Dr Florian Jabbour
email: fjabbour@mnhn.fr

Msc Student

Tatiana Giraud

Blandine Doligez

Frédéric Jiguet

J Fuchs

Erwan Cherel

Julian Lazard

MNHN. Antoine Larré M1

PhD Julien Massoni

Bader Kezzim

Boris Domenech

Investigation of floral architecture, development and genetic bases 2006

Keywords: c. Expert assessments VIII.d. Science popularization morphology, development, molecular phylogeny, and biogeography Phylogeny, molecular dating

Curriculum Vitae I. My scientific trajectory (or how I became a lifelong student of plant and flower form evolution) II. Morphology: The renaissance of an old discipline III. Investigating the diversity and evolution of plant and floral forms: One for all and all for one! IV. Supervision of students and of research projects V. Evolution of flower form in a phylogenetic context: Selection of past and current research projects V.a. Flower symmetry: Development, evo-devo, and evolution V.b. Systematics and flower evolution in Delphinieae (Ranunculaceae) V.c. Flower dimorphism in Nigella damascena L. (Ranunculaceae) V.d. Systematics of Stephania (Menispermaceae) V.e. Flower disparity in Ericales VI. Future and prospective research VI.a. Amending the description of the genus Delphinium (Ranunculaceae) VI.b. Flower development in the genus Staphisagria (Ranunculaceae) VI.c. Identifying the sister group of the Delphinieae (Ranunculaceae) VI.d. A synthesis about the research done so far on Delphinieae (Ranunculaceae) VI.e. Evo-devo of perianth dimorphism in Nigella damascena (Ranunculaceae) VI.f. Comparative study of petal development in Ranunculaceae VI.g. Evolution of extra-and intrafloral nectaries in Angiosperms VI.h. Phylogeography of Delphinium in the East African Rift System VI.i. Long-term projects VII. The many futures of plant morphology at the MNHN (and beyond) VIII. Involvement in the different tasks of the MNHN VIII.a. Research team responsibility VIII.b. Curation at P Herbarium VIII.c. Teaching VIII.

FOREWORD

As a child and teenager, I used to collect almost everything: gemstones and minerals, fossils, shells, feathers, dried plants, wine labels. Each of these collections was classified according to simplistic (and very personal!) systems. I remember I regularly reorganized my collections to optimize space and to be able to find back a particular object the fastest possible. It was a game, and with hindsight, an unconscious attempt to bring "order out of chaos". I sometimes think that me working in a Museum of Natural History is no coincidence, when looking at the influence two men had on me. On the one hand, my father taught me how interesting it can be to establish and take care of a collection. On the other hand, my maternal grandfather gave me the opportunity to accompany him during his walks in parks and forests. He could identify all kind of birds and plants, and was a true lover of natural history (see Fig. 1). He had to quit school relatively early, but as a self-taught person, and thanks to his endless curiosity and enthusiasm, he was one of the most cultivated people I have ever known. I dedicate this humble thesis to him.

STUDENT OF PLANT AND FLOWER FORM EVOLUTION)

Morphology, or the study of organismal forms, is at the crossroads of genomes, developmental and historical (phylogenetic) constraints, and environment. By definition, it is an integrative discipline building bridges between, for instance, evodevo and ecology, systematics and physiology.

I consider myself as a student of plant and flower form evolution, and I organized this thesis around this scientific topic. In this manuscript, I made the choice to use the term morphology rather than phenotype for two reasons: 1) these terms are not synonymous. A phenotype is the set of observable features of an organism, such as its morphology, but also, for instance, its metabolism and behaviour; 2) I think it is important to highlight the very long history of morphology as a discipline.

In this thesis, I will first say few words about the history of plant and flower morphology, and highlight its integrative nature. Second, I will give some ideas about the supervision of students, before developing a selection of past, current, and prospective research projects. Third, I will explain my involvement in the other tasks of the MNHN.

In the following paragraph, I will shortly explain how I became a researcher dedicated to the study of plant and flower morphology. In my opinion, it got funded by the FWF (Austrian Science Fund) mainly because the biological question was relevant (see Chartier et al. 2014[START_REF] Chartier | How (much) do flowers vary? Unbalanced disparity among flower functional modules and a mosaic pattern of morphospace occupation in the order Ericales[END_REF], but also because the project uniquely integrated botany and the mathematics of a morphospace approach. Before even starting this second postdoc (which was brilliantly taken in charge of by Marion Chartier), I got hired at the MNHN for an Assistant Professorship focusing on plant radiations on islands (I had studied the radiation of Delphinieae Warm. on the Himalayas and the mountains of SW China, [Jabbour and Renner 2012b]). While studying radiations on islands (Soulebeau et al. 2015), I pursued my studies on flower morphology, getting trained in plant anatomy, and supervising a PhD student on a flower-morphology-based topic. In January 2016, some colleagues and I got together in a team entitled Morphology, Anatomy and Development of Plants, in order to gather our expertise and know-how.

II. MORPHOLOGY: THE RENAISSANCE OF AN OLD DISCIPLINE

Since [START_REF] Goethe | Versuch die Metamorphose der Pflanzen zu Erklären[END_REF], morphology as a discipline has been not only descriptive but explanatory too. Johann Wolfgang von Goethe proposed that the different parts of a flower result from the "metamorphosis" (meaning transformation) of a basic organ, the "ideal leaf". Goethe's hypothesis was validated by the students of the ABC model, in which combinatorial interactions between different classes of floral homeotic genes (A, B, and C, among others) determine the identity of the floral organs [START_REF] Theissen | Plant biology: floral quartets[END_REF].

Morphology and its central concept, homology, are pre-Darwinian in origin, and as a result obsolete and arguable in the eyes of many [START_REF] Richter | A research program for evolutionary morphology[END_REF].

The pre-Darwinian idealistic homology concept was not anchored in organismal evolutionary history. According to this concept, two characters were thought to be homologous because they were built based on the same plan or archetype. This concept, already used by Goethe, is older than the term homology, which was defined in a modern way by Owen [START_REF] Mayr | The growth of biological thought: Diversity, evolution, and inheritance[END_REF]. Two structures are now called homologous if they represent corresponding parts of organisms, which are built according to the same body plan. The existence of corresponding structures in different species is explained by derivation from a common ancestor that had the same structure as the two species compared [START_REF] Wagner | The biological homology concept[END_REF].

Descriptions, comparisons and explanations of forms refer primarily to single organisms, morphemes (parts of an organism; units of description without reference to evolution; see [START_REF] Richter | A research program for evolutionary morphology[END_REF] or semaphoronts (characters expressed at any stage of ontogeny, [START_REF] Sharma | Hennig's semaphoront concept and the use of ontogenetic stages in phylogenetic reconstruction[END_REF].

Modern morphology is the heir of a long tradition. Many famous morphologists deeply influenced the evolution of concepts and techniques in their field. Old theories are still currently discussed and contribute to the maturity of the discipline. For instance, Haeckel's theory of recapitulation proposes that parts of the evolutionary history of a lineage are conserved in the ontogeny of its constituting extant species (Haeckel 1866). Although this concept received some criticism later on (e.g. [START_REF] Gould | Ontogeny and phylogeny[END_REF], its essence is still the basic principle of evo-devo [START_REF] Niklas | From Goethe's plant archetype via Haeckel's biogenetic law to plant evo-devo 2016[END_REF], namely that it is generally possible to reconstruct organismal evolution and potentially phylogenetic relationships by comparing developmental processes (see semaphoronts)

of different organisms [START_REF] Wanninger | Morphology is dead -long live morphology! Integrating MorphoEvoDevo into molecular EvoDevo and phylogenomics[END_REF].

However, being an old discipline may have its drawbacks. We may think everything was described, we might hear that morphology cannot teach us anything anymore in our days of omics-driven science, students often run away from plant morphology because the word is spread that it is a classical and obsolete discipline, that one has to read many old publications before initiating any study, and most of all, because "no one gets any position doing (only) morphology".

In the context of a steady decline in interest in morphology both at research institutions and funding agencies [START_REF] Schönenberger | Modern theoretical and technical approaches in plant morphology[END_REF], where morphology is mostly perceived as a provider of characters for systematic circumscription [START_REF] Kaplan | The science of plant morphology: definition, history, and role in modern biology[END_REF], there has been a trend in recent years to justify the necessity of research in this field and to provide evidence that the discipline has renewed itself (while on the one hand, this behaviour of calling into question is healthy, on the other hand it is a sign that morphologists feel insecure in the current scientific world). Some publication titles or paragraph titles are explicit in this respect: "Why is it important to maintain a research program in morphology?" [START_REF] Giribet | Morphology should not be forgotten in the era of genomics -a phylogenetic perspective[END_REF], "Morphology is dead -long live morphology!" [START_REF] Wanninger | Morphology is dead -long live morphology! Integrating MorphoEvoDevo into molecular EvoDevo and phylogenomics[END_REF], and linguistic innovations were proposed to modernize the field of morphology (and probably to compete with the disciplines that are usually the most funded), such as "high-throughput morphology", "next-generation histology", "phenomics" [START_REF] Giribet | Morphology should not be forgotten in the era of genomics -a phylogenetic perspective[END_REF], "MorphoEvoDevo" (for evolutionary developmental morphology; parallel with EcoEvoDevo [START_REF] Gilbert | Ecological developmental biology: the environmental regulation of development, health, and evolution[END_REF];

Wanninger 2015).

After a long period of absence from mainstream evolutionary biology, morphology experienced a renaissance within the field of evo-devo [START_REF] Richter | A research program for evolutionary morphology[END_REF]Wirkner 2014, Specht and[START_REF] Specht | Flower evolution: the origin and subsequent diversification of the angiosperm flower[END_REF]. Model-based reconstruction of ancestral states (e.g. [START_REF] Sauquet | The ancestral flower of angiosperms and its early diversification[END_REF], taking into account the phylogenetic framework and recent paleobotanical discoveries, and morphospace analyses (Chartier et al. 2014[START_REF] Chartier | How (much) do flowers vary? Unbalanced disparity among flower functional modules and a mosaic pattern of morphospace occupation in the order Ericales[END_REF] have also breathed new life into the field of morphology. A striking feature of morphological evolution is that due to developmental, functional, and phylogenetic constraints, not all theoretically possible architectures have been explored during evolution.

Morphospaces are theoretical spaces describing and relating organismal phenotypes.

Although widely applied in zoology, morphospace analyses have so far been largely disregarded in botany (but see Mander 2016), with notable exceptions in the field of pollination biology [START_REF] Ollerton | A global test of the pollination syndrome hypothesis[END_REF], Schönenberger et al. 2016).

We now are in need of a new generation of morphologists and anatomists (ca. 2,000 plant species described each year [START_REF] Christenhusz | The number of known plants species in the world and its annual increase[END_REF]). These scientists have to be able to understand genetics, genomics, molecular phylogenetics, and evodevo, in addition to mastering the old literature (often in English, but also in German and French). More than understanding the results, they should be able to communicate with the researchers in those disciplines, and design research projects hand in hand with them. Hardly any morphology-based project will get funded nowadays (at a national or international level) and morphology has to be combined with other approaches to become appealing.

The following boxes present the biographies of a selection of botanists who were influential in the fields of plant (and flower) morphology and anatomy. Three of them had strong links with the MNHN. A last box deals with the evolution of imaging techniques, most of which revolutionized the study of plant morphology. The plates illustrating the successive developmental stages of the meristems or of selected organs are of high quality, and are still useful nowadays.

Most of the biological material he studied was provided by the MNHN. For this reason, he followed the classification of Angiosperms that the MNHN was using back then.

Payer acknowledges in le Traité the influence of Charles-François Brisseau de Mirbel, who was, according to him, the father of plant organogenesis. de Mirbel started his career at the MNHN before settling at the Sorbonne. He is also known for his deep impact on the fields of cytology and physiology that were emerging at that time.

The Friederich Wilhelm Benedikt Hofmeister is known as the discoverer of the alternation of generations (sporophyte-gametophyte) and for his fundamental studies of plant embryology.

Hofmeister was an autodidact in botany; he had no university education, much less a Ph.D. degree, but ultimately was appointed Professor of Botany at Heidelberg at age 39.

Hofmeister investigated the causal bases for the morphological diversity of seed plants. To answer this question, he introduced in his research concepts and tools from physics and chemistry, all of which he also taught himself.

As one of the earliest plant biophysicists, Hofmeister was so far ahead of his time that he was not understood by his contemporaries or many who followed him.

Hofmeister's principal contribution to the field of plant morphology was his book Allgemeine Morphologie der Gewächse (General morphology of growth;

Hofmeister, 1868). In this complex, hardly accessible book, he presented an analytical view of the developmental basis underlying the diversity of plant form, but also biophysical interpretations of a range of phenomena including phyllotaxis and the effects of gravity and light on plant morphology.

Hofmeister's contribution to biology is still far from widely acknowledged.

However, [START_REF] Kaplan | The genius of Wilhelm Hofmeister: the origin of causal-analytical research in plant development[END_REF] conclude that "his reputation became eclipsed because he was so far ahead of his contemporaries that no one could understand or appreciate his work". The study of Hofmeister's work is also limited because it is published in German, though translations for some articles have been made. was the first one to systematically organize this field of botany. Through his publications and teaching, he defended an approach of plant anatomy integrated into the more general discipline of evolutionary biology. His work is mainly based on highlighting the geometrical properties of the observed structures, as it was a usual practice in science in the middle of the 19 th century (Bachelard 1938).

All the slides prepared by Van Tieghem and his students are kept at the MNHN.

This collection consists of ca. 20,000 slides, which corresponds to one third of the slide library of the MNHN. Many samples come from rare or important specimens (type specimens and herbarium specimens collected in the 18 th and 19 th centuries).

Almost all the samples are in good state and are cited in Van Tieghem's publications (available at the libraries of the MNHN). The project of digitizing this collection has still to be funded. Although this collection is of an immense scientific importance, it is so far rarely used.

BOX 4:

Biography and works of Michel Guédès (1942Guédès ( -1985, †43) , †43) Illustration: picture of a book cover taken at the Library of Botany at the MNHN.

Portrait from [START_REF] Aymonin | Michel Guédès (1942-1985)[END_REF].

Michel Guédès was a self-trained morphologist. He joined the staff of Paris Herbarium in 1971. He adopted a generalist approach of botany, investigating many taxonomic groups within Angiosperms, and being keen on morphology, plant architecture, systematics, teratology, floristics, nomenclature, history of science (e.g. [START_REF] Guédès | Goethe et Geoffroy Saint-Hilaire[END_REF]). He had an excellent knowledge of the German botanical literature and his work on floral structures reflected the theory of Goethe.

Although he passed away quite young, he published >170 publications, often in foreign journals such as Taxon, Flora, Botanische Jahrbücher, and Phyton. His book "Morphology of seed plants" (1979), with about 900 references, stimulated a renewed interest in the subject. He also published popular books for the general public (Les bienfaits des plantes; Les plantes sauvages; Baies et fruits de nos bois et jardins).

According to the biographical information available at the library of the MNHN, Michel Guédès had a complicated and thwarted career history. He did not seem to be a man of compromise, and was convinced of his hierarchy's disregard. In his opinion, he did not get any promotion at the MNHN because his directors believed he could not travel far and long, and because of his lack of servitude.

People who knew him add that he mastered the botanical literature (in French and other languages), and that he always found time to discuss students' or colleagues' projects, address their conceptual or technical problems, and give them advice.

Reference

 Bibliographic and publication records, Bibliothèque de Botanique (MNHN)

 Aymonin GG (1987) Michel Guédès (1942Guédès ( -1985)). Taxon 36(3): 699-700. Imaging technique (approximate date of first use)

1. Principle 2. Objective

Selected example

Light or optical microscopy (17 th century)

1. Uses visible light and a set of lenses to magnify the image of a sample.

2. Simple 2D observation at a magnification reaching 1,500x (product of the powers of the ocular and the objective lens).

III. INVESTIGATING THE DIVERSITY AND EVOLUTION OF PLANT AND FLORAL FORMS: ONE FOR ALL AND ALL FOR ONE!

Although studying plant (and floral) morphology and development requires investing a lot of time to make observations at the binocular or at the SEM, endlessly dissecting floral buds and reading a fairly abundant literature, morphology is everything but a solitary discipline. Interacting with students, interns, and collaborators indeed makes the daily work more rewarding and interesting, and incomparably more fruitful.

I would suggest to the students of floral morphology to design the most integrative studies possible. This may sound like a simplistic statement but it is not easy to achieve. To carry out an excellent morphological study, one would need to have knowledge -and be able to design and conduct research -in various fields, such as anatomy, paleobotany, geometric morphometrics, phylogenetics, and evo-devo (Fig. 2, [START_REF] Debat | Analysing phenotypic variation: when old-fashioned means up-to-date[END_REF]. Indeed, the morphology of an organism is at the interface of genome and environment, and its thorough analysis requires investigating its molecular bases, the developmental processes underlying it, and its functions in a given environment. Adapted from [START_REF] Kaplan | The science of plant morphology: definition, history, and role in modern biology[END_REF].

This list may seem restricted to methods, but each of the fields cited over is linked to a set of concepts and schools of thought, and is hence much more than a method.

Students of plant (and floral) morphology have to deal with ancient and old literature in depth (e.g. As it is almost impossible to master all these disciplines, especially at an early career stage, collaborations are crucial to achieve a thorough research project focusing on plant (and floral) morphology, in order to gather huge amount of data [START_REF] Sauquet | The ancestral flower of angiosperms and its early diversification[END_REF], Fig. 3), analyze morphological evolution using innovative methods (Chartier et al. 2017), or interpret morphological evolution in the light of other disciplines [START_REF] Mouly | The emblematic "Captaincookia" in New Caledonia (Rubiaceae), a divergent Ixora species showing adaptive variability from humid to sclerophyll forests[END_REF]. For instance, in the study led by Hervé Sauquet and Jürg Schönenberger, designed to shed new light on the structure of the ancestral flower of all living Angiosperms, 36 researchers worked together during six years to gather the largest data set of floral traits ever assembled (13,444 referenced data points), sampling 792 species from 63 orders (98%) and 372 families (86%) of Angiosperms. We reconstructed the ancestral Angiosperm flower as bisexual and actinomorphic, with more than two whorls of three separate perianth organs each (undifferentiated tepals), more than two whorls of three separate stamens each, and more than five spirally arranged separate carpels (Fig. 3a). Training students and future researchers is a duty and a pleasure. Due to the diversity of personalities and to the different perceptions of science, supervising a student is always a challenge, but a very rewarding task as well. My objective is to help students become more autonomous and develop their curiosity, and most of all encourage them to cultivate their own skills and scientific affinities.

Among the projects I organized and led (eight papers as last author + a pair of papers for which I chose to be the first author [Jabbour and Dressler, 2015;Jabbour et al. 2015a]), three are special to me. The first one was a collaborative study I initiated after I arrived at the MNHN. It was a scientometric study about radiations, focused on insular systems, and not restricted to plants (Soulebeau et al. 2015). Anaëlle Soulebeau was then a PhD student, and six other colleagues (including a postdoc, four Assistant Professors, and one Professor) contributed to the study. It was quite challenging to standardize a protocol for extracting data from the literature and to make everybody respect the deadlines.

The second was a neat and thorough nomenclatural revision for an afro-montane species of Delphinium, co-authored with Stefan Dressler. This work was linked with an ongoing research project but also with my curatorial activities. Stefan was at that time a colleague/curator from Frankfurt Herbarium, and we only knew each other from writing emails to each other. Since then, he visited Paris Herbarium, we designed a project that we submitted to the National Geographic Society, and we will do our first field trip together in Kenya next year.

The third was the first paper of Felipe Espinosa as a first author, Felipe being the first PhD student I am in charge of. Besides the fact that Felipe had to write for the first time a full manuscript, he successfully dealt with collaborators from the MNHN on the one hand, and from Turkey, China, and the Netherlands on the other hand. He is supervised by Prof. Sophie Nadot (Univ. Paris Sud) and me.

As a MSc student under my supervision, he co-authored an article in Flora (Jabbour et al. 2015a). His first PhD paper got accepted in April 2017 in Int. J.

Plant Sci. [START_REF] Espinosa | The Turkish endemic Pseudodelphinium turcicum (Ranunculaceae) : an unusual population of Delphinium with peloric flowers that has persisted in the wild for 20 years[END_REF].

After his PhD defence (planned in May 2019), Felipe will go back to Colombia. Prof. Shahin Zarre (Univ. Tehran) would be a third co-supervisor. These projects were conducted and completed thanks to students I supervised (Fig. 5)

V. EVOLUTION OF FLOWER FORM IN

and in collaboration with a group of reliable and enthusiastic collaborators.

Staying in close contact with some fields (e.g. evo-devo) and initiating projects in This PhD project combined two approaches: 1) phylogenetic comparative analyses of the evolution of selected floral traits to detect correlates of and constraints on the evolution of zygomorphy in Asteridae, and 2) a candidate-gene evo-devo investigation of floral symmetry in selected species of Ranunculaceae.

We demonstrated that the transition between different states of floral symmetry in Asteridae is conditioned by both androecium organ number and perianth merism, and that zygomorphy is a prerequisite to the emergence of spurs (Jabbour et al. 2008).

In selected actinomorphic and zygomorphic species of Ranunculaceae, we described the developmental events leading to the establishment of floral symmetry, and identified the developmental stages at which zygomorphy is settled in Delphinieae (Jabbour et al. 2009).

In the same family, 28 species (18 genera) sampled throughout the phylogeny of Delphinieae were used for characterizing the CYC-like gene sequences and infer their evolution in the tribe and family, and three species with actinomorphic flowers and a couple of species with zygomorphic flowers (Aconitum carmichaelii Debeaux and Consolida regalis Gray) were used to study the expression of the different paralogues during development.

We found that a gene duplication possibly predated the emergence of the Delphinieae lineage, resulting in up to four gene copies in zygomorphic species (a first duplication may have predated the divergence between Lardizabalaceae and Ranunculaceae).

The paralogs were expressed early in floral buds and the duration of their expression varied between species and paralog class. At most one paralog was expressed during the late stages of floral development in the species with actinomorphic flowers whereas all paralogs from the species with zygomorphic flowers were expressed, composing a species-specific identity code for perianth organs (Jabbour et al. 2014; I used for the first time herbarium specimens and got familiar with the herbarium work (see [START_REF] Malécot | L'utilisation des herbiers en phylogénie moléculaire[END_REF]. I built a molecular phylogeny of the tribe Delphinieae, reconstructed its biogeographical history, and found that the particular floral morphology of the Consolida (DC.) Gray/Aconitella Spach genera was the result of a postgenital fusion of two petal primordia (Jabbour and Renner 2011a, 2011b, 2012a, 2012b;Fig. 9). Since then, I continuously updated the phylogeny of Delphinieae, and made the necessary taxonomical changes. This work has been done in close collaboration with Wei Wang [START_REF] Espinosa | The Turkish endemic Pseudodelphinium turcicum (Ranunculaceae) : an unusual population of Delphinium with peloric flowers that has persisted in the wild for 20 years[END_REF], Xiang et al. 2017; Fig. 10). Numbers in parentheses correspond to the number of species in the taxon. Numbers on the branches correspond to the number of fully developed petals in the flowers of the corresponding lineage (see Fig. 9). Coloured bars refer to the corolla type (first column next to the tips of tree; see Fig. 9) and to the sepal shape (second column; spurred, hooded, cylindrical hood, flat).

My research on Delphinieae has been pursued and widened after I got hired at the MNHN. I still use molecular phylogenetics and biogeographical methods to address issues such as lineage evolution and morphological evolution (thanks to morphometrics applied to herbarium flower samples). Together with students, I look now at the evolution of Delphinieae at the infra-subgeneric level. Felipe Espinosa (second year PhD student) investigated the phylogenetic relationships and flower morphology and anatomy in a newly described species from Turkey, Delphinium turcicum [START_REF] Espinosa | The Turkish endemic Pseudodelphinium turcicum (Ranunculaceae) : an unusual population of Delphinium with peloric flowers that has persisted in the wild for 20 years[END_REF].

Marion Chartier (former postdoc at Univ. Vienna) was involved in the study of the afro-montane species of Delphinium (Chartier et al. 2016). Whilst I designed the study and conducted the molecular biology experiments and morphometric measurements on herbarium samples, Marion was in charge of the statistical and morphospace analyses based on the morphometric data (Fig. 11). This study was co-authored by Alfonso We gathered people to work on this very exciting topic, and tackled all these issues in the past few years. Some aspects are still under investigation.

Flower morphology (Jabbour et al. 2015), anatomy (Deroin et al. 2015), and development [START_REF] Zhao | Floral organogenesis of Helleborus thibetanus and Nigella damascena (Ranunculaceae) and its systematic significance[END_REF], Jabbour et al., 2015b) were described and compared between the two morphs, and the molecular basis of the floral dimorphism was unravelled (Gonçalves et al. 2013, Fig. 14). We demonstrated that the floral dimorphism is controlled by a single locus, which perfectly co-segregates with the N. damascena AP3-3 gene. This gene is not expressed in the apetalous morph and exhibits a dynamic expression during early floral development in the petalous morph.

We hypothesized that NdAP3-3 plays a role not only in petal identity but also in meristem patterning, possibly through regulation of perianth organ number and the perianth/stamen boundary.

Although we suffered from competition from another research group (Hongzhi Kong's lab: [START_REF] Zhang | Disruption of the petal identity gene APETALA3-3 is highly correlated with loss of petals within the buttercup family (Ranunculaceae)[END_REF], Wang et al. 2015), we continue our study in order to make it Also, we want to analyze the evolution of the distribution of each morph, to test for the conservation of particular geographic patterns through time. We hence georeferenced more than 3,000 herbarium specimens from 45 herbaria, and could reconstruct distribution maps for each morph, during the last two centuries, and by intervals of 50 years. The gene responsible for the floral dimorphism (NdAP3-3) is currently being sequenced from a sample of herbarium specimens collected from different areas in the Mediterranean Basin, to test whether the apetalous phenotype is always determined by the same molecular factor.

V.d. SYSTEMATICS OF STEPHANIA (MENISPERMACEAE)

The research project about the taxonomical circumscription within Stephania was initiated by my colleague Sovanmoly Hul (Assistant Professor and curator at P). She was the one starting the collaboration with a team of researchers in pharmacology from Marseille, aiming at demonstrating the pharmacological activity of the tubers of some Cambodian species from this genus.

We co-supervised a PhD student, Chhavarath Dary, who studied the systematics of the genus (Hul et al. 2014;Dary et al. 2015;Hul et al. 2017) and the chemical properties of selected species (Dary et al. 2016). Chhavarath reconstructed a molecular phylogeny of the genus (species sampling ratio = 64%) under my supervision. The manuscript has still to be worked on before submission. Building on the world checklist of Stephania we published (Hul et al. 2017), Sovanmoly, Chhavarath and I will now work on the Menispermaceae Flora of Cambodia, Laos and Vietnam.

Stephania is the most species rich of all the genera of Menispermaceae recorded in this region. Chhavarath went back to Phnom Penh where he works as an Assistant

Professor.

During the last years, I had the opportunity to work on Chinese Stephania with other collaborators. We published a molecular phylogeny (sampling ratio = 74% for China, 48% worldwide; Xie et al. 2015), and a study about the diversity of leaf epidermal features in the subset of Chinese species [START_REF] Kang | Leaf epidermal features of Chinese Stephania Lour. (Menispermaceae) and their systematic significance[END_REF].

V.e. FLOWER DISPARITY IN ERICALES

After a PhD in evo-devo (molecular biology techniques, focus on Ranunculaceae), a first postdoc in systematics (molecular biology and herbarium work, focus on Delphinieae), I was eager to address more theoretical issues in the field of flower morphological evolution. This idea was already in my mind during my PhD, and it led to an article about modelling flowers and inflorescences (Jabbour and Citerne 2010).

Reading articles from the abundant literature about morphological evolution in various kinds of organisms (e.g. [START_REF] Foote | Morphologic patterns of diversification: examples from trilobites[END_REF][START_REF] Niklas | Morphological evolution through complex domains of fitness[END_REF][START_REF] Mcghee | Theoretical morphology: the concept and its applications[END_REF]) helped me maturate the project. The idea was to apply mathematical methods to ordinate and analyze the discrete and continuous botanical data.

With this idea in mind, and following the advice Elvira Hörandl (Univ. Göttingen) gave me, I contacted Jürg Schönenberger (Univ. Vienna). Visiting his lab in 2012 was a great opportunity to meet Maria von Balthazar and Yannick Staedler, two of Jürg's collaborators. My idea to look at the profuse botanical data using a mathematical framework was nothing new, and Jürg and Peter Crane (Yale) had already thought about working on Angiosperm morphospace, reanalysing the data set of [START_REF] Stebbins | Natural selection and the differentiation of angiosperm families[END_REF] in the light of the APG classification and applying refined statistical methods.

This project was designed by Jürg Schönenberger and me and got funded by the Austrian Science Fund. It consisted in investigating the evolution of the floral morphospace occupation in the order Ericales. As I got appointed at the MNHN before the start of this project, Jürg and I hired Marion Chartier as a postdoc. Marion started with a review of the studies having looked at plant morphospaces, and reinterpreted under the APG3-light and with modern statistical analyses the pivotal findings of [START_REF] Stebbins | Natural selection and the differentiation of angiosperm families[END_REF]Chartier et al. 2014). Based on a limited number of floral traits, he determined the frequency of each combination of traits that is observable in nature, and he pinpointed the floral organizations that could not be possibly observed, either because he considered them "nearly impossible" (from a developmental or architectural point of view) or "with a low survival". Marion's postdoc project led her to study the order Ericales, an order of 381 extant species with many economically important species (e.g. tea [Theaceae], kiwi [Actinidiaceae], or rhododendrons

[Ericaceae]) and a huge flower disparity. She demonstrated in a very high quality paper (Chartier et al. 2017, Fig. 15) that the Ericales floral morphospace is organized as a continuous cloud in which most clades occupy distinct regions in a mosaic pattern, that disparity increases with clade size rather than age, and that fossils fall in a narrow portion of the space. She also found that among functional modules, it is the androecium that contributes most to total floral disparity in Ericales. Marion became a very good collaborator, and we together analysed the floral morphological disparity of the afro-montane species of Delphinium (Chartier et al. 2016). Now that a species of Delphinium with apetalous peloric flowers [START_REF] Rudall | Evolutionary change in flowers and inflorescences: evidence from naturally occurring terata[END_REF] was found [START_REF] Espinosa | The Turkish endemic Pseudodelphinium turcicum (Ranunculaceae) : an unusual population of Delphinium with peloric flowers that has persisted in the wild for 20 years[END_REF], the description of the genus Delphinium has to be amended and should include, among other characteristics, the following character states: zygomorphy or actinomorphy, dorsal sepal spurred or not, corolla present or absent (petals not developed), corolla consisting of four or a single fully developed petal, when present.

This will be an important taxonomical act as this genus consists of ca. 300 species distributed worldwide. This manuscript will be prepared together with Felipe Espinosa, Thierry Deroin, Pierre-Emmanuel du Pasquier, and James Byng. Felipe and I will be the first and last authors, respectively.

VI.b. FLOWER DEVELOPMENT IN THE GENUS STAPHISAGRIA (RANUNCULACEAE)

The Seeds from the two other species will be sown in order to get floral buds in spring 2018. The comparative analysis of the floral development in the three species will then be conducted. Some very interesting population genetic and pollination ecology studies focusing on this group of species were published (Orellana et al. 2009a(Orellana et al. , 2009b) ) but the floral morphology of the three species was never investigated in detail and compared. Staphisagria macrosperma has the shortest outer spur in the whole tribe Delphinieae.

Describing the floral development in this genus will be crucial to provide a good model to investigate the evo-devo of nested spur formation, the rhythm of spur development in species with contrasted spur lengths, and the physical constraints exerted by the inner spurs on the outer one, and reciprocally.

VI.c. IDENTIFYING THE SISTER GROUP OF THE DELPHINIEAE (RANUNCULACEAE)

To understand flower evolution in Delphinieae and reconstruct the ancestral flower of the tribe, it is required to include the sister group of the tribe in the analyses. The most recent phylogenies of Ranunculales [START_REF] Wang | Phylogeny and classification of Ranunculales: evidence from four molecular loci and morphological data[END_REF]; matrix of molecular [plastid and nuclear] and morphological traits) and Ranunculaceae (Cossard et al. 2016; eight markers from the three genomes; [START_REF] Wang | The rise of angiosperm-dominated herbaceous floras: Insights from Ranunculaceae[END_REF]; one nuclear and five plastid markers) published to date did not solve this issue.

A methodology inspired from [START_REF] Lehtonen | Sensitive phylogenetics of Clematis and its position in Ranunculaceae[END_REF] who found, by simulating many alternative phylogenetic tree topologies under different cost regimes, that Nigella L.

was likely the sister group of the tribe Delphinieae, could be used here with a taxonomic sampling better adapted to our question. This work will be conducted in collaboration with Wei Wang.

VI.d. A SYNTHESIS ABOUT THE RESEARCH DONE SO FAR ON DELPHINIEAE (RANUNCULACEAE)

Many studies about Delphinieae have been published in the last ten years. A wide array of questions have been answered, in fields as diverse as molecular phylogeny (Jabbour and Renner 2012b), flower morphology and anatomy (Novikoff and Jabbour, 2014), evo-devo [START_REF] Rasmussen | One size fits all? Molecular evidence for a commonly inherited petal identity program in Ranunculales[END_REF], Jabbour et al. 2014), teratology (Jabbour et al. 2015a), pollination ecology [START_REF] Duan | Insect and wind pollination of an alpine biennial Aconitum gymnandrum (Ranunculaceae)[END_REF], karyology [START_REF] Bosch | DCDB: an updated on-line database of chromosome numbers of tribe Delphinieae[END_REF], and pharmacology [START_REF] Xiao | A pharmacophylogenetic study of Aconitum L. (Ranunculaceae) from China[END_REF].

The taxonomic circumscription of many groups within Delphinieae has been particularly impacted by the regular updates of the phylogeny of the tribe. For instance, the genus Staphisagria was resurrected (Jabbour and Renner 2011b),

Pseudodelphinium was sunk into Delphinium [START_REF] Espinosa | The Turkish endemic Pseudodelphinium turcicum (Ranunculaceae) : an unusual population of Delphinium with peloric flowers that has persisted in the wild for 20 years[END_REF], Consolida and Aconitella are nested in Delphinium (Jabbour and Renner 2011a) and will be synonymized with Delphinium, Gymnaconitum was taken out from Aconitum [START_REF] Wang | Gymnaconitum, a new genus of Ranunculaceae endemic to the Qinghai-Tibetan Plateau[END_REF], and Delphinium sect. Delphinium was erected to the subgenus level (Xiang et al. 2017). The advances in floral evo-devo, and the new data about the morphology and anatomy of Delphinieae flowers, and the very abundant literature about the chemical properties of species from this tribe now have to be synthesized.

Furthermore, new data about flower modularity and integration were obtained by Yiyun Chen (Masters student, co-supervised by Sylvain Gerber and I) at the end of the summer 2017 (Fig. 16). These results obtained with geometric morphometrics on flowers from herbarium specimens will be included in the Special Issue of Botany Letters I co-organize with Guillaume Besnard (CNRS Toulouse).

On the long term, Wei Wang and I plan to revise the taxonomy of the genus Delphinium (ca. 300 species), building on our publications on the group. The last treatment of the genus was made by [START_REF] Malyutin | The system of the genus Delphinium (Ranunculaceae) based on the morphological features of seeds[END_REF] based on seed characters. The recent re-circumscription of some infragenerical taxa will be taken into account, and we expect that many species will need to be synonymized.

VI.e. EVO-DEVO OF PERIANTH DIMORPHISM IN NIGELLA DAMASCENA (RANUNCULACEAE)

The teams of Catherine Damerval and of Sophie Nadot, and the MADP team will submit a project to the ANR in September 2017. This project will include an investigation of perianth organ anatomy and development in both floral morphs, as our recent anatomical observations showed that the composition of the perianth of the apetalous morph is more complex than previously thought. Additionally, after having reconstructed the evolution of the distribution area of both floral morphs during the last three centuries thanks to an excellent sampling of the world collection of N.

damascena herbarium specimens (Jabbour et al. in prep), we will test whether the apetalous morph is always (in time and space) caused by the insertion of a transposon at the NdAP3-3 locus. Preliminary results suggest that it is not the case.

This project will be extended to other models of Ranunculaceae in order to compare the morphogenesis of perianth organs, and especially petals, at the family level.

VI.f. COMPARATIVE STUDY OF PETAL DEVELOPMENT IN

RANUNCULACEAE

This project was designed by Catherine Damerval and me to investigate the developmental and cellular processes involved in petal form diversity. The genesis of petal morphology in some representative genera of Ranunculaceae will be studied by combining morphological and evo-devo approaches.

For this purpose, and to understand the mechanisms responsible for the cellular and morphological development and growth of petals, the PhD student will apply anatomical investigations and histological analyses using microscopic observations, quantitative measurements as well as geometric morphometrics (or CT-scanning if possible). The sampling would include ca. 10 species representing various tribes of the Ranunculaceae differing in corolla aspect and petal morphology. Species from the Delphinieae, from the genera Aconitum, Consolida, and Delphinium will be selected.

Interestingly, this tribe comprises more than one-quarter of the species of the family, all being characterized by zygomorphic flowers but differing in perianth structure and composition.

For comparison, species will be selected from other genera showing a broad range of petal morphology (e.g. Adonis L., Aquilegia, Helleborus L.). In order to examine how different petal forms might evolve in correlation with changes in the regulation of cell division and expansion, the student will carry out in situ hybridization experiments to discriminate territories of gene expression relating to the cell division and cell differentiation processes by tracking histone H4 in DNA-replicating cells [START_REF] Fobert | Patterns of cell division revealed by transcriptional regulation of genes during the cell cycle in plants[END_REF]).

This and their location in the phylogenetic tree. This result will inspire evo-devo studies investigating the development of such secretory structures and will facilitate the selection of the taxa with similar morphologies (but from different lineages) to be investigated. Although we have at hand a preliminary database, this project has not started yet.

I was asked by the journal Botany Letters (formerly Acta Botanica Gallica, and initially known as Bulletin de la Société Botanique de France) to pick in their archives one of their highly cited papers, to translate it into English, and to add a concise synthesis of the research conducted so far on the same topic.

In order to fuel this project on secretory structures in Angiosperms, I selected an article entitled: "Etude sur l'anatomie et la physiologie des nectaires", by Gaston Bonnier (1878, Bulletin de la Société Botanique de France).

VI.h. PHYLOGEOGRAPHY OF DELPHINIUM IN THE EAST AFRICAN RIFT SYSTEM

The investigation of flower morphological diversity, phylogeny and biogeography of The newly collected data will be analysed together with my collaborators in this project, as well as with Guillaume Besnard (CNRS Toulouse, http://www.edb.upstlse.fr/Besnard-Guillaume.html) who is an expert in conducting genomic studies on herbarium specimens.

These prospective studies will only be possible with the contribution of my European, Asian, and African collaborators. Each of these studies will give us the opportunity to train students in various disciplines, from plant anatomy to phylogeny, and from taxonomy to the history of science.

VI.i. LONG-TERM PROJECTS

On the long term, I would like to train myself in embryology and palaeobotany in order to be able to answer in a more integrative way questions in the fields of plant systematics, phylogeny, and morphological evolution. I am already in contact with a paleobotanist (Dario de Franceschi) and an archeobotanist (Aurélie Salavert) at the MNHN, and it would be relatively easy for me to learn from them and look at the very rich fossil plant collection. I feel that getting familiar with embryology would be a more difficult task. It would probably be necessary to spend some time in a specialized laboratory where I could learn the techniques and get intensive training.

I would also like to invest some time in thinking about science and its evolution Choosing to build a team based on research projects about plant morphology, anatomy and development have some obvious effects. Even if this is not applicable to all our research projects, publishing in high impact factor journals can be difficult. Getting funding for plant morphology-oriented projects could be even more difficult in the current scientific context in France, or even in Europe (Jürg Schönenberger and I were extremely happy to get a FWF funding from the Austrian government to work on the evolution of the floral morphospace).

Students of plant morphology have to face many prejudices and have to repeatedly answer the same critics and explain why this discipline is important in the botanical research of the 21 st century. How many times did I hear the following sentences:

 Plant morphology is so old-fashioned!  When you study plant morphology, obtaining results is such a long process!  You cannot obtain ground-breaking results (anymore) in this field.

 You use simple (simplistic) and old-fashioned methods.

 The objects you study are less noble or less complex then, let's say, a genome, or a phylogenetic tree with hundreds of accessions.

This discipline may seem unrewarding at first sight, owing to the time one should invest in the dissections, experiments, and observations, and to the difficulty to publish the results in high impact journals. However, studying the morphology, anatomy, and development of plants can be fascinating. The community of botanists studying flower morphology is not large, and it is possible for a flower morphologist in his/her middle career to know and to have contacts with all the other members of the community (Fig. 20). of plant morphologists will be trained and will find positions in line with their expertise. The opposite scenario would involve a decrease in the number of experts in plant morphology, and consequently of teachers and mentors. The new generations of botanists would overlook the discipline of plant morphology (this trend is already noticeable as some courses in botany disappear in many Universities). Locally, this trend may lead to the loss of expertise in plant morphology-anatomy. This situation may last several decades before this discipline would be re-discovered, knowing that building new group of experts is not straightforward, especially in plant morphologyanatomy. Indeed, (but I am perhaps wrong), the role of mentors and colleagues is crucial in this discipline, for the transmission of the experimental know-how and of the theoretical knowledge (evolution of concepts, schools of thought, knowledge of recent and old publications, possibly in different languages).

In my opinion, the most realistic scenario would be a compromise between the two previous ones, and highly depends on the personal involvement of individuals rather than on an improvement of the funding policies or the scientific priorities of Universities, although these two aspects are important. Researchers in the field of plant morphology should 1) work on multiplying the connections of their network, 2) teach and train more students and make them travel within their network to get familiar with different methods and "families" of researchers, 3) build collaborations to be more successful when applying for funding. Teaching is essential as this is the only way to show the students that their prejudice against plant morphology and anatomy are unfounded. Students and interns should be wisely trained, i.e. trained in a way that they know as well the bases of evolutionary biology, evo-devo, systematics, physiology, and paleobotany. Such disciplines are a chance for plant morphology, as they highlight its importance and arouse new approaches and concepts. Researchers should invest in the young ones at early stages in their career.

As plant morphologists-anatomists, it is our duty to go towards the other researchers from other disciplines. If they do not include any investigation of plant morphologyanatomy-development, it is often because they do not know how crucial the results would be to better understand their model system and target the most useful structure and/or developmental stage for their own study.

VIII. INVOLVEMENT IN THE DIFFERENT TASKS OF THE MNHN

Figure 1 :

 1 Figure 1: Personal memories. A book about trees and shrubs that belonged to my grandfather, in which a drawing of a Gingko leaf, handwritten notes, and dried plant parts were slipped.

  Starting in 2005 with a MSc on the molecular phylogenetics of phytoparasitic fungi, I embarked on a PhD project focusing on the evo-devo of floral symmetry under the supervision of Sophie Nadot and Catherine Damerval, before doing a postdoc in plant systematics. I can now say that my passion for flower morphology and development was triggered by a visit to the Royal Botanic Garden Edinburgh in 2017, where I conducted my first study of floral development together with Louis Ronse De Craene. I could not invest more time in morphology-development during my PhD, as the evodevo experiments were time-consuming. During my postdoc in the lab of Susanne Renner, I conducted a developmental study, after re-building on my own a criticalpoint drying system from separate parts, and I insisted on writing a paper based on flower morphological evolution (Jabbour and Renner 2012a). The second postdoc project I designed with Jürg Schönenberger was exclusively based on flower morphology.

BOX 1 :

 1 Biography and works of Jean-Baptiste Payer (1818-1860, †42) Illustration: Payer JB (1857) Traité d'organogénie comparée de la fleur. Planche 56, Nigelles. Portrait: http://www.asfeldjuzancourt.fr/le-saviez-vous.html Jean-Baptiste Payer comes from a family of millers from the North-East of France. He starts his academic career in 1840 as professor of geology and mineralogy in Rennes. In 1841, he becomes Assist. Prof. in Botany at the Ecole Normale Supérieure in Paris. In 1848, he is involved in the French Revolution, and takes afterwards political responsibilities until the 1851 coup. He then dedicated himself fully to Botany. In 1852, he becomes professor of plant organography at the Faculty of Sciences of Paris, taking over from Auguste de Saint-Hilaire. Although Payer's contributions to botany are numerous, his most famous work is indisputably the Traité d'organogénie comparée de la fleur. This book, compiling personal observations on floral development for most Angiosperm orders, was published in 1857 (Payer was then 39!), after 13 years of dissecting floral buds, observing the preparations under a binocular, comparing and synthesizing his observations and presenting hypotheses of homology among structures from different taxa.

BOX 2 :

 2 Traité d'organogénie has been cited more than 500 times (G-Scholar, August 2017), in articles focusing on floral development, morphology, floral evolution, plant systematics, or evo-devo. Payer's Traité is an unmissable work for all students of flower morphology and development. References  Bibliothèque National de France: http://data.bnf.fr/10533055/jean_baptiste_payer/  Payer JB (1857) Traité d'organogénie comparée de la fleur. Paris: Masson. Biography and works of Wilhelm Hofmeister (1824-1877, †53) Portrait: https://fr.wikipedia.org/wiki/Wilhelm_Hofmeister Illustration: Hofmeister W (1859) Neue Beiträge zur Kenntniss der Embryobildung der Phanerogamen. XXIV Lamium purpureum.

BOX 3 :

 3 Biography and works of Philippe Van Tieghem(1839-1914, †75) Portrait from[START_REF] Bonnier | L'oeuvre de Philippe Van Tieghem[END_REF]. General and close view of the histological slides of the Van Tieghem collection at the MNHN. Philippe Edouard Van Tieghem started to work on microscopic fungi in the laboratory and under the supervision of Louis Pasteur. He gave his name to a device -the Van Tieghem cell -that allows following the mycelium development under a microscope. Van Tieghem joins the MNHN in 1878. His contribution to the study of plant anatomy in France was essential in many aspects (Van Tieghem 1891). Looking at the inner structures of plants, he aimed at elaborating hypotheses of homology. He



  Guédès M (1973) Goethe et Geoffroy Saint-Hilaire. Histoire et Nature 1: 27-45.  Guédès M (1979) Morphology of seed plants. J. Cramer. BOX 5: The evolution of imaging techniques for the study of plant morphology Left: Ranunculaceae flower meristem showing the bract and the bracteoles. Drawing from Payer (1857, plate about Nigella) and SEM image of an Aquilegia alpina L. flower bud. Right: Stereomicroscope (bought in 2013) and antique microscope (19 cm high, bought on a flee market in 2017).

Figure 2 :

 2 Figure 2: Relations of (plant) morphology with other disciplines. Links among disciplines are shown with double lines. All disciplines in relation with morphology are potentially linked to one another.

Figure 3a :

 3a Figure 3a: The hypothetical ancestral flower of Angiosperms (Sauquet et al. 2017).

Figure 3b :

 3b Figure 3b: The hypothetical ancestral flower of Angiosperms. Maximum likelihood ancestral state reconstruction of functional sex of flowers in Angiosperms, showing that bisexual flowers are ancestral and that unisexual flowers evolved many times independently. The pie charts at the centre of the figure indicate the proportional likelihoods for reconstructed ancestral states at 15 key nodes.

Figure 3c :

 3c Figure 3c: The hypothetical ancestral flower of Angiosperms. Scenario for the earliest phase of floral diversification as inferred from the analyses of Sauquet et al. (2017). Floral diagram colour code: light green = undifferentiated tepals; green = sepals; yellow = petals; red = stamens; blue = carpels.

  ., Univ. Phnom Penh The PhD project of Chhavarath focused on the systematics of the genus Stephania Lour. (Menispermaceae) and on the pharmacological properties of some species of Stephania found in Cambodia. He was supervised by Prof. Evelyne Ollivier and Dr. Sok-Siya Bun at the University of Marseilles, and by Dr. Hul and me at the MNHN. His PhD work led to four publications so far, and we still collaborate on his PhD topic to publish his last results. He is since September 2016 an Assistant Professor at the University of Phnom Penh. evolution of floral structures and floral organization in the Ranunculaceae, combining morphology-anatomy, development and molecular phylogeny.

  a MSc student under the supervision of Dr. Sylvain Gerber (MNHN) and me. She conducts a comparative analysis of floral shapes in the tribe Delphinieae, using geometric morphometrics, and based on herbarium specimens. She will defend her MSc thesis in September 2017. Yiyun will then go back to China where she will work at a panda conservation centre. PhD contract at the MNHN Masoumeh contacted me one year ago to build a PhD project combining floral development and evo-devo. Together with Dr. Catherine Damerval (CNRS, Le Moulon-Génétique Quantitative et Evolution), we designed a project addressing the possible genetic causes of differential petal development in Ranunculaceae. Masoumeh was invited to the competition of the Doctoral School of the MNHN in July 2017. She was unfortunately not selected, but she would still have the possibility to work in our team in case we get the ANR grant in early 2018.

  the evolution of flower forms in Angiosperms since my time as a PhD student. I tackled this issue through different disciplines such as comparative analyses, evo-devo, systematics, and morphology-anatomy. Figure4shows the different Angiosperm taxa I have investigated so far.

Figure 4 :

 4 Figure 4: The taxa I have investigated so far. The Angiosperm tree topology is from APG IV (2016).

  fields I was not trained in (e.g. pharmacological and chemical properties of plants) are important to me. It allows getting a more comprehensive understanding of the evolutionary processes having shaped current biodiversity. On a more personal ground, it helps me discovering new aspects of science and meeting new collaborators with different ways of apprehending morphological evolution. Whilst supervising students and transmitting the knowledge I have accumulated through the years is very satisfactory, I like to remain in the mental position of the student, pushing myself to discover new areas of botany (see paragraph VI.i.), and always trying to question my own reasoning, especially in fields where my personal scientific routine got settled.

Figure 5 :

 5 Figure 5: Synthesis of the main research projects I was involved in or led so far. The involvement of my former and current students in each project is shown.

Fig

  Fig.6).

Figure 6 :

 6 Figure 6: RANACYL genes: Phylogenetic relationships and expression pattern. Phylogenetic tree of the CYC-like sequences in Ranunculaceae (the lineages of species with zygomorphic flowers are highlighted in pink) and expression level of the RANACYL paralogs for Aconitum carmichaelii and Consolida regalis in the different perianth compartments (Jabbour et al. 2014). D: dorsal, L: lateral, V: ventral.

Figure 7 :

 7 Figure 7: Theoretical developmental trajectories combining different states for three processes (organ initiation, growth, and differential elaboration) resulting in different types of floral symmetry. See Citerne et al. (2010) for details.

Figure 8 :

 8 Figure 8: Delphinieae floral groundplan; a. Inflorescences of Delphinium elatum L. (Munich botanical garden), b. Antique dolphin fountain, c. Delphinium flower bud, resembling the shape of dolphin as depicted in antique art, d. Flower of Delphinium elatum, e. Dorsal spurred sepal of the same flower, put aside to show the pair of dorsal spurred petals. One lateral and one ventral sepal were removed, f. Petal organisation in Delphinium, showing the proportion of each part of the petal. Red: spur, green: labium, blue: stalk, g. Aconitum lycoctonum subsp. moldavicum (Hacq.) Jalas, h. Dorsal spurred sepal of the same flower, put aside to show the pair of dorsal spurred petals, i. Petal organisation in Aconitum L., showing the proportion of each part of the petal. Red: spur, green: labium, blue: stalk.

Figure 9 :

 9 Figure 9: The types of corolla development identified in Delphinieae (Jabbour et al. 2009, Jabbour and Renner 2012a); a. Delphinium elatum (credit: F. Espinosa), b. Staphisagria macrosperma Spach, c. Aconitum napellus L., d. Gymnaconitum gymnandrum (Maxim.) Wei Wang & Z.D.Chen (Wang et al. 2013), e. Consolida regalis (credit: A. Gröger), f. Aconitella hohenackeri Soják (credit: A. Gröger), g. Delphinium turcicum (H.Duman, Vural, Aytaç & Adigüzel) Espinosa (credit: F. Espinosa). Photographs are not to scale. Green: developing petals; orange: nectar spur.

Figure 10 :

 10 Figure 10: Improvements in the phylogeny of Delphinieae since 2010. This date corresponds to the start of my postdoc during which I built the first molecular phylogeny of the Consolida+Aconitella group (Jabbour and Renner 2011a), and then the first molecular phylogeny of the tribe Delphinieae (Jabbour and Renner 2011b, 2012a). My colleague Wei Wang built on my results one year later (Wang et al. 2013), focusing on the phylogenetic placement of the monotypic genus Gymnaconitum (Stapf) Wei Wang & Z.D.Chen. The subsequent studies about infratribal taxa in Delphinieae were conducted in collaboration with Wei Wang and his team (Espinosa et al. 2017, Xiang et al. 2017). The three species of Staphisagria J.Hill were once included in the genus Delphinium. Delphinium turcicum was formerly published as Pseudodelphinium turcicum H.Duman, Vural, Aytaç & Adigüzel (Vural et al. 2012).

  Rojas, a student I supervised when I was a postdoc. Other collaborators from the MNHN (Corinne Sarthou) or from other institutions (Stefan Dressler [Frankfurt], JürgSchönenberger [Vienna], WeiWang [Beijing]) were associated to this study.I remember an advice Sophie Nadot and Catherine Damerval gave me when I was a PhD student, and this advice had a great impact on the trajectory of my research in the few years following my defence. I hope I translate it here without too many changes: "Don't spend so much time doing development or morphology. Even if this is interesting, this will not bring big papers. Better focus now on your evo-devo experiments, you need molecular papers in your CV. Nobody will offer you a permanent position nowadays with only morphology and development in your CV". I followed this advice, and kept the proportion of morphological studies relatively low during my PhD and postdoc, although it was important for me to build some minimal expertise in this field(Jabbour et al. 2009; Jabbour and Renner 2012a). I now have the opportunity to dive into the fields of morphology, development and anatomy, to train students in such disciplines (Felipe Espinosa, Lucile Jourdain-Fiévet), and to build projects with collaborators who are keener on the investigation of the molecular causes of floral phenotypes.

Figure 11 :

 11 Figure 11: The morphospecies of afro-montane Delphinium and their phylogenetic relationships. PCA showing the circumscription of the three morphospecies based on the analysis of 13 metrics taken from flowers from herbarium specimens. See Chartier et al. (2016) for details.

Figure 11 :

 11 Figure 11: Continued. Plastid phylogeny of the afro-montane morphospecies of Delphinium. The two clades (blue and grey) are depicted on the map of Africa. The asterisks on the map represent monophyletic sets of accessions. The hypothesis of a convergent evolution towards a D. leroyi flower type in both the blue and grey clades due to ecological constraints was proposed.

Figure 12 :

 12 Figure 12: Floral vascular organization in three species representative of the three main floral types (with a developed corolla) found in Delphinieae (Novikoff & Jabbour 2014). Credits: A. lasiocarpum Gáyer (http://www.kul.pl/gal_12532.html), D. elatum (Felipe Espinosa).

Figure 13 :

 13 Figure 13: The corolla of a developing flower of Delphinium elatum. View from above. Dissected floral bud showing the corolla consisting of four ventral petals with an arrested development, two flat dorsolateral petals, and two spurred dorsal petals.

Figure 15 :

 15 Figure 15: The floral morphospace of Ericales. (a) Phylogenetic relationships among ericalean families. Pictures of six of the fossil genera included in the analyses. Assigned positions of the fossils are highlighted in (a) by superscript letters on the family names. (b) Disparity. (c) 2D-morphospace representation using PCA. Black dots: species of suprafamilial clades or families; grey dots: all remaining ericalean species. (d) Floral diversity in Ericales: from top to bottom: Satyria sp. (Ericaceae), Sarracenia flava L. (Sarraceniaceae), Symplocos pendula Wight (Symplocaceae), Schima superba Gardner & Champ. (Theaceae), Anneslea fragrans Wall. (Pentaphylacaceae), Primula officinalis (L.) Hill (Primulaceae), Cantua quercifolia Juss. (Polemoniaceae), Couroupita guianensis Aubl. (Lecythidaceae), Impatiens paucidentata De Wild. (Balsaminaceae), Mitrastemon matudae Yamam. (Mitrastemonaceae). (e) Position of the nine fossil species (black dots) in the morphospace (grey dots).

  genus Staphisagria is the earliest diverging in the tribe Delphinieae. It consists of three Mediterranean species, two of them (S. requienii (DC.) Spach and S. picta (Willd.) F.Jabbour) being narrow endemics. Material from S. macrosperma was collected in spring 2015 and spring 2017 in the Botanical Garden of the MNHN, Paris.

Figure 16 :

 16 Figure 16: Geometric morphometric study of Delphinieae floral shape based on herbarium specimens: Example of species with Aconitum-like flowers; a. Specimen of A. japonicum Thunb. from KRA Herbarium (Jagiellonian University, Kraków, Poland), b. Trans-illuminated inflorescence, c. Landmarks put by Yiyun Chen on a dorsal petal, d. 2D geometric morphometric analysis of shape variation.Principal component 1 (PC1) and PC2 account for 49% and 13% of the total variance in the sample, respectively. Virtual flower shapes were produced to visualize the sample mean shape (centre of the graph), as well as modes of floral shape variation along PC1 and PC2. Pictures in a and b, credit: Andriy Novikoff.

  comparative study on growth and development of petals should help reconstructing ancestral and derived developmental modules involved in the evolution of floral form in Ranunculaceae. Compared with results in other Angiosperm species, this study will help understanding how the orientation and timing of cell division and predominant growth directions contribute to the origin of flower variation within Angiosperms.VI.g. EVOLUTION OF EXTRA-AND INTRAFLORAL NECTARIES INANGIOSPERMSMany of the plant taxa I use for my research are species that reward their pollinators with nectar. For instance, Aconitum and Delphinium flowers have nectar spurs, whereas small pocket-like petals are found in Nigella. Many more examples of genera and species characterized by flowers with secretory structures can be found in the literature (e.g. Diascia Link & Otto, Satyrium Sw.,[START_REF] Johnson | Diverse pollination systems of the twin-spurred orchid genus Satyrium in African grasslands[END_REF] Monocots, Smets et al. 2000, see Fernández-Mazuecos and Glover, 2017, Fig. 17

  ).I would like to gather all existing data relative to extra-and intrafloral secretary structures from the literature, and map the different traits on a phylogeny of Angiosperms. We would get crucial information about the number of transitions towards each type of structure (e.g. nectar pouch, nectar spur, oil spur, nectar canals)

Figure 17 :

 17 Figure 17: Evolution of spurred flowers in Angiosperms. Pictures of four species producing twinspurred flowers. a. Diascia integerrima E.Mey ex Benth. (Scrophulariaceae) and its pair of oil spurs (http://www.randomharvest.co.za/South-African-Indigenous-Plants/ShowPlant/PlantId /259?Plant=Diascia+integerrima+Pink), b. Delphinium elatum (Ranunculaceae) and the pair of inner nectar spurs, c. Satyrium sceptrum Schltr. (Orchidaceae) with its pair of nectar spurs, d. Satyrium macrophyllum Lindl. with its pollinator, the long proboscid fly Prosoeca ganglbaueri. c and d from Johnson et al. (2011). Mirror trees of the Asteridae showing the evolution of perianth symmetry and the number of spurs optimized using the maximum parsimony method. (A) Evolution of perianth symmetry. Black: actinomorphy; red: zygomorphy. (B) Evolution of the number of spurs. Black: 0 or equal to perianth merism; red: inferior to perianth merism. Coloured lines and numbers refer to the orders of Asteridae: 1,

  the afro-montane Delphinium provides excellent bases for further studies focusing on the population genetics and on the phylogeography within the group, possibly at finer geographical scales. In addition, the hypotheses of hybridization among morphospecies, and of the evolutionary convergence towards the D. leroyi-like flower morphology in the two clades of afro-montane Delphinium, have still to be tested. New collections are needed, and the first trip in East Africa will take place in Kenya and Malawi, thanks to a funding form the National Geographic Society, in collaboration with Stefan Dressler (Senckenberg Museum, Frankfurt; Fig. 18), Itambo Malombe (Director of the EA Herbarium, Nairobi), and Esther Kioko (Head of Zoology Department, National Museums of Kenya; for the observations of floral visitors).

Figure 18 :

 18 Figure 18: Collecting afro-montane Delphinium in Kenya and Malawi in 2018. Distribution area of the three morphospecies of afro-montane Delphinium. The high altitude areas (in Kenya and Malawi)

(

  probably with my colleagues Thierry Deroin and Corinne Sarthou, see[START_REF] Lamy | Auguste de Saint-Hilaire (1779-1853): un botaniste français au Brésil / Auguste de Saint-Hilaire (1779-1853): um botânico francês no Brasil[END_REF]), particularly about the evolution of concepts in plant morphology andVII. THE MANY FUTURES OF PLANT MORPHOLOGY AT THE MNHN (AND BEYOND)Many of the botanists at the MNHN study plant morphology to complement a study focusing on a phylogeny of a group of taxa or to bring evidences for a taxonomic treatment. So the knowledge about plant morphology exists, is shared among colleagues, and is transmitted to the interns and students. However, detailed studies designed to answer a question about the evolution of morphological or anatomical structures, or studies aiming at discussing a concept in the fields of plant morphology and plant anatomy are less common, as they require genuine expertise. The team Morphology-Anatomy and Development of Plants is the only team in the Botany section, and even in the Institute of Systematics, Evolution and Biodiversity that makes clear that its members are the local experts in these fields. We made this choice to highlight our assets, and show our originality, although we conduct research in the fields of plant systematics, plant development in space, and history of sciences.

Figure 20 :

 20 Figure 20: Contemporary (floral) morphologists-anatomists throughout the world. See p. 80 for a focus on Europe. I offer my apologies to those I have forgotten.

  At the MNHN, the Assistant Professors and Professors are assigned five missions: conducting research, curating collections, teaching, making expert assessments, and popularizing science. I kept this part deliberately short as the Habilitation thesis focuses on research. VIII.a. RESEARCH TEAM RESPONSIBILITY My research projects are developed in § V and VI. I add here information about my recent experience as a team leader. Since January 2016, I share with Thierry Deroin the responsibility of the MADP (Morphology, Anatomy, and Development of Plants) team. The team consists of five permanent members: three Assistant Professors, one engineer, and one technician. We investigate the morphology, anatomy and development of plants (mostly flowers, but also roots) in a phylogenetic framework, in model (Arabidopsis thaliana (L.) Heynh., Aquilegia sp.) or non-model (e.g.Hyacinthoides sp., Delphinium sp.) species. Our objectives are to reconstruct the evolution of our structures of interest (e.g. the shoot meristem, the floral meristem, the carpel) and to study the impact of the environment (contrasted habitats, microgravity and spatial radiations) on their development.We submitted earlier in 2016 a project to a call for proposals of the city of Paris, in order to get an additional funding for our study on the impact of radiations on the development of the plants in spatial conditions.Next spring, the MADP team will ask the MNHN for the opening of an Assist. Prof. position, to fill important needs in teaching plant morphology and anatomy at the MNHN. One option would be to design the research part of the job description focusing on the evolution of the morphological-anatomical structures of plants and on their development, in relation with environmental constraints, and based on living material and on the herbarium, spirit and slide collections. The taxonomic and geographical focus of the position will be specified if necessary. The Assist. Prof. would work as a curator as well.VIII.d. EXPERT ASSESSMENTSI reviewed manuscripts submitted to 18 scientific journals, and proposals submitted to different authorities at the Museum and to the Polish National Science Centre. I am currently an associate editor for the journal Botany Letters (formerly Acta Botanica Gallica), and a member of the commission for scientific and technical literature at the French Centre National du Livre. This public organization financially helps the work of editors, authors and translators. In each commission, a dozen of members are designated for three years.VIII.e. SCIENCE POPULARIZATIONTogether with my colleagues at the MNHN, I contributed to the permanent exhibition in the Botany building. The team of curators also worked on a book entitled "L'Herbier du Muséum"[START_REF] Aupic | L'Herbier du Muséum -L'aventure d'une collection[END_REF], and since I got hired at the Museum, I participated each year in the "Fête de la Science" national event, during which labs open their doors to the public and present their work. In 2018, an open-air exhibition in the Jardin des Plantes will highlight projects currently conducted at the Museum.Our project about the floral dimorphism of Nigella damascena got selected (see Fig.21).

Figure 21 :

 21 Figure 21: Science popularization at the MNHN. The poster and panel (SEM picture) that will be displayed in the Jardin des Plantes at the MNHN, explaining the floral dimorphisms in Nigella damascena and the purpose of studying the development of the floral buds. The bottom picture was taken during a previous exhibition in 2016 called "Planète Grenouille" (Credit: Cyril Ruoso, http://www.cyrilruoso.com/?Planete-grenouille).

Figure 21 :

 21 Figure 21: Continued.

of the Dept. of Structural and Functional Botany,

	Vienna, Austria. Talk Jabbour F and Renner S. Evolutionary trends in
	Delphinieae (Ranunculaceae). A focus on life histories and perianth
	structure.
	 Botany 2011, St. Louis, USA. Talk
	Jabbour F, Rojas A, Renner S. A phylogeny of the Delphinieae
	(Ranunculaceae) reveals that Aconitum is embedded in Delphinium:
	implications for flower evolution.
	Mediterranean area.
	 25 th Plant Biology symposium, University of California, Riverside,
	USA. Poster and talk
	Jabbour F, Nadot S, Damerval C. Developmental features and
	evolution of zygomorphy in the tribe Delphinieae (Ranunculaceae).
	 2 nd Euro Evo Devo meeting, Ghent, Belgium. Poster
	Jabbour F, Ronse de Craene L, Nadot S, Damerval C. The
	developmental bases of floral symmetry in Ranunculaceae. Constraints
	on the evolution of zygomorphy in basal eudicots.
	 11

Jabbour F and Renner S. Systematics, biogeography, and perianth evolution in the Delphinieae (Ranunculaceae). ** Seminars  7 th Int. Congress of Systematic and Evolutionary Biology, Berlin, Germany. Talk Jabbour F and Renner S. The spatiotemporal unfolding of the Delphinium/Consolida clade (Ranunculaceae) in the eastern

th Evolutionary Biology Meeting, Marseille, France. Talk

	Jabbour F, Damerval C, Nadot S. Comparative evolution of floral
	symmetry in monocots, basal and core eudicots. A morphological and
	phylogenetic approach.
	 6 th

Biennial Meeting of the Systematics Association, Edinburgh,

	Scotland. Poster
	Jabbour F, Nadot S, Damerval C. Diversity and evolution of
	CYCLOIDEA-like genes in Ranunculaceae -Papaveraceae.
	 Annual

meeting of the French Systematics Society, Paris

  , France.

	Jabbour F, Le Gac M, Giraud T. Inferring phylogenies of
	Microbotryum violaceum and of its host plants for delimiting parasite
	species and study cospeciation.
	TABLE 1: Publications
	Total number = 34, number with IF ≥ 1 = 28, G Scholar: H = 11, citations > 420
	Poster

Position in the sequence of authors Journal IF # First Contributing author

	I
						Last	Following a
							supervision
	Int. J. Plant Dev. Biol.	-	1	1
	Modern Phytomorphology	-	2			2
	Edinburgh Journal of Botany	-	1			1	1
	Kew Bulletin	0.6 1		1
	Australian Systematic Botany	0.8 1		1
	Phytokeys	1.1 1	1
	Comptes Rendus Biologies	1.1 1	1
	Flora	1.1 1	1			1
	Phytotaxa	1.2 3	1		2	2
	Plant Systematics and Evolution	1.2 1		1
	Advances in Botanical Research	1.3 1		1
	International Journal of Plant Sciences	1.7 2	1		1	1
	Natural Product Research	1.8 1		1		1
	Organisms Diversity and Evolution	2.3 1			1
	Botanical Journal of the Linnean Society 2.5 1	1
	Taxon	2.5 3	1	1	1	2
	Plos One	2.8 2	1	1
	BMC Evolutionary Biology	3.2 1		1
	Annals of Botany	4.0 2	2
	Molecular Phylogenetics and Evolution	4.4 2	1	1
	Nature Scientific Data	4.8 1		1
	Proceedings of the Royal Society B	4.9 1		1		1
	Plant Journal	5.9 1		1
	New Phytologist	7.3 1		1		1
	Nature Communications	12.1 1		1
	Webpage (will be transferred to a new website in autumn 2017):
	http://isyeb.mnhn.fr/annuaire-et-pages-personnelles/pages-personnelles/Florian-
	JABBOUR,325
	Team webpage (will be transferred to a new website in autumn 2017):
	http://isyeb.mnhn.fr/departements/departement-systematique-et-172/morpho-anatomy-
	and-development-of/

. MY SCIENTIFIC TRAJECTORY (OR HOW I BECAME A LIFELONG

In vivo live imaging based on Epi-Illumination Light Microscopy, ELM

  Allows in vivo observations and retrieves high structural details. It allows for instance the direct measurement of phyllotactic parameters. 3. Barabé D, Lacroix C, Jeune B (2007) Following the initiation and development of individual leaf primordia at the level of the shoot apical meristem: the case of distichous phyllotaxis in Begonia. Annals of botany 99(3): 555-560. (2014) 1. Non-destructive, it requires a single growing meristem. The drawback of live ELM is the difficulty in keeping the tissue healthy for several days. 2. It reveals the absolute time of organ initiation and duration of growth.

	2. 2D observations at a magnification reaching 500,000x.
	3. Antoń S, Kamińska M (2015) Comparative floral spur anatomy and nectar
	secretion in four representatives of Ranunculaceae. Protoplasma 252(6): 1587-
	1601.
	3. Jerominek M, Bull-Hereñu, K, Arndt M, Claßen-Bockhoff R (2014) Live
	Confocal microscopy (1950's) imaging of developmental processes in a living meristem of Davidia involucrata
	1. It is an optical microscope reconstructing 3D structures from sets of images Light sheet microscopy (2000's) (Nyssaceae). Frontiers in Plant Science 5: 613.
	obtained at different depths within a thick object. It uses reflected light or 1. It achieves optical sectioning by the selective illumination of a single plane of the
	fluorescence. The user does not directly observe the object, he or she rather sees a sample with a sheet of laser light while simultaneously recording emitted
	computer-generated image. fluorescence orthogonally to the illumination plane. A 3D image of the sample can
	then be generated with a temporal resolution ranging from seconds to several days,
	and at scales ranging from subcellular to whole organ.
	3. Melo-de-Pinna GF, Hernandes-Lopes J, Ogura AS, Santos LK, Silva DC, 2. 3D images at a magnification reaching 100,000x. It allows reconstructing
	Haevermans T (2016) Growth patterns and different arrangements of vascular developmental steps of an ontogenetic sequence. 2. It delivers detailed 3D phenotypical data and allows virtual dissections in all
	tissues in succulent leaves. International Journal of Plant Sciences 177(8): 643-3. Ren Y, Chang HL, Endress PK (2010) Floral development in Anemoneae directions.
	660. (Ranunculaceae). Botanical Journal of the Linnean Society 162(1): 77-100. 3. Staedler YM, Masson D, Schönenberger J (2013) Plant tissues in 3D via X-ray
	tomography: simple contrasting methods allow high resolution imaging. PloS one
	Replica molds (1970's) 8(9): e75295.

Transmission Electron Microscopy,

TEM (1930's)

1. A beam of electrons is transmitted through a sample (less than 100 nm thick) to form an image. The image is magnified and focused onto an imaging device.

2. 2D observations of fixed material or living material, using the GFP (Green Fluorescent Protein). It can reveal gene expression patterns. 3. Harrison CJ, Rezvani M, Langdale JA (2007) Growth from two transient apical initials in the meristem of Selaginella kraussiana. Development 134(5): 881-889.

Scanning Electron Microscopy,

SEM (1970's)

1. Destructive method, it requires many samples to obtain a consecutive series of states. Samples are critical point dried and coated with conductive materials (e.g. gold, platinum, graphite). In the SEM, an electron beam is emitted under vacuum and scans the surface of a sample. The most common SEM mode is detection of secondary electrons emitted by atoms excited by the electron beam. The number of secondary electrons that can be detected depends, among other factors, on specimen topography. By scanning the sample and collecting the secondary electrons that are emitted using a special detector, an image displaying the topography of the surface is created. SEM can achieve resolution better than 1 nm.

1. Indirect method of capturing the form of a structure. An impression mould of a structure is filled with epoxy. Once removed from the mould, an impression is retrieved, which is a positive reproduction of the initial structure. 2. 2. It provides fast imaging, and low phototoxicity, two key criteria for live imaging under physiological conditions. 3. Berthet B, Maizel A (2016) Light sheet microscopy and live imaging of plants. Journal of microscopy 263(2): 158-164. High Resolution X-ray Computed Tomography, HRXCT (2000's) 1. It is based on the property of low X-ray absorption of most plant tissues. It uses contrasting agents to reveal inner structures. The maximal resolution that can be obtained is 30 nm with synchrotron radiation. Compared with HRXCT, Micro Magnetic Resonance Imaging (microMRI) requires large and expensive magnets and cannot currently achieve resolution below 50-25 µm, and Optical Projection Tomography (OPT) requires that samples be transparent (treatment with caustic soda), a sample size of a few mm and allows a resolution of ca. 15-10 µm. HRXCT requires minimal sample preparation and is not destructive.

  Jabbour et al. 2015a), as well as with conceptual adjustments (Ronse De Craene 2010, Ronse De Craene et al. 2014) and technical advances (e.g. High Resolution X-ray Computed Tomography, Stuppy et al. 2003, Staedler et al. 2013; In vivo live imaging of meristem development, Jerominek et al. 2014; see Box 5; Davies et al. 2017).

  Model systems to study the evolution of nectar spurs. Ordinal-level phylogeny of flowering plants, with red branches indicating orders in which nectar spurs have evolved (Fernández-Mazuecos and Glover 2017). Three phylogenetically disparate genera in which evolution of nectar spurs is being investigated are shown. a, Aquilegia: species with different pollination syndromes display contrasting spur lengths, as exemplified by A. sibirica Lam. (c. 10 mm, bee-pollinated, top), A. formosa Fisch. ex DC. (ca. 20 mm, hummingbird-pollinated, centre) and A. chrysantha A.Gray (ca. 70 mm, hawkmoth-pollinated, bottom). b, Linaria Mill.: L. salzmannii (ca. 13 mm, left) and L. clementei Haenseler ex Boiss. (ca. 3 mm, right) are two closely related species with nearly identical floral morphology but contrasting spur lengths. c, Platanthera Rich.: intraspecific variation in spur length in P. bifolia (L.) Rich. (ca. 15-40 mm) correlates with the proboscis length of local pollinators.This work could be suitable for a MSc student or a PhD student, and will be done in collaboration with Thierry Deroin (MNHN), Sophie Nadot (Univ. Paris Sud),

	Catherine Damerval (CNRS Le Moulon), and potentially with Beverley Glover (Univ.
	Cambridge) and Eberhard Fischer (specialist of Impatiens L.; Univ. Koblenz,
	Germany).

1,

Garryales; 2, Untitled Group 1; 3, Gentianales; 4, Solanales; 5, Lamiales; 6, Aquifoliales; 7, Apiales; 8, Dipsacales; 9, Untitled Group 2; 10,

Asterales; 11, Ericales; 12, Cornales. From Jabbour et al. (2008)

.

  Stefan Dressler and I will visit in August 2018 are circled in white. The project gathers partners from the MNHN, the Senckenberg Museum (Frankfurt), and from the National Museums of Kenya. The field trip is funded by the NGS. Picture of D. macrocentrum Oliv. in its natural habitat on Mt. Elgon (Kenya; credit: David Gwynne-Evans). Picture of D. dasycaulon Fresen. in its natural habitat, Tala Hill Kasense,

	Rukwa (Tanzania; credit: Robert v. Blittersdorff; http://www.africanplants.senckenberg.de).
	The continuation of this project calls for more field trips, in priority in Ethiopia and
	Tanzania, but also in Cameroon (contact: Thomas Couvreur, IRD, Montpellier,
	https://sites.google.com/site/plantbiodiversityadaptation/members/permanents/thomas-
	couvreur).

ACKNOWLEDGEMENTS

I warmly thank Julien Bachelier, Vincent Debat, Pierre-Henri Gouyon, Paula Rudall, and Erik Smets for being part of the jury of my Habilitation defence. I thank Catherine Damerval, Thierry Deroin, and Emily Meineke for proofreading this thesis. I thank the members of the MADP team for their support and for making the daily work (experiments, supervisions of students, meetings, writing manuscripts and proposals) such a pleasure. I must as well say few words about five researchers, who influenced my way of conducting research and being a researcher. Most of them were former supervisors; many of them are current collaborators. Tatiana Giraud, Sophie Nadot, Catherine Damerval, Susanne Renner, and Thierry Deroin -thank you! I thank all my former and current students for regularly making me reappraise the way I supervise them. Special thanks are due to Felipe Espinosa, Lucas Batory, and Lucile Jourdain-Fiévet for showing such an enthusiasm for science and botany in particular. I thank Evelyne Ollivier, Sophie Nadot, and Manuela Lopez-Villavicencio for giving me the opportunity to co-supervise PhD students. I thank the colleagues from the MNHN, be they botanists, ornithologists, paleontologists, or entomologists, technicians or researchers, for all the fruitful and pleasant interactions. I heartily thank my very close collaborators Wei Wang, James Byng, and Stefan Dressler for putting their trust in me.

Many thanks to my friends Andreas and Marie, for their loyalty and support.

I thank Stephan for all the sacrifices he made and for his unconditional support. The Earth will always be "bleue comme une orange".

the most integrative possible. Results about mating systems, fitness, and pollination ecology were obtained and have now to be published. development, adapting the methodology of [START_REF] Fisler | Categorizing ideas about trees: A tree of trees[END_REF] who studied the evolution of the use of trees in systematics (Fig. 19). In the most optimistic (but unlikely) scenario, priority will be put on exploring, describing, and analysing the evolution of biodiversity, more funding opportunities will become available, the number of taxonomists will increase, and a new generation Plant morphology-anatomy is neither a minor discipline nor a discipline from the past.

On the contrary, the roots of plant morphology-anatomy are very deep. Many advances, especially in the imaging techniques, have been made in the last 50 years, and morphology-anatomy deserves a place among the -omics. Why could not we argue that we are conducting studies in the field of morphomics and anatomics, if this is a way to get more funding and visibility from other fields of science? Few disciplines allow a single researcher to conduct all the steps of the research (from material sampling to writing a manuscript through the experimental part) his-/herself.

Plant morphology is one of them. In this context, one should focus even more on collaborating with colleagues, locally and internationally, training students tirelessly, and integrating his/her research in the broad context of evolutionary biology.

VIII.b. CURATION AT P HERBARIUM

I am a curator for 12 orders (ca. 750,000 specimens) of flowering plants (e.g.