function.

S.2.6 The seed germination
spectrum of Mediterranean
plants

Diana María Cruz Tejada¹, Eduardo
Fernández Pascual², Efisio Mattana³,
Angelino Carta¹

1 University of Pisa, Pisa, Italy. 2 IMIB Biodiversity Research Institute (University of Oviedo - CSIC - Principality of Asturias), University of Oviedo, Mieres, Spain. 3 Royal Botanic Gardens, Kew, Wakehurst, UK.

The understanding of seed biology of wild plant
species is fundamental to address ecological, evo-
lutionary, and biogeographical questions, as well
as to the success of seed-based conservation and
restoration actions. Despite recent attempts to syn-
thesise seed germination knowledge across major
temperate biomes, most assumptions about the
germination ecology of Mediterranean plants are
based on individual species or local studies, miss-
ing an integrated view of germination patterns for
the whole Mediterranean Basin biodiversity hotspot.
Here, we performed a binomial phylogenetically in-
formed Bayesian meta-analysis, considering ca.
7,000 germination tests of ca. 400 species charac-
teristic of EUNIS Mediterranean habitats (Europe-
an habitat classification system) from laboratory
experiments conducted across the Mediterranean
Basin. We estimated the effect on germination pro-
portions of five environmental cues, including mean
incubation temperature, temperature regime, light,
and dormancy-breaking treatments for typical
Mediterranean plants (i.e., species that only occur
in lowland habitats of the Mediterranean Basin) and
generalist plants (i.e., distribution wider than the Mediterranean Basin). Overall, typical Mediterranean species exhibit a cold-cued germination response, a positive effect of scarification, and are insensitive to light and alternating temperatures, in accordance with previously described germination syndromes for the area. Generalist species are also stimulated by cool temperatures but show positive effects of alternating temperatures, light, and cold stratification. Our results suggest that specific germination responses could vary according to species macroclimatic constraints (typical vs. generalist species) and secondarily local habitat conditions. Indeed, we

found consistent patterns across different habitats (forests, scrublands, and grasslands) with few exceptions, including anthropogenic habitats, in which opportunistic germination strategies were found. Finally, whilst phylogenetically related species exhibit similar germination responses, our results also suggest a functional convergence of the seed germination patterns across Mediterranean species. Our integrative methodological approach emphasised the significance of utilising extensive datasets to formally test eco-evolutionary and biogeographical research questions.

# S.3 APOCYNACEAE - ON THE WAY TO A MODEL FAMILY

## S.3.1 Phylogenomic analysis of Apocynaceae based on the Apoc836 probe set: toward resolution of recalcitrant nodes

Chelsea R. Smith¹,², Shannon C. K. Straub³, Tatyana Livshultz¹,²

1 Academy of Natural Sciences of Drexel University, Philadelphia, USA. 2 Drexel University, Philadelphia, USA. 3 Hobart and William Smith Colleges, Geneva, USA

The broad outline of Apocynaceae phylogeny, divided into three subfamilies (Periplocoideae, Secamonoideae, Asclepiadoideae) and two informal grades: rauvolfioids (11 tribes) and apocynoids (9 tribes) has been stable since the earliest single gene plastid analyses: a paraphyletic rauvolfioid grade subtending a monophyletic APSA clade (apocynoids, Periplocoideae, Secamonoideae, Asclepiadoideae). Accumulation of data, culminating in the first phylogenomic analysis of the family (Antonelli, et al. 2021) has highlighted areas of continued topological uncertainty including relationships within the APSA clade, specifically the positions of Periplocoideae and Rhabdadenieae, and an earlier radiation that includes the rauvolfioid tribes Melodineae, Hunterieae, Amsonieae, and Alyxieae. We revisited Apocynaceae phylogenomics with expanded sampling of taxa and loci (Apoc836; Straub, et al. 2020). We constructed trees using summary coalescent

and concatenation methods and conducted sensitivity analyses to evaluate the robustness of our topologies to sampling and analytical decisions. An intriguing outcome of these analyses is the frequent placement of Periplocoideae as sister to the well-established clade of Baisseeae, Secamonoideae, and Asclepiadoideae, harkening back to the abandoned Asclepiadaceae (comprised of Periplocoideae, Secamonoideae, and Asclepiadoideae) and re-opening questions about homology of pollen aggregation, translators, and coronas in these taxa. This topology is sensitive to both sampling and analytical choices, and we conclude that the weighted AS-TRAL analysis most accurately represents remaining uncertainty. This new Apocynaceae species tree suggests a revised classification for the family at the subfamily and tribal levels.

References: Antonelli, A., et al. (2021). Settling a family feud: a high-level phylogenomic framework for the Gentianales based on 353 nuclear genes and partial plastomes. https://doi.org/10.1002/ajb.2.1697. Straub, S. C. K., Boutte, J., Fishbein, M., & Livshultz, T. (2020). Enabling evolutionary studies at multiple scales in Apocynaceae through Hyb-Seq. Applications in Plant Sciences, e11400. https://doi.org/10.1002/aps3.11400.
<figure><img src="image_25.png" /></figure>

## S.3.2 Biome lability and substrate conservatism fuelled plant evolution in xeric ecosystems

Cássia Bitencourt¹; Conal Aitken¹; Matilda Brown¹; Leonel Herrera-Alsina²; Olivier Maurin¹; Natalia Przelomska¹; Moabe F. Fernandes¹³; Mohammad Vatanparast¹; Alessandro Rapini⁴; Mary Endress⁵; Sigrid Liede-Schumann⁶, David Goyder¹; Brian Schrire¹; Kyle Dexter⁷; Nicolai Nürk⁸; Jan Hackel⁹; Mike Gilbert¹; Raquel Negrão¹; Alexandre Zuntini¹; Paul Kersey¹; Ilia Leitch¹; Eve Lucas¹; Oscar A. Pérez-Escobar¹; Melanie J. Howes¹; William J. Baker¹; Eimear Nic Lughadha¹; Alexandre Antonelli¹,¹⁰,¹¹; Félix Forest¹

1 Royal Botanic Gardens, Kew, London, UK. 2 University of Aberdeen, Aberdeen, UK. 3 University of Exeter, Exeter, UK. 4 Universidade Estadual de Feira de Santana, Feira de Santana, Brazil. 5 Institute of Systematic Botany, Zürich, Switzerland. 6 University of Bayreuth, Bayreuth, Germany. 7 University of Edinburgh, Edinburgh, UK. 8 University of Bayreuth, Germany. 9 Philipps-Universität Marburg, Marburg, Germany. 10 University of Gothenburg, Gothenburg, Sweden. 11 University of Oxford, Oxford, UK.

The expansion of arid environments following the Eocene-Oligocene climate transition is thought to have promoted increased species diversification rates across several plant lineages. In this study, we use the Apocynaceae as a model to assess whether a combination of biotic (e.g., lifespan and dispersal traits) and abiotic (e.g., climate and soil) predictors favoured diversification during the expansion of dry biomes. To accomplish this, we reconstructed a comprehensive time-calibrated phylogenetic tree for Apocynaceae using the Angiosperms353 bait set to establish a backbone. Subsequently, we merged all Apocynaceae species with sequence data (for ITS, trnL-F, and rps16) available on GenBank. Finally, distribution and trait data were compiled to test three hypotheses: [H1] Biome shifts between Apocynaceae species primarily occur from rainforest to savanna, whereas transitions between tropical and temperate environments are rare; [H2] Diversification in Apocynaceae was driven by "key synnovation" involving anemochory,

presence of pollinia and climbing habit; and [H3] High speciation rates favoured the diversification of Apocynaceae lineages adapted to xeric environments. Our study suggests that lineages in more xeric environments exhibit higher speciation rates as they adapt to new, harsher conditions. These findings suggest that substrate, rather than biome, might be the primary factor regulating evolutionary rates in plants adapted to dry ecosystems.

## S.3.3 Marsdenieae (Apocynaceae – Asclepiadoideae) – phylogenetic analysis clarifies generic concept and unveils abundant parallelisms

Michele Rodda¹, Sigrid Liede-Schumann²

1 Singapore Botanic Gardens, National Parks Board, 1 Cluny Road, Singapore 25969, Singapore. 2 University of Bayreuth, Department of Plant Systematics, Bayreuth, Germany

## S.3.4 Biogeography and multiple origins of Indian open-flowered Ceropegia

Siddharthan Surveswaran¹

1 Department of Plant Sciences, University of Hyderabad, Hyderabad, India.

The open-flowered type of *Ceropegia*, formerly *Brachystelma* is a rare and elusive group of plants in India. They occupy the relatively dry habitats of peninsular India which is influenced by the highly seasonal monsoonal climate. The diversity of *Brachystelma* is about 35 species in peninsular India. Molecular phylogeny reveals that within the radiation of *Ceropegia* (tubular-flowered species). *Brachystelma* has evolved at least three times from closed tubular-flowered. In this study, I examine the distribution of the Indian open-flowered lineages based on the most up-to-date and comprehensive sampling of the genus. The diversity of the genus is linked to open grassland in the Eastern Ghat region of India. Using climatic niche modelling, I discuss the climatic requirements which
<figure><img src="image_26.png" /></figure>

might have led to the diversity of the open-flowered genotypes. Furthermore, using transcriptomic approaches, I examine the genes responsible for the shift from tubular flowers to open type flowers. I also discuss preliminary data on pollinators of Indian open-flowered genera of *Ceropegia*.

A densely sampled molecular phylogeny of the pantropical, predominantly forest-dwelling tribe Marsdenieae, including almost a quarter of the ca. 740 species shows that the tribe is divided in two large clades, with two genera *Rhyssolobium* (southern Africa) and *Sicyocarpus* (Madagascar) placed outside of both clades. The first of the two large clades has a native distribution exclusively in Asia and the Pacific area. The type genus of the tribe, *Marsdenia* R.Br., is sister to all remaining genera of this clade, and comprises ca. 10 species. This clade harbours two species-rich radiations, *Leichhardtia* (ca. 80 species) in the Pacific realm and the *Hoya*-alliance (>300 species). The second large clade shows a cosmopolitan distribution with several small Asian lineages as sisters to some larger genera of African-Asian distribution, as well as the New World genus *Ruehssia* (ca. 130 species). As far as known, all larger clades comprise species adapted to various types of pollinators (small and larger flies, bees, butterflies, moths), forcing highly similar floral structures. For the *Hoya*-alliance, we present a densely sampled phylogeny obtained using target enrichment and including ca. 80 % of the known diversity of *Hoya* and *Dischidia*, all taxa of *Oreosparte* (3 species) and *Papuahoya* (3 species). *Dischidia*, *Oreosparte* and *Papuahoya* are confirmed as monophyletic, but *Hoya* is paraphyletic, with a smaller clade sister to *Papuahoya* and a larger clade (*Hoya* s.s.) sister to *Dischidia* + *Oreosparte*. Within *Hoya* s.s., some clades are characterised by uniform flower morphologies while others, including the main radiation of species in New Guinea, independently developed a large range of flower morphologies. The largest molecular and morphological diversity of *Hoya* s.s. is found in Borneo. Our results will contribute towards a modern, stable classification of the *Hoya*-alliance, and studies on its origin and diversification.

### S.3.5 Tackling tough taxonomy: genomic, morphological, and ecological approaches to Asclepias (Apocynaceae) in North America

Mark Fishbein¹, Shannon C. K. Straub², Victor Andreev¹,³, Julien Boutte⁴, Carl Wagner², David Kunkel¹, Therese Mandracchia², Alessandra Bryan², James Riser⁵

1 Oklahoma State University, Stillwater, USA. 2 Hobart and William Smith Colleges, Geneva, USA. 3 University of Arizona, Tucson, USA. 4 A2Bio2, Molecules, France. 5 University of Wisconsin-Platteville, Platteville, USA

Rapid diversification may present taxonomic challenges resulting from the gradual nature of speciation. Processes including incomplete fixation of ancestral polymorphisms and partial establishment of reproductive barriers permitting gene flow after speciation result in conflicting information about species boundaries and phylogeny. The rapid diversification of >130 species of American *Asclepias* provides a good system to demonstrate that integration of genomic data, population genetics, ecological modeling, and traditional approaches can resolve longstanding taxonomic challenges. We present two case studies. The “dwarf milkweed” clade has been variously treated as 1-4 species (*A. eastwoodiana*, *A. ruthiae*, *A. sanjuanensis*, *A. uncialis*) distributed across semiarid western North America. Populations exhibit limited morphological variation, largely in the shape and vestiture of leaves. Previous research supported recognition of four species based on population genetics and ecological niche modeling. We support this taxonomy with a plastome phylogeny and newly discovered floral differences and resolve the phylogeny of the four species using sequence data from >2000 nuclear genes. In the second example we address circumscription of *A. hirtella* and *A. longifolia*. Populations of these taxa are found in the eastern United States, with coastal populations assigned to *A. longifolia* and interior populations to *A. hirtella* (or *A. longifolia* subsp. *hirtella*). In the western Gulf Coast (Texas and Louisiana) populations are morphologically intermediate and have been commonly assigned to *A. longifolia*, which they most resemble. Using population genomic data
<figure><img src="image_27.png" /></figure>

from >1400 SNP loci and a plastome phylogeny, we found evidence for two largely discrete gene pools. However, the western Gulf Coast populations belong to the interior *A. hirtella*, not the coastal *A. longifolia*, and we present supporting morphological evidence for this circumscription. These case studies pave the way for achieving deeper understanding of evolutionary and biogeographic history of *Asclepias*.

### S.3.6 Citizen science and its impact on systematics and natural history of Apocynaceae of Mexico

Leonardo O. Alvarado-Cárdenas¹

1. Departamento de Biología Comparada, Facultad de Ciencias, Universidad Nacional Autónoma de México, Ciudad de México, México.

By 2003, humanity had created the equivalent of 5 exabytes of information. Today, this information is generated in 2 days. Although most of it is entertainment, some of it has allowed us to benefit and connect the daily life of citizens and scientists. Social networks have enabled a proximity between science and society, but citizen science platforms have created a very productive link, an example of which is INaturalist. In Mexico, INaturalist integrates

more than 3000 photos daily, many of them exotic or cultivated organisms, but others of great importance for the information they contain. This has contributed to aspects such as systematics, biography, plant-animal interactions, biological invasions and, of course, the relationship between people and scientists. One of the plant families that has benefited from these platforms is the Apocynaceae. This is one of the most diverse families in the world, with more than 5000 species, and in Mexico there are about 500 species, making it the 12th most diverse family in the country. Since 2018, the INaturalist database has been managed as the "Apocynaceae of Mexico" project. The project includes about 60,000 photos, the presence of 4 specialists in the group and a curation of more than 80 %. As a result, the distribution of many taxa has been adjusted. Over 400 species have been documented, including some with flowers and fruits. The documentation includes the presence of five species that have not been collected in over 30 years, the discovery of five new species, and the identification of invasive species within natural areas. Likewise, floral visitors and potential pollinators have been documented, many of them unknown to the country. Proper utilization and management of this information will enhance the comprehension of the biology and systematics of the organisms under investigation.

## S.4 BIOGEOGRAPHY AND DIVERSIFICATION OF INTERCONTINENTAL TROPICAL PLANT DIVERSITY. SESSION 1

### S.4.1 The biogeography of tropical plants with megafaunal fruits

Renske E. Onstein¹,², Seheno

Andriantsaralaza³, Onja Razafindratsima⁴,

Andressa Cabral², Anna Traveset⁵, Mauro

Galetti⁶

1. Naturalis Biodiversity Center, Leiden, the Netherlands. 2. German Centre for Integrative Biodiversity Research, Halle-Jena-Leipzig, Leipzig, Germany. 3. University of Antananarivo, Antananarivo, Madagascar. 4. University of California Berkeley, Berkeley,

USA. 5 Instituto Mediterráneo de Estudios Avanzados (CSIC-UIB), Mallorca, Spain. 6 São Paulo State University, São Paulo, Brazil

Plants with megafaunal fruits (>4 cm) have relied on dispersal by megafauna and other large-bodied vertebrates (> 40 kg) throughout their million-year evolutionary history. How megafruit plants have persisted in contemporary ecosystems after the Pleistocene extinction of most megafauna - especially in the Neotropics, Madagascar, and Australasia - remains an evolutionary enigma. By integrating global data on traits, seed dispersal interactions, phylogenies, and geographical occurrences for >1500 megafruit plants, we show that many megafruit species (613 species, 40%) interact with contempo-
<figure><img src="image_28.png" /></figure>

orary dispersers, including humans, megafauna (e.g., elephants), secondary dispersers (e.g., rodents), or water, providing an explanation for their persistence. These megafruit species often have large geographical range sizes, strongly contrasting the 128 (up to 300) megafruit species that are currently threatened with extinction, suffer from small range sizes, and lack suitable co-occurring dispersers, primarily on (isolated) tropical islands in the Australasia, IndoMalay and Madagascar regions. Our results illustrate how the Pleistocene extinctions and ongoing defaunation limit seed dispersal of many tropical plant populations with megafaunal fruit, but humans may have replaced megafaunal dispersal functions for its subset of taxa.

## S.4.2 The spatio-temporal co-diversification of frugivores and three vertebrate-dispersed tropical plant families

Andressa Cabral¹, Yaowu Xing², Renske E. Onstein¹,³

1. German Centre for Integrative Biodiversity Research (iDiv) Halle - Jena - Leipzig, Germany. 2. Xishuangbanna Tropical Botanical Garden, CAS Menglun, Mengla, Yunnan, China. 3. Naturalis Biodiversity Center, Leiden, The Netherlands.

Fleshy-fruited plants and their fruit-eating and seed-dispersing animals (frugivores) have originated millions of years ago, but it remains unclear when and where their mutualistic interactions evolved, and whether this led to co-diversification and co-adaptation of frugivory-related traits. Here, we hypothesize that shifts in diversification and trait evolution rates of tropical, fleshy-fruited plant lineages coincided with the evolution of frugivorous vertebrate lineages and their traits, as a result of co-evolutionary selective pressures from interacting partners. To test this hypothesis, we focus on three pantropical keystone plant families comprising >6,200 species with vertebrate-dispersed fruits: custard apples (Annonaceae), palms (Arecaceae) and mulberries and figs (Moraceae). We align their evolution with that of their primary seed dispersers based on data from extant and Late Quaternary mammals and birds (~5,200 species). Because fruit and seed size are constrained by frugivore gape widths and body sizes, with the largest fruits depending on dispersal by the largest animals, we also collected information on

fruit and/or seed sizes and body sizes/gape widths/diet. We integrated these data in a phylogenetic framework and used comparative methods to identify the origin and biogeography of major frugivore lineages and their food plants. Using diversification and trait evolution models, we showed that several plant and frugivore lineages co-diversified in the Cenozoic (i.e., last 66 million years), with increased rates of diversification coinciding with increases in fruit and body size evolution. These findings suggest that reciprocal adaptations in frugivory-related morphological traits between plants and frugivores have influenced their mutualistic-dependent co-diversification across the tropical realms, explaining high contemporary diversity and 'trait matching' in tropical rainforests.

## S.4.3 Climatic niche and functional shifts after Neotropics colonization drive diversification in the amphi-pacific genus Dendropanax

Angélica Gallego-Narbón¹, Jun Wen², Luis Valente³,⁴, Gabriel Johnson², Marina Coca-de-la-Iglesia¹, Nagore G. Medina¹,⁵, Mario Fernández-Mazuecos¹,⁵, Virginia Valcárcel¹

1. Universidad Autónoma de Madrid, Madrid, Spain.
2. Smithsonian National Museum of Natural History, Washington DC, USA.
3. Naturalis Biodiversity Center, Leiden, The Netherlands.
4. University of Groningen, Groningen, The Netherlands.
5. Centro de Investigación en Biodiversidad y Cambio Global (UAM), Madrid, Spain.

Dendropanax is a species-rich genera of Araliaceae and a classic example of the Asian-Neotropical disjunction or amphi-pacific tropical disjunction. Previous biogeographic reconstructions of the genus point to an East Asian origin and posterior colonization of the Neotropics, with higher diversity in the Neotropical clade. Here we investigated the role of climatic niche evolution and colonization events in the diversification of the functionally and climatically heterogeneous genus Dendropanax. We used Hyb-Seq phylogenomics on 237 Araliaceae species to obtain time-calibrated nuclear and plastid phylogenies used to perform biogeographic analyses, diversification models, and climatic niche and functional trait reconstructions (habit, leaf type, dimor-
<figure><img src="image_29.png" /></figure>

phism, and area, stipule and trichome presence) for *Dendropanax* and phylogenetically close genera. Curated databases were used to estimate current and ancestral climatic niches (6,496 occurrences) and leaf areas (910 measurements) using a novel probabilistic approach that considers niche heterogeneity. Diversification in *Dendropanax* followed a temperature-based model with a decrease of diversification towards the present. After an Asian origin, successive colonizations of Central and South America preceded climatic niche expansions and shifts towards warmer and drier climates coupled with an increase in leaf area and variability and associated with the persistence of high diversification rates in the Neotropical *Dendropanax* clade in comparison with the Asian *Dendropanax* clade. Given that the evolution of leaf area followed the colonization of the Neotropics, we believe that this trait could be associated with the adaptation to the climatic conditions in the colonized area. Our results support the relevance of colonization events, climatic niche evolution and functional trait shifts to explain phylogenetic diversification in the intercontinental tropical *Dendropanax*.

### S.4.4 Phylogenetic insights into the origins of tropical rainforest hyperdiversity: Big Data versus model groups

Wolf L. Eiserhardt¹,², William J. Baker¹,², Paola de Lima Ferreira¹, Lars Emil S. F. Hansen¹, Oscar B. Wrisberg¹, Melanie Tietje¹

1 Department of Biology, Aarhus University, Aarhus, Denmark. 2 Royal Botanic Gardens, Kew, Richmond, Surrey, UK.

Rainforests occur across the tropics and are consistently the most biodiverse biome on Earth. Despite centuries of scientific interest, the eco-evolutionary drivers of rainforest hyperdiversity remain elusive. That said, it seems clear that explanations have to be sought in deep time, involving dynamics of diversification, migration and adaptation over tens to hundreds of millions of years. Phylogenetic trees can provide a unique window on those deep-time dynamics, but are notoriously hard to obtain at the scale needed to unpick rainforest hyperdiversity. Robust and well-sampled phylogenetic trees tend to be only available for small "model groups" that may or may not be representative for the assembly of rainforest biotas. Meanwhile, "Big Data" approach-

es investigating major groups as a whole (e.g. seed plants) consistently suffer from limited, patchy and biased phylogenetic data. Similar trade-offs exist for the geographic and ecological data needed to assign species to biomes, a prerequisite for investigating biome assembly over time. Here, we discuss recent efforts to understand the evolutionary assembly of rainforest hyperdiversity using both "Big Data" and "model group" approaches, comparing their findings and highlighting the specific challenges, pros and cons of each approach. Specifically, we report our recent findings based on phylogenetic, geographic and biome association datasets for all seed plants (Spermatophyta), contrasting them with focused studies mostly from the palm family (Arecaceae), a well-established model group for tropical plant ecology and evolution.

### S.4.5 Reconciling deep and recent time to unravel global tropical rain forest diversification dynamics at unrivaled levels of precision

Laura Holzmeyer¹, Francis J. Nge¹,², Vincent Soulé¹, Thomas L.P. Couvreur¹, Annonaceae Phylogenetics Consortium³

1 IRD, UMR DIADE, Université de Montpellier, Montpellier, France. 2 National Herbarium of New South Wales, Australian Institute of Botanical Science, Royal Botanic Gardens and Domain Trust, Mount Annan, Australia. 3 Annonaceae Global Phylogenetic Consortium, http://www.couvurlab.org/agpc.

The evolution and processes leading to the remarkable biodiversity contained within tropical rain forests (TRF) have been subject to research for decades. Traditionally, at a macroevolutionary scale, two predominant contrasting hypotheses are generally debated, the museum hypothesis, where low extinction rates lead to a constant diversification and a steady increase of diversity through time, and the cradle hypothesis, where increased speciation rates lead to rapid bursts of diversity. To date, most studies were based on regional or limited global taxonomic sampling spanning either recent or deep time scales. These studies suggested a more nuanced view with a mixed or neutral model of high species turnover, implying both elevated speciation and extinction rates in TRF. Here, we reconcile deep and recent time scales to test these hypotheses by generating for the first time a global near
<figure><img src="image_30.png" /></figure>

complete species level dated phylogenetic tree of the major TRF family Annonaceae Juss. (ca. 2000 out of 2450 species). Using this framework, we tested both hypotheses using different macroevolutionary methods and statistics. We shall present the major results obtained from these analyses and implications for our understanding of TRF evolution through time at global scales. Our study marks a significant advancement in understanding TRF evolution, by presenting the first comprehensive species-level phylogenetic tree of one of the oldest rain forest angiosperm families on a global scale.

### S.4.6 The biogeography and evolution of derived woodiness in tropical and sub-tropical angiosperms

Alexander Zizka¹,²*, Renske Onstein²,³, Frederic Lens²,⁴

1 Department of Biology, Philipps-University Marburg, Marburg, Germany. 2 Naturalis Biodiversity Center, Leiden, The Netherlands. 3 German Centre for Integrative Biodiversity Research Halle-Jena-Leipzig, Leipzig, Germany. 4 Institute of Biology Leiden, Leiden University, Leiden, The Netherlands.
*alexander.zizka@biologie.uni-marburg.de.

The repeated evolution of woodiness in herbaceous lineages has long been perceived as defining feature of island floras, in particular in the Tropics and sub-Tropics ("insular woodiness"). However, scattered evidence indicated comparable transitions of herbaceous lineages towards woody growth on continents, for instance on tropical mountains ("derived woodiness"). Here, we contrast recent results on the evolution of insular woodiness with a novel dataset on derived woodiness on continents. We identified numerous independent transitions towards woodiness in 57 eudicot angiosperm families, resulting in more than 6,500 recent derived woody species. We found more transitions and derived woody species on continents than on islands but derived woodiness on islands was more common when accounting for total angiosperm richness (on average 0.77% of the total flora on continents vs 1.85% on islands). Furthermore, we identified four global continental centers of derived woodiness, mostly in the Tropics and sub-Tropics: The Andes, South Africa, parts of Australia and New Zealand ("Australasia"), and the Old-World dry belt. Our results reveal the re-evolution of woodiness in herbaceous lineages as widespread phenomenon on island and on continents driven globally by environmental conditions, albeit with important regional and systematic idiosyncrasies.

## S.5 ADVANCES IN SEED CONSERVATION OF WILD SPECIES. SESSION 1

### S.5.1 Status of European seed Conservation of Wild Plants in Seed banks

Andreas Ensslin¹, Adelaide Clemente², Elke Zippel³, Udayangani Liu⁴, Peter Glasnović⁵, Sandrine Godefroid⁶

1 Conservatory and Botanic Garden of the City of Geneva, Switzerland. 2 University of Lisbon, Portugal. 3 Botanic Garden and Botanical Museum Berlin-Dahlem, Germany. 4 Royal Botanic Gardens Kew, UK. 5 University of Primorska, Slovenia. 6 Meise Botanic Garden, Belgium.

Conserving orthodox seeds of wild plants through seed banking (at 15% RH and -20°C) has become a valuable tool in safeguarding threatened pop-

ulations and species, supporting conservation interventions such as plant translocations, but also in conserving genetic resources for plant breeding and food security. Despite over 90% of the European flora is likely to possess bankable seeds, there is no general assessment of how many taxa are secured in seed banks, where the gaps are and whether the countries met the objectives of global conservation initiatives such as the GSPC (i.e., target 8: 75% of threatened plants conserved ex situ). Within the framework of the COST Action Conserve-Plants (CA18201), a European-wide survey on seed bank collections was launched to gain an overview over the current state of ex situ conservation of wild plant taxa in seed banks across Europe. After a challenging process of taxonomic harmonization, we identified over 13,000 taxa conserved in 109 institutions from 29 countries across Europe and the
<figure><img src="image_31.png" /></figure>

Middle East. We mapped the taxonomic names and the original provenance of seed lots against the respective national red lists to identify the coverage met by each country concerning the GSPC target 8. We reveal gaps in species conservation and point out future directions on how to meet global conservation targets. Our study demonstrates the importance of collaborative approach for seed banking to support safeguarding threatened plants across Europe but also highlights that its potential has not at all been fully exploited yet.

## S.5.2 Research Progress on in vitro preservation of Wild Plants in China

Jun He Junbo¹, Yang Dezhu Li¹

1 Germplasm Bank of Wild Species, Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, Yunnan, People's Republic of China.

Both in situ and ex situ conservation are complementary and synergistic approaches that are essential for the protection of wild plants and their biodiversity. As a major national scientific infrastructure, the Southwest China Germplasm Bank of Wild Species has established a wild plant germplasm resource bank that includes a seed bank, plant tissue culture bank, DNA bank, animal germplasm bank, and microbial germplasm bank. The plant tissue culture bank is mainly based on the theory and methods of tissue culture to carry out species conservation, in order to achieve long-term conservation of "exceptional" species with low yield, poor quality, recalcitrance, and cannot be directly sown. Explants are obtained through field collection or exchange, and then cultured through various methods such as protocorn type, short branch cutting type, clustered shoots proliferation type, or organ type. Finally, strategies such as normal temperature or low temperature restriction are used to achieve in vitro preservation of germplasm resources. The use of in vitro conservation methods allows wild plant germplasm resources to obtain regenerability, repeatability, and sustainability in research and utilization, which is a major characteristic that distinguishes in vitro preservation from other ex situ conservation methods and provides new strategies to make up for the shortcomings of in situ and other ex situ conservation methods. The staff of the Southwest China Germplasm Bank of Wild Species has made certain progress in large-scale in vitro preservation of wild plants through years of accumulation. As of the end

of 2023, the plant tissue culture bank has preserved in vitro culture materials of more than 1,100 species and 27,100 samples, with orchid species accounting for over 60%. It has preserved one-third of the species and 55% of the genera of wild orchids in China, as well as other rare and endangered plants.

## S.5.3 Spectrometry and machine learning technologies to aid conservation of native dry forest seeds

Maria A. R. Ferreira¹, Raquel A. Gomes¹, Jamille C. Silva¹, Jailton J. Silva¹, Williane A. S. Oliveira¹, Marcos V. C. Freitas¹, Sérgio T. Freitas¹, Bárbara F. Dantas¹

1 Embrapa Semiárido, Petrolina, Brazil.

Seed-based conservation plays a critical role in the conservation, enrichment, and restoration of degraded environments. Climate change and agricultural pressure has enhanced the vulnerability to desertification of tropical dry forests. Plant biodiversity conservation is urgent, mainly in populations located in climate change threatening hotspots. Traditional stored seed quality assessment relies on destructive tests to evaluate viability, germination, and moisture content. These usually take more time and seeds than is convenient for restoration actions. Quick and accurate non-destructive methods allow efficient decision making in all native seed conservation activities, from harvest to storage, and also in seed-based restoration actions. Spectrometry allied to machine learning can be an effective method for evaluating the quality of native forest seeds, while maintaining the usually small bulk of stored seeds. Thus, we aimed to develop an easy and non-destructive method for evaluating stored seeds physiological quality and moisture content using a portable Vis-NIR spectrometer and a machine learning algorithm model. Stored and fresh seeds of different populations of two dry forest Fabaceae were individually evaluated for germination, moisture content and spectral data using a portable spectrometer. External and internal validation of the model were carried out by dividing the data into training (70%) and testing (30%) stages, using 10x cross-validation. The spectral data were processed using Weka 3.8.6 software. The discriminative algorithms applied were Support Vector Machine, Multilayer
<figure><img src="image_32.png" /></figure>

Perceptron, Random Forest and J48. For germination, the seeds were classified as YES (germinated) and NO (not germinated). Multilayer Perceptron algorithm obtained the best results for water content evaluation, with calibration and prediction correlation coefficients above 0.70. Random Forest and J48 algorithms showed best performances for seed germination classification. Although the algorithms were efficient in seed quality classification, increasing the number of sampled seeds may improve their prediction accuracy and aid seed-based conservation activities.

### S.5.4 Capturing representative genetic diversity in quality conservation germplasm collections

Marlien van der Merwe¹, Saphira Bloom-Quinn¹, Patricia Lu-Irving¹, Jason Bragg¹, Eilish McMaster¹, Maurizio Rossetto¹

1 Research Centre for Ecosystem Resilience, Botanic Gardens of Sydney, Sydney, Australia.

Seed banking of wild plants is a cost-effective conservation tool that allows for genetically representative germplasm collections to be stored for extensive periods of time. Such collections are becoming increasingly

more important particularly for species where anthropogenic activities are contributing towards local extinction and population decline, leading to a loss of genetic diversity and adaptive capacity. Seed collecting aimed at capturing genetic diversity across the range of wide-spread species can be daunting specially in parts of the world where resources are limited, and such germplasm collections are only now being initiated. Here we use a range of genomic datasets to provide guidelines for building representative conservation germplasm collections. Progeny datasets for multiple species that we obtained from seed collected across multiple sites and multiple maternal lines allowed us to explore how different collecting strategies (numbers of seed, families, and sites) capture genetic diversity. The results from this empirical data were encouraging indicating that a.) small quality collections can capture a large proportion of allelic variation and b.) we can use population genomic data to strategize seed collections to maximise genetic diversity. We apply the outcomes from the progeny analysis to multiple extensive population genomic datasets available through the Restore and Renew program and discuss guidelines for capturing various representative levels of allelic variation. Finally, based on these progeny data sets and other results from our teams' extensive work in taxonomy, conservation and restoration genomics, we prompt seed collectors and seed bank managers to maintain maternal lines during the life of the collection as this maximises the value and utility of these precious germplasm collections.

## S.6 NEW FRONTIERS IN PLANT PHYLOGENOMICS. SESSION 1

### S.6.1 New Views on Fern Phylogenetics. Updates from the GoFlag Targeted Enrichment Probe Set

Emily B. Sessa¹, Jessie A. Pelosi², Susan Fawcett³, J. Gordon Burleigh²

1 William & Lynda Steere Herbarium, New York Botanical Garden, Bronx, NY, USA. 2 Department of Biology, University of Florida, Gainesville, FL, USA. 3 University & Jepson Herbaria, University of California, Berkeley, CA, USA.

Ferns are the second largest group of vascular land plants, with ca. 10,000, species, and they are critical components of Earth's biodiversity — ferns can be found in nearly every type of ecosystem and habitat, from desert to rainforest. Ferns also occupy a pivotal evolutionary position as sister to the megadiverse seed plants, and they are thus the critical outgroup needed to understand the evolution of key seed plant features. Fern phylogenetics as a field has evolved considerably since the 1990s, when the first rbcL-based phylogenetic analyses of ferns were completed. Fern phylogenetics has presented researchers with a number of challenges, including the deep evolutionary history of the group and their propensity to undergo hybridization and whole genome duplication. We present a phy-
<figure><img src="image_33.png" /></figure>

logenomic approach to fern evolution, using the most highly resolved nuclear dataset to date (targeting 408 loci), and with highly targeted taxonomic sampling (including nearly all fern families and genera). This dataset allows us to explore a range of outstanding questions in fern phylogenetics, including resolving re-calcitrant nodes, and comparing results between our nuclear based trees and other recent large-scale fern phylogenies based on chloroplast loci.

## S.6.2 Completing the Angiosperm Tree of Life

William J. Baker¹, the PAFTOL Community

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

The tree of life is the fundamental biological roadmap for navigating the evolution and properties of life on Earth, and yet remains largely unknown. Even angiosperms are fraught with data gaps, despite their critical role in sustaining terrestrial life. Since 2016, the Plant & Fungal Trees of Life (PAFTOL) project at the Royal Botanic Gardens, Kew has targeted this knowledge gap in angiosperms by leveraging biological collections with novel genomic tools at unprecedented scale. Here, we describe our efforts to complete the angiosperm tree of life for all genera. Key achievements include: (1) the creation of Angiosperms353, an innovative, open-access, target sequence capture toolkit for tree of life research on any flowering plant; (2) the assembly of Angiosperms353 sequence data for 8,336 (61%) of the 13,600 genera of flowering plants, amounting to over 1.5 billion base pairs in total; (3) a tree of life containing all 7,514 genera of flowering plants sequenced to date, integrating novel data for over 8,000 samples, with data sourced from collaborators and public resources; (4) the Kew Tree of Life Explorer (https://treeoflife.kew.org/) launched in February 2021, an open data portal through which our tree of life and all data are made publicly available, with release 3.0 (April 2023) containing data for 10,377 samples; (5) a collaborative network of >300 individuals worldwide driving numerous projects using Angiosperms353 data; (6) outreach activities communicating the science of PAFTOL to the public. A wealth of primary research has also arisen from our data ranging from species-level studies, through to big picture analyses exploring the rise of the angiosperms. As the project approaches its goal of sampling all angiosperm genera, we welcome collaboration from all corners of the community and encourage anyone with interests in working with us on data generation, sharing or utilisation to get in touch.

## S.6.3 Angiosperms353: Version 2: improving gene recovery and locus length for universal single copy genes in flowering plants

Matthew G. Johnson¹, Chris Jackson², Alexandre R. Zuntini³, Nan Hu¹, Paul Bailey³, Felix Forest³, Norman J. Wickett⁴, William J. Baker³

1 Department of Biological Sciences, Texas Tech University, Lubbock, USA. 2 Royal Botanical Gardens Victoria, Melbourne, Australia. 3 Royal Botanical Gardens Kew, London, UK. 4 Department of Biological Sciences, Clemson University, Clemson, USA.

Angiosperms353 has gained popularity both as informative single copy genes and universal probes for cost-effective target capture in high-throughput sequencing. With data from over 10,000 species and 60% of extant genera now represented in the Kew Tree of Life Explorer (treeoflife.kew.org), Angiosperms353 has become an important tool for phylogenetics and population genetics studies. However, several shortcomings of the original gene targets and associated probe sequences have led to poorer than average recovery of sequences in certain angiosperm groups. We seek to improve Angiosperms353 probes by using expanded resources of high-quality genome references in angiosperms and data generated by the Plant and Fungal Tree of Life (PAFTOL) project at RBG-Kew and the Genomes of Australian Plants (GAP) project at RBG-Victoria. For Angiosperms353v2, we expanded the targeted sequence of each gene to include coding regions not captured in the first version. We also improved the regions covered in Angiosperms353v1 by using kmedioids clustering to select represented sequences at a per-exon level. We describe the in-silico testing of Angiosperms353v2 and associated resources, including curated clade-specific target files for use in gene recovery workflows. Our new probe design is intended to continue the success of Angiosperms353 with improved recovery and increased gene sequence length, thus enhancing future phylogenetic inference.
<figure><img src="image_34.png" /></figure>

### S.6.4 Integrative phylogenomics of the model tropical rain forest family Annonaceae: from herbaria to global scale diversity dynamics

Thomas L.P. Couvreur¹; Carlos Rodrigues-Vaz¹,²; Vincent R.C. Soulé¹; Laura Holzmeyer¹; Francis J. Nge¹; Serafin J. R. Streiff¹, Annonaceae Global Phylogenetics Consortium³

1. DIADE, Univ Montpellier, CIRAD, IRD, Montpellier, France. 2. Institut de Systématique, Evolution, Bio-diversité, Muséum National d'Histoire Naturelle-CNRS-SU-EPHE-UA, Paris, Cedex 05, France. 3. http://www.couvurlab.org/agpc.html.

Annonaceae is a major pantropical family with 2450 known species mainly occurring in rain forests worldwide. Based on decades of taxonomic research and resource generation, Annonaceae provide an excellent model-clade to understand the evolution of hyperdiverse tropical rain forests. We present ongoing integrative research combining phylogenomics, evolution and ecology of Annonaceae. Using a custom Annonaceae baiting kit, we sequenced hundreds of nuclear genes for over 85% of its diversity worldwide (> 2000 species). Over 75% of species were sequenced from herbarium specimens, the oldest dating from 1785. In parallel, we produced a massive morphological dataset documenting functional diversity, generated precise species-level distribution based on herbarium specimens documenting spatial diversity, and finally, used innovative near infrared scanning (NIRs) to estimate species specific spectra documenting chemical diversity. Most relationships were well resolved producing one of the first ever near-complete robust species level phylogenetic hypotheses of a major tropical rain forest plant family. The results mainly confirmed past systematic studies in Annonaceae, but also produced some surprises. By integrating these large-scale datasets with a robust dated phylogenetic tree at the species level, we present preliminary results about global rain forest diversification. Finally, we discuss the advantages and challenges of leading a large-scale project on a model clade, and how herbaria are (and will be) central to tackling numerous questions about rain forest evolution.

### S.6.5 A phylogenomic analysis of Salsoloideae (Amaranthaceae). An example of extreme paralogy and discordance

Diego F. Morales-Briones¹, Gudrun Kadereit¹

1. Ludwig-Maximilians-Universität München, Munich, Germany.

Caroxyloneae and Salsoleae constitute a species-rich lineage of Amaranthaceae containing around 40 genera and 300-400 species, including the largely polyphyletic *Salsola*. Members of this lineage are primarily distributed in the steppes and deserts of Eurasia, northern and southern Africa. It includes leaf- and stem-succulent halophytic, xerohalophytic, xerophytic, and ruderal plants, where C₄ photosynthesis is prominent. Generic boundaries in Caroxyloneae and Salsoleae have been historically a matter of long-standing debates, with numerous transfers, reinstatements, or newly established genera in recent years. To date, phylogenetic analyses of the tribe have been based on a limited number of DNA regions, leaving most phylogenetic relationships unclear in an already highly taxonomic challenging group. To build a comprehensive backbone phylogeny of the group to clarify generic boundaries and investigate patterns of C₄ photosynthesis, we used 483 putative single-copy loci from a custom target enrichment bait set and sampled 175 spp. from 36 genera from Caroxyloneae and Salsoleae. We recovered many paralogs for all species, including many pseudo-gene copies. These paralogs were spread across all Chenopods II, with several species even having gene copies as far as nested within Polycnemoideae (one of the other subfamilies in Amaranthaceae). This extreme paralog pattern largely halted the ability to infer orthologs using tree-based methods because the many duplications found in the gene trees resulted in mainly small orthologs, which ultimately resulted in a phylogeny with massive discordance and low to no support for most nodes. Here, we explore the challenges and strategies for overcoming the issue of rampant paralogy to attempt to build a backbone of Caroxyloneae and Salsoleae.
### S.6.6 Applying the phylogenomics package Captus to polyploid, contaminated, and degraded samples

Edgardo M. Ortiz¹, Gentaro Shigita¹,², Alina Höwener³, Mustafa Raza¹, Hanno Schaefer¹

1 Technical University of Munich, Freising, Germany.
2 University of Tsukuba, Tsukuba, Japan. 3 Ludwig Maximilian University of Munich, Munich, Germany

Our recently developed software package, Captus (available at https://github.com/edgardomortiz/Captus), greatly simplifies processing high-throughput sequencing data into alignments ready for phylogenetic analyses. Captus can remove paralogs from these alignments for use in traditional species tree estimation methods or include the complete set of recovered paralogs for use in more recent methods such as ASTRAL-Pro. Even though Captus normally searches for previously known sets of orthologs (e.g., Angiosperms353, BUSCO lineage databases) it can also be used to discover new putative homologs across the samples analyzed, making it

possible to extract phylogenetically informative loci from any group of organisms. Besides recovering accurately and rapidly a greater number of more complete loci across more samples, Captus is also capable of dealing with several common issues that arise in current phylogenomics practice. We show that the accuracy of recovery helps minimizing the methodological sources of conflict between gene trees and the species tree revealing clear patterns of reticulation or hybridization. Captus facilitates the combination of sequence data from samples of varying quality (i.e., silica-preserved, herbarium), different data types (i.e., target capture, high and low depth whole genome sequencing, and RNA-Seq), and pre-assembled genomes and transcriptomes to build a single phylogenomic dataset for analysis. Captus can also be used to identify outlier samples containing an excess of paralogs due to polyploidy or potentially contamination. Captus can be used to remove phylogenetically distant contaminants from a sample. Finally, we show an additional feature of Captus, an easy-to-use workflow to find putative homologs and design baits for target enrichment and how we used it to create a set of baits to capture 989 protein coding genes (854 single-copy and 135 associated with morphological traits of interest) as well as 177 non-coding regions for the family Cucurbitaceae.

## S.7 MECHANISMS AND CONSEQUENCES OF DISPERSAL IN LAND PLANTS: TOWARDS A MORE UNIVERSAL UNDERSTANDING. SESSION 1

### S.7.1 Seed dispersal mismatches under global change: what can we apply to the rest of terrestrial plants?

Irene Mendoza¹

1 Estación Biológica de Doñana (CSIC), Sevilla, Spain

Seed dispersal implies the movement of a seed at a certain distance from the maternal plant, which favours propagation towards new environments. Many Spermatophyta rely on animals for seed dispersal, especially those with fleshy fruits which offer

a nutritious award for the disperser. The unprecedented environmental change produced by human activities (including climate and land-use changes) is not only reducing biodiversity in terms of the number of species, but it is also changing the probabilities of interactions among plants and their dispersers. Seed dispersal by avian species necessarily implies that the phenology of birds and fruit production matches in time. Whereas we have evidence that species are already changing their geographic distribution ranges and their life cycles' timing (e.g., bird migration, fruiting time) as a consequence of climate change, we still have a knowledge gap in understanding whether these changes will result in mismatches among interacting species. Network analytical advances and new technologies
<figure><img src="image_35.png" /></figure>

applied to biodiversity monitoring have paved the road to undertake this challenging objective. In this presentation, I focus on these specific questions 1) What phenological changes are currently taking place in Southern Spain as a consequence of global change? 2) Are these phenological shifts resulting in a mismatch between the phenology of birds and fleshy-fruited plants? 3) How can we forecast phenological mismatches? For answering these questions, I use methods from classical field monitoring and new technologies such as digital images from phenocams or unmanned aerial vehicles that have barely been applied to research on ecological interactions, combined with cutting-edge analytical tools including network analyses and forecasting modeling. These methods and conceptual advances can be easily extended to other terrestrial plants and will help understanding the ultimate consequences for ecosystem functioning and biodiversity conservation of seed dispersal mismatches induced by global change.

## S.7.2 How can dispersal by endozoochory by waterbirds influence the genetic structure of a dry-fruited wetland plant?

M. Angeles Ortiz¹, Raul Sanchez², Iciar Jim², Karin Tremetsberger³, Casper van Leeuwen⁴, Adam Lovas-Kiss⁵, Andrew J. Green².

1 Dpto. Biología Vegetal y Ecología, Universidad de Sevilla, Spain 2 Dept of Wetland Ecology, Doñana Biological Station EBD-CSIC, Sevilla, Spain. 3 Institut für Botanik, Universität für Bodenkultur Wien, Wien, Austria 4 Netherlands Institute of Ecology (NIOO-KNAW), Wageningen, The Netherlands 5 Wetland Ecology Research Group, Centre for Ecological Research, Danube Research Inst., Debrecen, Hungary

One of the main challenges when reconstructing the evolutionary history of a plant species is the lack of information about its dispersal mechanisms. These mechanisms shape the distribution of plants, favoring long-distance dispersal events or confining them to much more restricted areas. Some species exhibit obvious dispersal syndromes, like plants with fleshy fruits. However, plants without clear modifications in their fruits or seeds promote controversy regarding whether they have a specialized dispersal system, fall passively near the mother plant, or

depend on random events to colonize new habitats. A specific case involves dry-fruited plants in wetlands, traditionally assumed to have hydrochorous dispersal due to their distribution in or near waterbodies. Recent studies have shown that aquatic birds can disperse dry-fruited plants through endozoochory, after feeding on the plants or in the seed bank, egesting viable seeds in their faeces. But how does this type of dispersal affect the genetic structure of plants? Does it promote long-distance dispersal along migratory flyways, and/or connectivity between nearby wetlands?

We will test the dispersal processes shaping the population structure and gene flow in Eleocharis palustris (Cyperaceae). This species inhabits wetland borders and temporarily flooded areas, being native to the Northern Hemisphere. Endozoochory of this species has been recorded for many different waterbird species. Morphological and ploidy level studies have been conducted to characterize its variability. To delve into this topic, population genetics was investigated applying GBS (genotyping by sequencing) in 188 plants from 20 populations in seven European countries. Finally, the relationship between the migratory flyways and the population networks will be discussed, as well as the role of their ecological niche. The understanding of how endozoochory can shape the distribution of a species could be key to management and conservation policies for native taxa.

## S.7.3 Autorotating diaspores of flowering plants: an inspiring strategy in wind dispersal

Drisya V.V.¹, C. Pramod¹,², A.K. Pradeep², T.P. Suresh³

1 Department of Botany, Government Brennen College, Dharmadam P.O., Thalassery, Kerala, India. 2 Department of Botany, University of Calicut, Calicut University P.O., Kerala, India. 3 Department of Physics, Government Brennen College, Dharmadam P.O., Thalassery, Kerala, India

Plants are excellent source of inspiration for design innovation as they are already figured out what works to spread over the planet and provide numerous examples of stable design and specialized functionality that are worth to study (Pandolfi & Izzo, 2013). One amazing demonstration of 'intelligent' behavior of plants for wind
<figure><img src="image_36.png" /></figure>

dispersal is the production of autorotating diaspores. Dispersal units, such as fruits, seeds, and other structures which constantly rotate in the air during dispersal basically due to aerodynamic forces for moving away from the parent plant to establish the progeny as an independent sapping are known as autorotating diaspores. Based on the morphology and type of rotation during the fall, three types of autorotating diaspores are identified: autogyros, rolling autogyros, and helicopters (Augspurger, 1986). In a study conducted on the winged diaspores of the flowering plants of South India, a total of about 120 species (55 autogyros, 24 helicopters, 41 rolling autogyros) were recorded, which produce autorotating diaspores with well-defined morphological adaptations. Macromorphological, micromorphological, developmental, anatomical, flight behavioural, and aerodynamic studies of a subset of diaspores from each category show that each of them varies in its origin, pattern of development, and configuration of the wings, which account for the variation in the overall morphology and eventually the dispersal potential. The autorotating diaspores of the flowering plants of South India display a variety of morphology and design for the efficient mechanism of dispersal, also effective as natural inspiration for design innovation. The present paper aims to provide a better understanding of the morphology, geometry, underlying mechanisms of flight, specificity of the direction of rotation, and biomimetic application of the autorotating diaspores of flowering plants of south India.

### S.7.4 Linking pollen limitation and seed dispersal effectiveness

Laura C. Leal¹,², Matthew H. Koski²

1 Universidade Federal de São Paulo, Diadema, São Paulo, Brazil. E-mail: laura.leal@unifesp.br. 2 Clemson University, Clemson, South Carolina, USA

Seed production and dispersal are crucial ecological processes impacting plant demography, species distributions, and community assembly. Plant-animal interactions commonly mediate both seed production and seed dispersal, but current research often examines pollination and seed dispersal separately, which hinders our understanding of how pollination services affect downstream dispersal services. To fill this gap, we propose a

conceptual framework exploring how pollen limitation can impact the effectiveness of seed dispersal for endozoochorous and myrmecochorous plant species. We summarize the quantitative and qualitative effects of pollen limitation on plant reproduction and use Optimal Foraging Theory to predict its impact on the foraging behavior of seed dispersers. In doing so, we offer a new framework that poses numerous hypotheses and empirical tests to investigate links between pollen limitation and seed dispersal effectiveness and, consequently, post-dispersal ecological processes occurring at different levels of biological organization. Finally, considering the importance of pollination and seed dispersal outcomes to plant eco-evolutionary dynamics, our framework opens new avenues for future studies exploring the current and future functioning of these key interactions, both deeply threatened by human landscape modification and ongoing climatic changes.

### S.33.3 How can dispersal by endozoochory by waterbirds influence the genetic structure of a dry-fruited wetland plant?

M. Ángeles Ortiz¹, Raul Sanchez², Iciar Jiménez², Karin Tremetsberger³, Casper van Leeuwen⁴, Adam Lovas-Kiss⁵, Andrew J. Green²

1 Departamento de Biología Vegetal y Ecología, Universidad de Sevilla, Spain. 2 Dept. of Wetland Ecology, Doñana Biological Station -CSIC, Sevilla, Spain. 3 Institut für Botanik, Universität für Bodenkultur Wien, Wien, Austria. 4 Netherlands Institute of Ecology (NIOO-KNAW), Wageningen, The Netherlands. 5 Wetland Ecology Research Group, Centre for Ecological Research, Danube Research Inst., Debrecen, Hungary

One of the main challenges when reconstructing the evolutionary history of a plant species is the lack of information about its dispersal mechanisms. These mechanisms shape the distribution of plants, favoring long-distance dispersal events or confining them to much more restricted areas. Some species exhibit obvious dispersal syndromes, like plants with fleshy fruits. However, plants without clear modifications in their fruits or seeds promote controversy regarding whether they have a specialized dispersal system, fall
<figure><img src="image_37.png" /></figure>

passively near the mother plant, or depend on random events to colonize new habitats. A specific case involves dry-fruited plants in wetlands, traditionally assumed to have hydrochorous dispersal due to their distribution in or near waterbodies. Recent studies have shown that aquatic birds can disperse dry-fruited plants through endozoochory, after feeding on the plants or in the seed bank, egesting viable seeds in their faeces. But how does this type of dispersal affect the genetic structure of plants? Does it promote long-distance dispersal along migratory flyways, and/or connectivity between nearby wetlands? We test the dispersal processes shaping the population structure and gene flow in *Eleocharis palustris* (Cyperaceae). This species inhabits wetland borders and temporarily flooded areas, being native to the Northern Hemisphere. Endozoochory of this species has been recorded for many different waterbird species. Morphological and ploidy level studies have been conducted to characterize its variability. To delve into this topic, population genomics was investigated applying GBS (genotyping by sequencing) in 188 plants from 20 populations in seven European countries. Finally, the relationship between the migratory flyways and the population networks will be discussed, as well as the role of their ecological niche. The understanding of how endozoochory can shape the distribution of a species could be key to management and conservation policies for native taxa.

### S.7.5 Pollen dispersal in flowering plants. Importance of pollinators from single visits to regional patterns

Javier Valverde¹

1 Universidad de Sevilla, Sevilla, Spain

Pollen dispersal in flowering plant species results from the joint action of the pollinators at each plant population. These dispersal agents determine the mating patterns between plants and ultimately the genetic composition of the populations as well as their degree of genetic isolation. Here, results from a series of studies will be introduced to highlight the importance of pollinators for the genetic dispersal of flowering plants at different scales. At the level of individual plants, a study demonstrates how pollinators can widely differ in their capacity of transporting pollen of different genetic diversity and origin. At the population level, a study is used to evidence the role of these animals in generating contemporary patterns of pollen flow between plants. At the regional level, a study performed on several plant species demonstrates how pollinator diversity correlates with the genetic diversity and connectivity of plant populations. Taken together, these studies demonstrate that addressing the mechanisms of pollen flow is essential to explain the genetic patterns found within and between plant populations that ultimately provide insights into plant evolution.

## S.8 SAPINDALES: UNDERSTANDING ANGIOSPERM EVOLUTION BY INTEGRATION ACROSS DATA, SPACE AND TIME

### S.8.1 Hybrid Capture resolves the phylogeny of the pantropically distributed Zanthoxylum (Rutaceae) and reveals an Old World origin

Lee Ping Ang¹, Salvatore Tomasello¹, Jun Wen², Marc S. Appelhans¹,²

1. Department of Systematics, Biodiversity and Evolution of Plants, Albrecht-von-Haller Institute of Plant Sciences, University of Goettingen, Untere Karspuele 2, Goettingen, Germany. 2. Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

With about 225 species, *Zanthoxylum* L. is the second most species rich genus in Rutaceae. It is the only genus with a pantropical distribution. Economically, it is used in several Asian countries as traditional medicine and spice. In the past *Zanthoxylum* was
<figure><img src="image_38.png" /></figure>

divided into two genera, the temperate Zanthoxy-
lum sensu strictu (s.s.) and the (sub)tropical Fagara,
due to the large differences in flower morphology:
heterochlamydeous in Fagara and homochlamyd-
eous in Zanthoxylum s.s. This genus is much under
studied and previous phylogenetic studies using
Sanger sequencing did not resolve the relationships
sufficiently. In this study, we use Hybrid Capture with
a specially designed bait set for Zanthoxylum to se-
quence 347 putative single-copy genes. The taxon
sampling has been largely improved as compared
to previous studies and I present preliminary results
based on 371 specimens representing 133 species
from all continents and major island groups. Our
preliminary results reveal similar tree topology as
the previous studies while providing more details to
the backbone of the phylogeny. The phylogenetic
tree consists of four main clades: a) African/Mala-
gasy clade, b) Z. asiaticum clade - a clade consist-
ing widespread species occurring in (sub)tropical
Asia and Africa as well as Madagascar, c) Asian/
Pacific clade and d) American clade, which also in-
cludes the temperate Asian species. The merging of
Fagara and Zanthoxylum is supported by our results
and the homochlamydeous flowers of Zanthoxylum
s.s. are likely derived from heterochlamydeous flow-
ers. Several of the morphologically defined sections
within Zanthoxylum are not monophyletic. In this
presentation, we (1) introduce the framework of this
project; (2) present preliminary results and (3) the
ongoing progress of the study.

S.8.2 Systematics and
morphological diversity
of the cashew family
(Anacardiaceae)

Susan K. Pell¹, John D. Mitchell²

1 United States Botanic Garden, Washington, USA.
2 The New York Botanical Garden, New York, USA

Anacardiaceae is an ecologically and economically important plant family of about 800 species in more than 80 genera worldwide. Most well-known for its commercial crops, cashew (Anacardium occidentale), mango (Mangifera indica), and pistachio (Pistacia vera), and its many contact dermatitis-causing species (e.g., poison ivy and oak; Toxicodendron spp.), the family includes many other regionally important taxa and exhibits substantial morphological diversity. Anacardiaceae is particularly diverse in leaf architecture, flower and

fruit morphology, and pollination and dispersal syn-
dromes. Fruit diversity is the primary reason 30 of the
genera are monotypic and that so many genera are
recognized in the family. Recently published Sapin-
dales phylogenies and newly discovered Anacardi-
aceae fossils have shed new light on the evolution
of structural diversity in the family. This presentation
provides an overview of the noteworthy morpho-
logical diversity of Anacardiaceae in the context of
systematics and paleobotany. Several newly recog-
nized taxa, including newly published and yet-to-be
published genera, are discussed.

S.8.3 Meliaceae (mahogany family): model group for a better understanding of tropical angiosperm evolution?

Alexandra N. Muellner-Riehl¹,², Meliaceae
Taxonomic Expert Network

1 Leipzig University, Leipzig, Germany. 2 German Centre for Integrative Biodiversity Research, Halle-Jena-Leipzig, Leipzig, Germany

Meliaceae comprises woody plants widely dis-
tributed throughout the tropics and subtropics,
extending into temperate zones. With over 740
species in 58 genera, they constitute a medi-
um-sized family in Sapindales. The Indo-Male-
sian region is the geographic centre of diversi-
ty. Africa-Madagascar is almost as diverse as
Indo-Malesia, followed by Australasia. Only eight
genera are present in the Neotropics, but they
are as diverse as the Africa-Malagasy region as
for the number of species. Comparing geological
evidence with the fossil record as well as bioge-
graphic studies, there is indication that the now-
adays pantropically distributed family has made
use of three routes to acquire its current distri-
bution: land connections (i) between Eurasia and
North America, (ii) between North America and
South America, and (iii) dispersal paths between
Africa and South America that have existed since
the proposed evolutionary origin of modern Meli-
aceae (Cretaceous). Although important new in-
sights into Meliaceae evolution have been gained
during the past two decades, there are also still
open questions. A careful revision of the entire
fossil record of Meliaceae will be an important
step into a more comprehensive understanding
of the biogeographic history of this family, which
may serve as a prime example for understanding
the evolution of rainforest (and related) taxa on
a global scale. A more or less completely sam-
pled species-level phylogeny, accompanied by
molecular clock dating making use of the rich
fossil record, complemented by biogeographic
reconstructions (incl. the investigation of niche
evolution), will only be achievable by a scientific
community effort, bringing together researchers
from different disciplines, who have a keen inter-
est in Meliaceae evolution. The vision of this lon-
ger-term research agenda will be to establish
this—also economically very important—family
as a model group for a better understanding of
angiosperm evolution.

S.8.4 The evolution of
Meliaceae: systematics,
genome duplications and
implications for taxonomy

Monika M. Iskander¹, Elizabeth M. Joyce²

1 Rhenish Friedrich Wilhelm University of Bonn, Germany. 2 Ludwig Maximilian University of Munich, Germany

Meliaceae is a widespread angiosperm family of c.
740 species in 58 genera, mainly comprising tropi-
cal and subtropical trees. Given its distribution and
diversity, Meliaceae is considered a useful lineage
for understanding the evolution of tropical forests;
however, our understanding of the family's evo-
lution and infra-familial relationships remain un-
clear. Here, we present the first genus-complete
phylogenies for the family, generated with Angio-
sperms353 and chloroplast loci, and including 139
taxa with good geographic representation. In the
nuclear trees, many relationships were retrieved
with maximum support (PP=1, BS=100) and sug-
gest that some genera are non-monophyletic and
should be recircumscribed, while some relation-
ships remained poorly supported or conflicted with
the topology of the chloroplast phylogeny, partic-
ularly in the Melioideae. Analysis and placement of
paralogs suggest that this poor resolution is likely
caused by a history of hybridisation and genome
duplication events. Our results indicate that this
may confound our ability to reconstruct relation-
ships in bifurcating trees, raising interesting ques-
tions about how we might classify tribes and gen-
era within the family.

S.8.5 Phylogeny of Galipeeae (Rutaceae, Sapindales), a group of the Citrus family. with insights of critical morphological traits.

Milton Groppo¹, Jacquelyn Ann Kallunki²,
Carla P. Bruniera³, José Rubens Pirani¹,
Carolina Ferreira¹, Paola de Lima Ferreira⁴

1 University of São Paulo, Brazil. 2 New York Botanical Garden, USA. 3 Federal University of São Paulo, Brazil. 4 Aarhus University, Denmark

Tribe Galipeeae (Rutaceae, Subfamily Zanthoxy-
loideae) is the largest group of Rutaceae in the
Neotropical Region, with 33 genera and c. 181 spe-
cies. The tribe is composed of two subtribes, Gali-
peeae (28 genera and c.130 species) and Pilocar-
pinae (5 genera, 51 species). Subtribe Galipeinae
is different from Pilocarpinae by the possession
of zygomorphic flowers (v. actinomorphic flowers
in Pilocarpinae) and by the constant presence
of staminodes (v. all fertile stamens in Pilocar-
pinae.) A phylogenetic analysis was conducted
to test the monophyly of Galipeeae and its two
subtribes. A total of 120 terminals, representing
24 genera and 80 species of both subtribes were
sampled. Species of Balfourodendron, Choisya,
Helietta, Hortia, and Plethadenia, all from gen-
era traditionally not included in Galipeeae were
also used; Zanthoxylum, a phylogenetically dis-
tant genus was used as outgroup in all analyses.
Molecular data included four regions, two from
the cpDNA (trnL-trnF and rps-16) and two nuclear
(ITS1 and ITS2) analyses. Parsimony and Bayes-
ian analysis was conducted. Results showed
all Galipeeae grouped in a strongly supported
clade (Bootstrap Percentages BP-100% BP, Pos-
terior Probabilities PP = 1); Subtribes Galipeeae
and Pilocarpinae appear as sisters, both clades
with strong support. Balfourodendron and Heliet-
ta appear as part of the Pilocarpinae and Hortia
along with the Galipeinae. Critical morphologi-
cal characteristics such as zygomorphic flowers,
presence of staminodes, embryo and fruit char-
acteristics are discussed in the light of the new
produced phylogeny.
# S.9 TOWARDS INTEGRATIVE TAXONOMY. THEORY AND PRACTICE. SESSION 1

## S.9.1 Integrated Taxonomy

R.W. Scotland¹

1 University of Oxford, Oxford, UK.

Tropical plants are in the most urgent need of taxonomic revision as the majority of tropical genera have never been monographed, that is they have not been the subject of detailed study across their entire geographical range. To complete the inventory of tropical flowering plants phylogenetics and monophyly are important but need to be integrated with nomenclature, description, morphology, geography, literature, interpretation of specimens, plant collectors and protologues. In the absence of this integration the inventory of tropical plants remains partial and problematic to tackle the biodiversity crisis.

## S.9.2 Taxonomic updates to Caribbean Pitcairnia (Bromeliaceae) based on genomic, morphological, and ecological data

Natalia Ruiz-Vargas¹, Roberta Mason-Gamer¹

1 Department of Biological Sciences, University of Illinois at Chicago, Illinois, United States

Pitcairnia is the second largest genus in the family Bromeliaceae, distributed from Mexico to northern Argentina, and the only one with presence in Africa. There are approximately 15 species reported for the Caribbean islands, most of which are island endemics; one species, Pitcairnia angustifolia, has wider distribution. Among the islands, Ayiti (Hispaniola), shared by the Dominican Republic and Haiti, stands out with six proposed species, while most of the islands have one. Using a target Sequence Capture approach with the Bromeliad 1776 baits set we analyze the relationships within the Caribbean group and found 1) that current species delimitation does not match the molecular data, and 2) that species diversity

has been overestimated, likely because of high phenotypic plasticity in the group. We analyzed morphological characters in the group to understand differences between taxa and intraspecific variation. Ecological factors that influence morphological variations and promote or inhibit isolation are also analyzed.

## S.9.3 Carex sect. Fecundae: a boreotemperate-origin adaptive radiation in the Neotropics

Raúl Lois¹, Anton A. Reznicek², Socorro González Elizondo³, Eduardo Ruiz Sánchez⁴, Marcial Escudero⁵, Ana Morales Alonso⁶, Carmen Acedo¹, Pedro Jiménez-Mejías⁷

1 Universidad de León, Spain. 2 University of Michigan Herbarium, USA. 3 Instituto Politécnico Nacional, CIIDIR Durango, Mexico. 4 Universidad de Guadalajara, Mexico. 5 Universidad de Sevilla, Spain. 6 Universidad Rey Juan Carlos, Spain. 7 Universidad Pablo de Olavide, Spain

Carex, a member of the Cyperaceae family, stands as a megadiverse genus encompassing over 2000 recognized species, primarily thriving within the boreal and temperate zones of the northern hemisphere. Despite this, certain lineages of Carex have successfully ventured into the tropical zones of the southern hemisphere on multiple occasions. Notably, Carex sect. Fecundae emerges as one such lineage, displaying a distribution entirely within the Neotropics, predominantly within high-altitude forests and grasslands spanning from Central Mexico to Northern Argentina. Comprising more than 30 species, Carex sect. Fecundae stands as one of the broadest lineages within Carex. It boasts perhaps the most intricate inflorescence patterns among its counterparts. However, akin to numerous Neotropical plant groups, the understanding of sect. Fecundae remains significantly limited. Only three treatments of this section exist to date, none of which are comprehensive. Our research addresses this gap through a comprehensive taxonomic revision of Carex sect. Fecundae, reviewing more than 1000
<figure><img src="image_39.png" /></figure>

specimens and statistically testing more than 100 characters. We established morphometric boundaries for species within the section by selecting the most distinguishing characters, such as inflorescence conformation, utricle, glumes, and achene dimensions. Furthermore, we undertook the first complete molecular phylogeny of the section, employing HybSeq to unravel the evolutionary relationships within *Carex* sect. *Fecundae*. Our combined analysis of morphological and molecular data led to the discovery of 18 previously overlooked species within the section. Additionally, we dated the recent origin of this section to approximately 5.77 Mya, affirming the rapid adaptive radiation witnessed within *Carex* sect. *Fecundae*. Through biogeographical analysis, we inferred the most recent common ancestor of the section between the Neotropical and South American Transition Zones. This origin was followed by two independent rapid radiations—one in Mexico and the other in South America—further elucidating its evolutionary history.

### S.9.4 Heuristics, species, and the analysis of systematic data - or why morphology is still king

Tom Wells¹,²

¹Herbarium, Royal Botanic Gardens, Kew, UK. ²Biology Department, University of Oxford, UK

The rise of molecular phylogenetics and “integrative taxonomy” have led to a sense that disagreements about the fundamental nature of the species category are no longer an issue in contemporary systematics. The implicit consensus is that increases in the volume, type, and precision of data and methods available to contemporary researchers will automatically lead to more accurate and consistent species delimitations, without the need for a widely accepted definition of what species are and how they should be identified and used in biological research. The chief challenge to this triumphalist view of ‘integrative taxonomy’ remains the heterogenous nature of speciation. Since speciation proceeds over extended periods of time and through different processes, it results in the inconsistent manifestation of different ‘contingent properties’ for its recognition, depending on the taxa in question. Simply combining multiple forms of data or methods under an ‘integrative approach’ cannot automatically overcome this fundamental obstacle because

of the continued lack of an overarching theoretical framework for species delimitation and how to treat incongruence between different types of Systematic data. Deciding what properties to assess, how to do so, and which to prioritise in the event of incongruence remains an unresolved issue, requiring a theoretical framework within which to make those decisions. In this talk, I explore how treating species as a “heuristic” can provide the basis of just such a framework. Species delimitation inherently involves the use of heuristics – through the study of representative specimens, the use of selected diagnostic characters, and strategic sampling of “tree space”. Meanwhile, species themselves are routinely used in a heuristic manner to investigate broader patterns of biodiversity and the processes underpinning them. Acknowledging these aspects of systematic research can therefore inform both operational ways of accurately and consistently delimiting species, and theoretical ideas about their nature.

### S.9.5 Challenges in defining genera: the limits of integrative taxonomy in the Funariaceae (Bryophyta)

Rafael Medina¹, Matthew G. Johnson², Nikisha Patel³, Nicholas Wilding⁴, Yang Liu⁵, Bernard Goffinet⁵

¹Departamento de Biodiversidad, Ecología y Evolución. Universidad Complutense de Madrid. Madrid. ²Department of Biological Sciences. Texas Tech University. Lubbock TX. USA. ³Department of Biology. Trinity College. Hartford CT. USA. ⁴UMR PVBMT, Pôle de Protection des Plantes. University of La Réunion. Saint-Pierre, Réunion. France. ⁵Department of Ecology and Evolutionary Biology. University of Connecticut. Storrs CT. USA

Traditionally, morphology played a pivotal role in shaping bryophyte taxonomy until the advent of the molecular phylogenetic revolution, substantially impacting their systematics. The integration of morphology and sequence data, while occasionally feasible, often remains elusive in select groups, likely due to the constrained morphospace of their anatomy and the prevalence of homoplasy. Here, we delve into the transformative influence of molecular investigations on classical generic concepts within the moss family Funariaceae. Despite the general conservation of moss sporophyte architecture driven by selective pressures for efficient spore dispersion, the Funariaceae exhibit an exceptionally diverse sporophytic gen-
eration. Historically, generic assignment relied on spe-
cific trait combinations, resulting in the recognition of
17 genera, including three specious ones—*Funaria*, *En-
tosthodon*, and *Physcomitrium*—and 14 paucispecific
genera. Many of the latter were established to reflect
intuitively unique morphological variations, typified by
single or very few species, such as for *Physcomitrella*,
harboring the model taxon *P. patens*. However, a series
of molecular phylogenetic studies at different scales
has substantially reshaped the taxonomic landscape
within the Funariaceae. These investigations have
unveiled novel insights: For example, most highly re-
duced sporophyte morphologies reflect a recurrent
syndrome within the family, lacking predictive value
of systematic affinities. Secondly, certain taxa arise
from hybridization events, exhibiting combinations of
morphological traits that may or may not align with
the progenitor species. Additionally, what seemed like
a coherent lineage, *Entosthodon*, reveals itself as two
separate groups (*Entosthodon* s.s. and *Physcomi-
trellopsis*) diverging 30 mya. While proposing a novel
generic synopsis consolidating the family's diversity
into seven genera was feasible, achieving a complete
integration between morphology and molecular sys-
tematics remains a partially realized goal. The refine-
ment of Funariaceae taxonomy remains an ongoing
challenge, prompting future inquiries to uncover over-
looked synapomorphies that complement the evolu-
tionary narratives elucidated by molecular data.

S.9.6 Modern taxonomic re-
visions: example of our ap-
proach in the big pantropical
genus Solanum (Solanaceae)

Tiina Särkinen¹, Rebecca Hilgenhof¹,
Edeline Gagnon², Sandy Knapp³, Leandro
Giacomin⁴

1 Tropical section, Royal Botanic Garden Edinburgh,
Scotland, UK. 2 Department of Integrative Biology,
University of Guelph, Ontario, Canada. 3 Life Sci-
ences Division, Natural History Museum, London, UK.
4 Department of Systematics and Ecology, Univer-
sidade Federal da Paraíba, Paraíba, Brazil

Agriculture has benefited from the large mor-
phological diversity found within the large genus
*Solanum* through the exploitation of tubers (po-
tatoes) and fruits (tomatoes, eggplants, and sev-
eral minor crops). We present an overview of how
taxonomic work on *Solanum* has developed over
the past decades, highlighting what tools and
approaches have been useful at different stag-

es and how "integrative taxonomy" has helped
our work. Our main aim is to provide a classifi-
cation framework for the megadiverse and ag-
riculturally important genus to be used across
research fields, including taxonomically authori-
tative molecular phylogenies, updated list of ac-
cepted species names and synonyms, and mor-
phological descriptions of all species. In our view,
phylogenies are important but the main core of
our work is around morphology, nomenclature
and field work as molecular sequence data (in-
cluding full reference genomes) are being pub-
lished more widely by non-taxonomists. We are
passionate to expand the number of taxonomic
specialists working in the genus across different
parts of the world, and to provide interactive and
visual identification tools that enable botanists to
identify *Solanum* species with higher confidence.
Our taxonomic expertise enables us to collabo-
rate with evolutionary developmental biologists
and genomics experts to understand the genetic
control of distinct morphologies in the group and
our understanding of evolution across *Solanum*.
Core part of this work is using our database of
taxonomically verified herbarium specimens from
across >400 herbaria to characterise species and
traits in ecological and geographical space to
better understand their biology and function.
<figure><img src="image_40.png" /></figure>

**S.10 FLORAL SPECIALIZATION AND THE**
**GENERALISED NATURE OF POLLINATION**
**INTERACTIONS. SESSION 1**

**S.10.1 Flowers on an adaptive**
**plateau: which floral traits**
**mediate specialization**
**in buzz-pollinated**
**Melastomataceae?**

Benjamin Lazarus¹, Ashlee Kerber², Agnes S. Dellinger¹

1 Department of Botany and Biodiversity Research, University of Vienna, Austria. 2 Ecology and Evolutionary Biology, University of Colorado Boulder, USA.

While the ecology and evolution of plant-pollinator interactions has often been studied through the comparison of closely-related species differing in functional pollinator groups (e.g., bees and birds), many plant lineages exhibit a single pre-dominant pollination strategy. Buzz-pollination is one such strategy, where, in the case of the plant family Melastomataceae, more than 5000 species are adapted to pollination by bees capable of producing specialized vibrations on flowers. While many buzz-pollinated flowers across angiosperms converge into a single main phenotype ('Solanum-type' flowers), Melastomataceae exhibit highly diverse flowers differing in color and scent profiles, corolla shapes, sizes, and the architecture and biomechanics of stamens. To date, we know comparatively little about how unbalanced patterns of flower diversity and uniformity arise across plant lineages, and how they play out in communities of co-flowering species. In this talk, we introduce patterns of macroevolutionary diversity of Melastomataceae flowers, and present new results on how flower traits may contribute to pollination niche partitioning and specialization despite substantial overlap in buzzing bee pollinators among closely-related, co-flowering Melastomataceae species. We compare trait and pollinator data across selected Melastomataceae communities from the South-Eastern USA, Ecuador and Brazil, covering a broad range of floral phenotypes and phylogenetic diversity of the group. By analyzing the links between buzz-pollination niche specialization, flower

traits commonly considered important in pollinator attraction (scent, color, reward) and pollen transfer (flower-pollinator fit), we aim at better resolving the processes underlying the evolution of floral diversity within pollination strategies.

**S.10.2 Pollinator shifts, floral**
**evolution, and geographic**
**range evolution in the**
**genetic divergence of three**
**Aeschynanthus** species

Jing-Yi Lu¹, Yaowu Xing², Hong Truong Luu³, Ming-Xun Ren⁴, Shao-Jun Ling⁴, and Richard H. Ree¹,⁵

1 Committee on Evolutionary Biology, University of Chicago, Chicago, USA. 2 CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, China. 3 Southern Institute of Ecology, Vietnam Academy of Science and Technology, Ho Chi Minh City, Vietnam. 4 Center for Terrestrial Biodiversity of the South China Sea, Hainan University, Haikou, China. 5 Negaunee Integrative Research Center, Field Museum, Chicago, USA

How do plants evolve under a geographic mosaic of pollinators? The Grant-Stebbins model predicts that local adaptation to different pollinators may drive ecological speciation. We test this hypothesis in the SE Asian genus *Aeschynanthus*, whose 160 species mostly overlap with their putative pollinators, nectar-specialist sunbirds. A conspicuous exception is *A. acuminatus*, a species widespread across mainland East Asia that also occurs in Taiwan, beyond the range of sunbirds. In Taiwan, *A. acuminatus* is exclusively pollinated by a mixed group of generalist passerines. Less is known about its pollination in mainland Asia, where both sunbirds and generalist passerines are available. In this study, we investigate the roles of pollinator shifts and range expansions in the genetic divergence of *A. acuminatus* from its closest relatives, *A. moningeriae* and *A. pedunculatus*. Pollinator obser-
<figure><img src="image_41.png" /></figure>

vations of *A. acuminatus* across its range showed varied degrees of visitation by both sunbirds and generalist passerines on mainland Asia. In contrast, its closest relatives are both exclusively pollinated by sunbirds. Genome-wide SNP analysis revealed a mainland origin in *A. acuminatus*, preceding its range expansion to Taiwan. The sunbird specialists have independent origins: *A. moningeriae* is nested within *A. acuminatus* and *A. pedunculatus* is reciprocally monophyletic. The origin of *A. acuminatus* was thus likely on the mainland, coincident with a shift from specialized sunbird pollination to a more generalized pollination system that facilitated its colonization of Taiwan. *A. moningeriae* represents a localized reversal to sunbird specialization on Hainan. The evolutionary history of *A. acuminatus* and its closest relatives was not consistent with the classic Grant-Stebbins model. Instead, our research exemplifies how multiple transitions between generalized and specialized pollination systems can influence the genetic divergence and range dynamics, and the potential for speciation.

## S.10.3 Pollination niche partitioning promotes coexistence of specialists in a Neotropical plant community

Simone Cappellari-Rabeling¹,⁴, Rosana Tidon², John L. Neff³ and Beryl B. Simpson⁴

1 Department of Plant Evolutionary Biology, Institute of Biology, University of Hohenheim, Stuttgart, Germany.

2 Department of Genetics and Morphology, University of Brasília, Brazil. 3 Central Texas Mellitological Institute, Austin, TX, USA. 4 Department of Integrative Biology, The University of Texas at Austin, USA.

Neotropical habitats harbor over 40% of the world's angiosperm diversity including several plant families with specialized pollination systems such as the Orchidaceae, Malpighiaceae, and the Plantaginaceae. The Malpighiaceae is considered one of the most common elements of the Neotropical Flora and is known for its specialized pollination system involving the production of floral oils (i.e., lipids) which are harvested by solitary, oil-collecting bees and used for larval provisioning or nest construction. Comprising 1,300 species, the Malpighiaceae has its center of diversity in South America, where the family is thought to have originated 65 Mya. Among South American biomes, the Brazilian Cerrado, a seasonally dry savanna-like vegetation considered one of the World's Biodiversity Hotspots for angiosperms, is especially rich in Malpighiaceae species. But how do these specialists coexist and persist successfully in symmetry if competition for pollination services and risk for hybridization (i.e., via interspecific pollen transfer) among close relatives is expected to be high? Here, we explore the niche ecology of 33 sympatric Malpighiaceae species, focusing on potential temporal, spatial, behavioral, and morphological partitioning mechanisms. Among other factors, we show the importance of temporal niche partitioning in phylogeny promoting the Malpighiaceae species richness that characterizes the Cerrado vegetation throughout its geographic range. We discuss the relevance of the emergent patterns observed in this system to improve our understanding of niche and community ecology in other Neotropical plant communities where specialized pollination is also widespread.

## S.10.4 Chemical divergence in floral scents of two Magnolia species from the Colombian Andean region

Juan Pablo Santa-Ceballos¹, Marcela Serna Gonzalez², Vinita Gowda³, Hiroshi Suzuki-Azuma⁴

1 Departamento de Ciencias Forestales, Universidad Nacional de Colombia, Sede Medellín, Medellín, Colombia. 2 Facultad de Ingeniería, Tecnológico de Antioquia-Institución Universitaria, Medellín, Colombia. 3 Department of Biological Sciences, Indian Institute of Science Education and Research Bhopal, Bhopal, Madhya Pradesh, India. 4 Faculty of Engineering, Toyama Prefectural University, Toyama, Japan

Floral scents serve as chemical signals to attract pollinators and thus play an essential role in the reproductive strategies of angiosperms. In Magnoliaceae, floral scents were identified as primary attractants for floral visitors. This study explores the chemical composition of floral scents in two Colombian endemics species, *Magnolia jardinensis* and *M. yarumalensis*, from the Andean cloud forests. Through scent collection by dynamic headspace methods, followed by Gas Chromatography-Mass Spectrometry (GC-MS) analysis, we identified distinct volatile compounds from flowers in their staminate and pistillate stages, in both the species. *Magnolia jardinensis* exhibited seven compounds, while *M. yarumalensis* displayed a more complex profile with sixteen compounds. Notably, five
<figure><img src="image_42.png" /></figure>

compounds were shared between the male and fe-
male floral stages of *M. yarumalensis*, reinforcing the
notion of specialized pollination systems. The chemi-
cal divergence between these two *Magnolia* species
aligns with previously observed differences in floral
visitors and phenological patterns. *Magnolia yarumal-
ensis* attracts beetles of the Staphylinidae family, while
*M. jardinensis* attracts beetles of the *Cyclocephala*
genus (Family Scarabaeidae). This work holds signif-
icance for conservation efforts. The specialized polli-
nation systems observed in these *Magnolia* species
highlight the vulnerability of these ecosystems to dis-
ruptions, underscoring the need for targeted conser-
vation measures to safeguard the intricate relation-
ships between *Magnolia* trees and their pollinators.

S.10.5 Sapromimesis as a
pollination strategy – floral
mimicry of the necrobiome

Katherine R. Goodrich¹, Jade Savage²

1 Widener University, 1 University Pl. Chester, PA, USA.

2 Bishop's University, Sherbrooke, Quebec, Canada

Some flowers use false advertisements to decep-
tively attract pollinators. Mimetic resemblance to
other organisms or their by-products can result in
pollinators cognitively misclassifying the flowers,
as evident by behavior such as mating attempts
or egg-laying. Floral mimicry of decomposing sub-
strates, frequently termed "oviposition-site mimicry,"
presents a particularly challenging study system.
Current studies on flowers mimicking decomposing
substrates (e.g. carrion, fermentation, dung) show
intriguing patterns of convergence of floral scents,
color, and texture. However, we do not yet fully un-
derstand the cues utilized by foraging and/or gravid
saprophagous insects which pollinate sapromimet-
ic flowers. Examining floral phenotype convergence
in sapromimetic species may help elucidate cru-
cial cues used by these insects. Traditional views of
mimicry suggest that increased phenotypic overlap
of the mimic with a specific model should increase
mimic fitness. But in sapromimesis, it is frequent-
ly unclear whether the flower imitates a general
set of cues used by saprophagous insects or more
specialized imitation of a particular model, such
as a species of fermenting fruit. I present ongoing
research for two sapromimetic species – *Asimi-
na triloba* (Annonaceae) and *Syplocarpus foeti-
dus* (Araceae) – occurring in the same habitats of
northeastern USA. Yeasty-smelling maroon flowers

of *A. triloba* overlap chemically with odors of a range
of fermenting environmental substrates and attract
a range of fermentation-seeking Diptera. Fetid, car-
rion-smelling inflorescences of *S. foetidus* chemi-
cally overlap with both carrion and fermenting sub-
strates, potentially attracting a broader range of
saprophagous insects. We highlight convergent flo-
ral characteristics between these species as well as
potentially important differences. We present sub-
strate chemical data for locally-occurring potential
models. Finally, we identify insect taxa attracted to
flowers of both species, and to carrion and fermen-
tation baits placed near blooming plants. This inves-
tigation helps elucidate boundaries and potential
overlap between substrate properties and attracted
insect communities within a close geographic area.

S.10.6 Future distribution
of Iranian Salvia and its
pollinators using ecological
niche modeling and
molecular data

Fatemeh Moein¹, Hojatolah Saeidi¹

1 Department of Plant and Animal Biology, Faculty of Biological Science and Technology, University of Isfahan, Isfahan, Iran

Climate change and anthropogenic drivers are widely considered as the most important threat to biodiversity. Pollinators play an important role in ecosystem regulation. Phenological mismatches between plant-pollinator interactions can threat at population or species level depend on the degree of specialization. Taken into account these adverse effect and limited time for conservation, providing maps of species distribution are required to identify where vulnerable species are at risk of decline or even extinction specially for plants depend on pollinators for reproduction. *Salvia*, the largest genus in Lamiaceae, includes approximately 1000 species in the world with around 200 species in southwestern Asia distribution. More than 80 percent of the species are characterized by special pollination system known as lever mechanism. *Salvia* species in southwest of Asia are melittophilous, pollinated by insects, mostly bees. (Apis mellifera, Bombus sp and Xylocopa sp.). Iran with 72 species of *Salvia* in which 19 of them are endemic is one of the main center of Salvia diversity in southwestern Asia, we focused on Iranain *Salvia* and relevant pollinators
<figure><img src="image_43.png" /></figure>

## S.11.2 Phenotypic diversity generated by transgenerational plasticity and transposable elements enhances population functioning

Javier Puy¹, Vit Latzel², Carlos P. Carmona³, Michael Thieme⁴, Etienne Bucher⁵, Lars Götzenberger²,⁶, Francesco de Bello⁶,⁷

1 Estación Biológica de Doñana, CSIC, Sevilla, Spain. 2 Institute of Botany, Czech Academy of Sciences, Průhonice, Czech Republic. 3 Institute of Ecology and Earth Sciences, University of Tartu, Tartu, Estonia. 4 Department of Plant and Microbial Biology, University of Zürich, Zürich, Switzerland. 5 Crop Genome Dynamics Group, Nyon, Switzerland. 6 Department of Botany, University of South Bohemia, České Budějovice, Czech Republic. 7 CIDE-UV-CSIC, Centro de Investigaciones sobre Desertificación, Valencia, Spain.

The observed positive effect of diversity on ecosystem functioning, explained by mechanisms of trait diversity, has rarely been assessed within populations. However intraspecific phenotypic variability should theoretically cause similar biodiversity effects. Intraspecific trait variation could stem both from underlying genetic diversity and non-genetic mechanisms such as plasticity in response to environmental cues. This plasticity can potentially last multiple generations and can derive, for example, from epigenetic modifications or mobilization of transposable elements. Using *Arabidopsis thaliana* as a proof-of-concept model, our work has been on unravelling the effect of phenotypic diversity generated by 1) heritable plasticity and 2) transposable elements on coexistence, population productivity, and resistance to stress, as well as on comparing their magnitude with the effect of genetic diversity. Across two glasshouse experiments, three different types of *Arabidopsis thaliana* populations were established: one type with differing levels of genetic diversity, another one with a different mixture of individuals that differed in their *ONSEN* retrotransposition insertions, and a last one, genetically uniform but with varying diversity levels of the parental environments (parents grown in the same or different environments). In all populations, we tested whether the increasing diversity increased the phenotypic diversity of populations and enhanced their functioning under different conditions. As expected, both

heritable plasticity and transposable elements created differentiation in ecologically important traits connected to different axes of the plant 'economics' spectrum. Such increased phenotypic diversity ameliorated the negative effect of competition between coexisting individuals and enhanced population productivity and stress resistance; and interestingly, by a similar magnitude as genetic diversity did. Our results highlight the ecological relevance of unexplored sources of phenotypic variability because as other sources of biodiversity, they could similarly affect ecosystem and population functioning.

## S.11.3 Intraspecific trait variability within and across generations: implications for coexistence and stability

Francesco de Bello¹, Vit Latzel², María del Mar Sobral Bernal³, Javier Puy⁴, Christina Richards⁵

1 Centro de Investigaciones sobre Desertificación (CSIC), Valencia, Spain. 2 Academy of Sciences of the Czech Republic, Institute of Botany, Pruhonice, Czech Republic. 3 University of Santiago de Compostela, Santiago de Compostela, Spain. 4 Estación Biológica de Doñana, CSIC, Spain. 5 University of South Florida, USA & University of Tübingen, Germany

To comprehend the responses of ecosystems to global change, unraveling the dynamics of biological diversity and ecosystem function over time is imperative. A key pursuit in this context is deciphering the factors fostering coexistence and resilience to environmental shifts within communities and populations. Ecologists have debated various mechanisms modulating local biodiversity effects on ecosystem properties although effects within populations remain largely unexplored. Ecological mechanisms act initially on individuals, scaling up to communities and ecosystems, shaping and being shaped by Intraspecific Trait Variability (ITV). These variations stem from diverse sources: genetic (attributed to DNA differences) or non-genetic (e.g., epigenetic mechanisms without DNA sequence modifications). Genetic diversity is believed to empower populations to adapt to changing environmental conditions by providing a broader array of phenotypes from which the fittest can be selected. Concurrently, the ability of an organism to adjust trait expression in response to the environment (i.e., phenotypic plasticity) can be also heritable. Plasticity, in fact, does not allow
only short-term responses within an individual's life-
time (within-generation plasticity) but also cause
inherited differences across generations (transgen-
erational plasticity). The relative impact of genetic
versus non-genetic heritable mechanisms, encom-
passing their temporal extents and responsiveness
to different environmental drivers, remains largely
unknown. Questions persist regarding the cumu-
lative nature of non-genetic effects in stable envi-
ronmental conditions across generations and the
reversibility of induced differences when environ-
mental conditions shift. Addressing these issues ne-
cessitates researchers to develop new approaches
to evaluate how genetic diversity and non-genetic
inheritance diversity intersect with population sta-
bility over time. Additionally, outlining experimental
approaches to tackle these questions is crucial. This
contribution seeks to synthesize ideas and perspec-
tives aimed at understanding the interplay between
genetic and non-genetic effects on short- and
long-term phenotypic variation within species. It ex-
plores how these effects influence coexistence and
ecological stability of growing conditions and traits.

S.11.5 Epigenetics and
transgenerational plasticity
in the face of climate
change: a matter of
reproductive mode?

Iris Sammarco¹, Zuzana Münzbergová ¹,²,
Vít Latzel¹

1 The Czech Academy of Sciences, Průhonice, Czech Republic. 2 Faculty of Science, Charles University, Prague, Czech Republic

Climate change poses a significant threat to plant
species, potentially pushing them beyond their adap-
tive capacities. Epigenetic modifications, such as DNA
methylation, have emerged as a key mechanism
enabling plants to quickly adapt to environmental
changes by generating locally adapted phenotypes.
These phenotypic changes can be even inherited
across multiple generations, a phenomenon known
as transgenerational plasticity. Specifically, the ex-
tent and ecological implications of this heritability are
known to potentially differ between sexual and clonal
reproductive modes. However, the impact of environ-
mentally induced epigenetic changes on transgen-
erational plasticity across these reproductive modes
remains poorly understood. Here, we assessed the

adaptive potential of woodland strawberry (*Fragaria
vesca*), a widely distributed herb with both clonal and
sexual reproductive capabilities, in the face of pro-
jected environmental conditions expected by the end
of the 21st century. These conditions included a tem-
perature rise of 4 °C, an atmospheric CO₂ concentra-
tion twice that of the present (800 ppm), and periodic
drought periods. We quantified ecologically relevant
phenotypic traits and examined whole-genome DNA
methylation patterns in both parental individuals and
their clonal and sexual offspring. We found evidence
for transgenerational plasticity induced by the pa-
rental environment, with a stronger overall effect ob-
served in the clonal offspring compared to the sexual
offspring. These results suggest that transgenerational
plasticity via DNA methylation could potentially facili-
tate plant adaptation to future environmental condi-
tions, particularly in clonal species.

S.11.6 Epigenetic memory of
climate change associated
stress in forest trees

Joe Q He¹, Rosa Sánchez-Lucas¹, Phoebe
Swift¹, Charlotte Jones¹, Adriane Esquivel-
Muelbert¹, Marco Catoni¹, Estrella Luna-Díez¹

1 University of Birmingham, Birmingham, UK.

Memory is the acquisition, retention and transmission
of information guiding future action. The imprinting of
memory in plants mostly happens by altering its epi-
genetic signature through DNA methylation, which has
been shown to contribute to both short-term pheno-
typic plasticity and the longer-term adaptive capacity.
In our large research project MEMBRA, we are studying
epigenetic changes and transgenerational memo-
ry because of climate change-associated stress in
key forest tree species present. Different abiotic (i.e.,
drought, frost, elevated CO2) and biotic stresses (i.e.,
insect infestation and different diseases) have been
selected based on our data showing that they alter im-
portant traits, such as growth, within UK forests. Leaf DNA
from selected trees exposed to stress has been extract-
ed and subjected to Whole Genome Bisulfite Sequenc-
ing (WGBS). A custom bioinformatic pipeline was es-
tablished to call differentially methylated points (DMPs)
and regions (DMRs) and to associate the differential
stresses to the methylome. Our long-term aim is to in-
tegrate the capacity of tree species to develop memo-
ry into the spectrum of tree form and function and test
its influence on forest resilience. Memory will be used as
a plant functional trait to help select the most resilient
<figure><img src="image_44.png" /></figure>

forests and populations to be targeted as a source of seeds for afforestation programme. Therefore, MEMBRA is concerned with the implications of establishing long-term memory of stress as a key characteristic of trees that can fundamentally change climate resili-

ience strategies. Through this multidisciplinary project working with artists, ethicists and policy experts, we are providing a paradigm shift in our understanding of the adaptation capacity of forests to the unprecedented rapid global-change-driven environmental shifts.

# S.12 METAL HYPERACCUMULATORS: NEW ADVANCES ON A BOTANICAL CURIOSITY.
## SESSION 1

### S.12.1 Trace element hyper accumulator plant traits: a call for trait data collection

Béatrice Gervais-Bergeron¹, Adrian Paul¹, Pierre-Luc Chagnon², Alan Baker³,⁴,⁵, Antony van der Ent³,⁵, Michel-Pierre Faucon⁶, Celestino Quintela-Sabarís⁷ and Michel Labrecque¹

1 Institut de recherche en biologie végétale, Université de Montréal, Canada. 2 Agriculture and Agrifood Canada, Canada. 3 Sustainable Minerals Institute, The University of Queensland, Australia. 4 School of BioSciences, The University of Melbourne, Australia. 5 Laboratoire Sols et Environnement, Université de Lorraine, France. 6 UniLaSalle, AGHYLE, France. 7 Grupo de Ecoloxía Animal, Universidade de Vigo, Spain

Hyperaccumulator plants exhibit extreme ecophysiological characteristics, which make them valuable for conservation and suited for phytoremediation. Understanding their ecological strategies might help identify the species and functions to be fostered in phytoremediation, restoration, and conservation projects for metalliferous sites. Here, we identified the hyperaccumulator species in the worldwide plant trait database TRY and cross-referenced these trait syndromes associated with the ability of plants to concentrate metals. We collected three traits (leaf area (LA), specific leaf area (SLA), and leaf dry matter content (LDMC)) on all available species on TRY (n= 4523), from which 20 were identified on the global hyperaccumulator database. This allows us to link trace element hyperaccumulation with broader plant ecological strategies, such as competitive, stress-tolerant, and ruderal strategies (CSR) or resource acquisition ones. With these primary data,

hyperaccumulators seem to have smaller leaves and poor competitive ability compared to non-hyper accumulator species. Contrary to expectations, we found no indication of hyperaccumulators being more resource-conservative on the leaf economics spectrum. However, these data remain fragmentary as only 2.7% of hyperaccumulators have their traits published in the TRY database. We thus proceed to a systematic review to retrieve around 40 additional new hyperaccumulator species present on TRY and call for an international collaborative sampling effort to measure traits in more hyperaccumulator species. The recent development of trait-based models to construct plant communities providing optimal ecosystem services (e.g., phytoremediation, restoration)³ requires further research to identify predictable trait-service relationships. This research provides a first synthesis of the research advancement on hyperaccumulator traits, as well as a first synthesis analysis of their strategies at a global scale.

### S.12.2 Functional syndromes of hyper-accumulating species: recent advances and perspectives

Florian Delerue¹

1 Université de Bordeaux, CNRS, Bordeaux INP, EPOC, UMR 5805, Pessac, France

Data bases of plant traits are now available at the global scale, and functional axes of variation among plants are well defined. Here, we present the synthesis of two recent studies revealing the position of herbaceous metallophyte species along these functional axes, with a special focus on hyper-accumulators. These studies covered the major kinds of metalliferous ecosystems either spontaneous or anthropogenic and in diverse
<figure><img src="image_45.png" /></figure>

biogeographical regions: Cu-rich environments including the "copper hills" in central Africa; Zn-Pb-Cd rich calamine substrates and Ni-Co-Cr rich serpentinic substrates in Western Europe. Shoot and root functional traits have been characterized following standardized protocols for 348 species present in 30 metalliferous communities, as well as leaf metal concentration and soil biogeochemical properties. In all cases metallophyte species exhibited small size like many species growing on other harsh substrates, some being among the shortest plants worldwide. This confirms a constant impact of metal-stress on plant stature. Additionally, an important variability regarding the leaf resource economics axis related to soil resource acquisition was found. Metallophyte communities are composed by a mix of conservative species (slow resource acquisition, long-lasting leaves) together with acquisitive species (fast resource acquisition and growth, short-lasting leaves). Remarkably, hyper-accumulating species were found among the most acquisitive species in the metalliferous communities studied. These results offer new perspectives regarding the evolution of metal-accumulation in plants. Fast-growing species maintain high soil resource acquisition on harmful substrates, leading to an important inflow of metals into their leaves. This may represent a first step towards hyper-accumulation if specific physiological adaptations are selected afterwards. Future studies should test the genericity of these results, by focusing on woody species present in some serpentinic regions. Additionally, the importance of the collaboration with mutualistic fungi (another functional axis related to both soil resources acquisition and plant-metal relationships) for hyper-accumulating species should be specified.

### S.12.3 Addressing the diversity of the mechanisms involved in metal hyperaccumulation in plants

Sylvain Merlot¹,², Celestine Belloeil¹, Melina Gallopin¹, Vanesa García de la Torre¹, Dubiel Alfonso Gonzalez³, Yohan Pillon⁴

1 Institute for Integrative Biology of the Cell, CEA, CNRS, Paris-Saclay University, France. 2 Laboratoire de Recherche en Sciences Végétales, Toulouse University Paul Sabatier, CNRS, INP Toulouse, France. 3 Jardín Botánico Nacional, Universidad de La Habana, Cuba. 4 Laboratory of Tropical and Mediter-

ranean Symbiosis, IRD, CIRAD, INRA, Montpellier University, France

The extraordinary ability to accumulate and tolerate tremendous amounts of metals such as nickel, zinc, or manganese in their leaves has been observed in about 700 plant species belonging to more than 50 families. For many years, these metal hyperaccumulator species have attracted the attention of scientists interested in understanding the evolution of this complex and peculiar trait. In the context of the decarbonization of energy, metal hyperaccumulators appear as an opportunity to develop phytotechnologies aimed at limiting the metal pollution associated with the growing demand for metals. However, our knowledge of the molecular mechanisms involved in metal hyperaccumulation, which is key to the development of these technologies, is still very limited. To extend our knowledge of the diversity of hyperac cumulator species, we participated in a global effort to identify new hyperaccumulator species using the X-ray fluorescence technology focusing on the flo ra of the neotropical region. Our herbarium and field studies revealed the first example of a zinc hyperac cumulator from the Amazonas region, as well as new nickel hyperaccumulators originating from Cuba. The identification of nickel hyperaccumulators and closely related non-hyperaccumulator species from different families has opened the possibility to perform compar ative cross-species transcriptomic studies to identify orthologous genes whose expression is linked to nick el hyperaccumulation. Among candidate genes, our analyses revealed that the high expression in leaves of metal transporters of the IREG/ferroportin family is a highly conserved mechanism involved in nickel hy peraccumulation across distant plant species. We are now developing the multi-metal hyperac cumulator *Noccaea caerulescens* of the Brassicaceae family as a genetic model to validate and study the role of can didate genes in metal hyperaccumulation.

### S.12.4 Deciphering the physiological and biochemical mechanisms of thallium hyperaccumulation in Biscutella laevigata

Mirko Salinitro¹, Sandrine Isnard², Dennis Brueckner³, Kathryn M. Spiers³, Mark G. M. Aarts⁴, Amelia Corzo Remigio⁵, Antony van der Ent⁴