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PROGRAMME « RETOUR POST-DOCTORANTS »

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Project MEvo-DivA

DOCUMENT SCIENTIFIQUE (VERSION ANGLAISE)

Important Ce document, hors annexes, ne doit pas dépasser 30 pages, corps de texte en police de

taille 11. Ce point constitue un critère de recevabilité de la proposition de projet. Les propositions de projets ne satisfaisant pas aux critères de recevabilité ne seront pas

évaluées.

Nom et prénom du coordinateur / coordinator’s name

Thomas Claverie

Acronyme / Acronym MEvo-DivA

Titre de la proposition de projet en français

Modularité et évolution de la diversité chez les arthropodes

Proposal title in english Modularity and evolution of diversity in arthropods

Domaine scientifique/scientific domain Evolution and development – Macroevolution

Type de recherche / Type of research

Recherche Fondamentale / Basic Research Recherche Industrielle / Industrial Research Développement Expérimental / Experimental Development

Aide totale demandée / Grant requested

164 340 €

Durée de la proposition de projet / Proposal duration

36 mois

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1. PRESENTATION OF THE CANDIDATE.........................................................31.1. Short Biography..........................................................................................31.2. Full presentation.........................................................................................31.3. List of publications......................................................................................9

2. CONTEXT AND RELEVANCE TO THE CALL..................................................92.1. Relevance of the proposal.........................................................................102.2. Presentation of the laboratory..................................................................11

3. SCIENTIFIC AND TECHNICAL DESCRIPTION...............................................143.1. State of the art..........................................................................................143.2. Progress beyond the state of the art.........................................................16

4. SCIENTIFIC AND TECHNICAL OBJECTIVES / PROJECT ORGANIZATION.............224.1. Scientific Programme, Project structure....................................................224.2. Project management.................................................................................244.3. Description of the tasks............................................................................24

4.3.1 Task 1 254.3.2 Task 2 264.3.3 Task 3 274.3.4 Task 4 28

4.4. Tasks schedule, deliverables and milestones............................................295. DISSEMINATION AND EXPLOITATION OF RESULTS......................................316. SCIENTIFIC JUSTIFICATION FOR THE MOBILISATION OF THE RESOURCES.........32

6.1. Equipment................................................................................................326.2. Personnel costs.........................................................................................336.3. Subcontracting..........................................................................................336.4. Travel........................................................................................................346.5. Expenses for inward billing (Costs justified by internal procedures of

invoicing)..................................................................................................346.6. Other working costs..................................................................................34

7. ANNEXES.........................................................................................347.1. References................................................................................................347.2. Involvement of the candidate in other contracts.......................................347.3. Additional administrative document programme Retour Post-doctorants

2011.........................................................................................................35

Avant de soumettre ce document :

- Supprimer toutes les instructions en rouge (par exemple en faisant Format Styles Menu contextuel du style « Instructions » Sélectionner toutes les occurrences suppr.)

- Mettre la table des matières à jour (bouton droit sur la table des matières mettre à jour les champs Mettre à jour toute la table).

- Donner toutes les références bibliographiques en annexe 7.1.

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1. PRESENTATION OF THE CANDIDATE(8 pages maximum)

1.1. SHORT BIOGRAPHY(10 lignes maximum) Fournir une brève notice biographique en français.

I began my career with a solid foundation in marine biology and evolution culminating in a Masters degree. Subsequently, my PhD focused on identifying selective pressures acting on the claw morphology of a crustacean (Anomura). During this, I perfected novel techniques to quantify morphology (geometric morphometrics) and utilised a multidisciplinary approach involving population biology, behaviour and biomechanics to identify the selective pressures acting on claw morphology. Currently I am investigating the evolution of the raptorial appendage of mantis shrimps as part of my post-doctoral research. During this project, I have measured more than 70 species and did a series of biomechanical measurement to correlate morphology to function. I have identified developmental modularity in this appendage and have linked each developmental module to a function [15].

1.2. FULL PRESENTATIONFournir un CV très précis et une notice de titres et travaux synthétique ainsi qu’une copie du diplôme de docteur.Remplir le formulaire au paragraphe 7.3 intitulé « additional administrative document for programme PDOC 2011 ».EDUCATION

2008 Ph.D. in Biology: “Cheliped morphology, behaviour and selective pressures in the squat lobster Munida rugosa (Fabricius, 1775).” Advisor: Dr I. Philip Smith, University Marine Biological Station, Millport (United Kingdom)

2003 D.E.A. (Master’s degree) in Biodiversity of Fossils and Actual Ecosystems University of Lille (France)Research advisor: Professor Peter M. J. Herman

2002 Maîtrise (Bachelor’s degree + 1) in Biology of Populations and Ecosystems, option Marine BiologyEuropean Sea University Institute (IUEM) (Brest, France)

2001 Licence (Bachelor’s degree) in Biology of OrganismsUniversity of Brest (France)

2000 D.E.U.G. (Two Year Degree) in Life Sciences, Biology and Earth SciencesUniversity of Bordeaux (France)

PUBLICATIONS

In preparation:

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McHenry, M. J., Claverie, T. & Patek, S. N. (In prep) Gearing for high speed generates a slow strike in the raptorial appendage of the mantis shrimp

Claverie, T. & Patek, S. N. (In prep) Correlated evolution of a power-amplified structure: modularity in mantis shrimp.

Staaterman, E. R., Clark, C. W., Gallagher, A. J., deVries, M. S., Claverie, T., Patek, S. N. (In prep) Rumbling in the benthos: the acoustic ecology of the California mantis shrimp and the presence of anthropogenic noise

Published: Claverie, T., Chan, E. & Patek, S. N. (2011) Modularity and scaling in

fast movements: power amplification in mantis shrimps, Evolution 65, 443-461

Claverie, T. & Smith, I. P. (2010) Allometry and sexual dimorphism in the chela shape of the squat lobster, Munida rugosa. Aquatic Biology 8, 179-187.

Staaterman, E. R., Claverie, T. & Patek, S. N. (2010) Disentangling defense: the function of spiny lobster sounds. Behaviour 147, 235-258.

Zack, T. I., Claverie, T. & Patek, S. N. (2009) Elastic energy storage in the mantis shrimp’s fast predatory strike. Journal of Experimental Biology 212, 4002-4009.

Claverie, T. & Smith, I. P. (2009) Morphological maturity and allometric growth in the squat lobster Munida rugosa, Journal of the Marine Biological Association of the UK 89, 1189-1194.

Claverie, T. & Kamenos, N. (2008) Spawning aggregations and mass movements in subtidal Onchidoris bilamellata (Mollusca; Opisthobranchia), Journal of the Marine Biological Association of the UK 88, 157-159.

Claverie, T. & Smith, I. P. (2007) Functional significance of an unusual chela dimorphism in a marine decapod: specialisation as a weapon?, Proceedings of the Royal Society B 274, 3033-3038.

Claverie, T. & Smith, I. P. (2007) A comparison of the effect of three common tagging methods on the survival of the galatheid Munida rugosa (Fabricius, 1775), Fisheries Research 86, 285-288.

Bouma, T.J., van Duren, L.A., Temmerman, S., Claverie, T., Blanco-Garcia, A., Ysebaert, T., & Herman, P.M.J. (2007) Spatial flow and sedimentation patterns within patches of epibenthic structures: Combining field, flume and modelling experiments, Continental Shelf Research 27, 1020–1045.

GRANTS AND FELLOWSHIPS

Claverie, T. (2010) Grant to attend conference: 2010 annual meeting of the Society for Experimental Biology, (SEB conference grants)

Claverie, T. (2006) Grant to attend conference: 3rd European Conference on Behavioural Biology, (ASAB conference grants)

Claverie, T. (2004) Sheina Marshall PhD studentship, University of London (University Marine Biological Station, Millport)

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PRESENTATIONS AND PUBLISHED ABSTRACTS

Claverie, T. & Patek, S. N. (2010) Society for Experimental Biology, Prague, Czech Republic

Claverie, T. & Patek, S. N. (2010) Society for Integrative and Comparative Biology, Seattle, WA, U.S.A.

Claverie, T. (2009) Division of Vertebrate Morphology, Northeast Regional Meeting, Providence, MA, U.S.A.

Claverie, T. & Patek, S. N. (2009) Society for Integrative and Comparative Biology, Boston, MA, U.S.A.

Claverie, T. & Smith, I.P. (2007) 11th Congress of The European Society for Evolutionary Biology, Uppsala, Sweden

Claverie, T. & Smith, I.P. (2006) 3rd European conference on Behavioural Biology, Belfast, United Kingdom

Claverie, T. & Smith, I.P. (2006) 11th International Behavioural Ecology Congress, Tours, France

Claverie, T. & Smith, I.P. (2005) 6th International Crustacean Congress, Glasgow, United Kingdom

SEMINARS

2010 – ISEM, Montpellier, France; Univ Bourgogne, Dijon, France; Univ Bath, United Kingdom; OSEB, Paris, France.

2007 – SAMS, Oban, United Kingdom.

REVIEWER FOR

2010 – Journal of Experimental Zoology, Aquatic Biology, Journal of Applied Ichthyology.2009 – Journal of the Marine Biological Association of the United Kingdom, Fisheries Research.

TEACHING EXPERIENCE

2010 Guest lecture: Department of Biology, UMass AmherstImmune response

2004-2007 Teaching assistant: seven intensive Marine Biology courses held at the University Marine Biological Station in Millport, for two weeks each in March/April and August/September:

2004-2005 Teaching assistant: two one-month introductory Marine Biology courses for high school students.

STUDENTS MENTORING

Christina Mitsock, Biology Undergraduate Research Apprentice, UMass AmherstDanielle Cloutier, Laboratory Assistant, UMass AmherstJoie Yonamine, Laboratory Assistant, UMass AmherstQuan Tran, Laboratory Assistant, UMass, Amherst

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Elliot Chan, Undergraduate Research Apprentice, UC BerkeleyAlexander Stubbs, Undergraduate Research Apprentice, UC BerkeleyTravis Zack, Undergraduate Research Apprentice, UC Berkeley

SKILLS

Field work: - Sampling at sea from various research vessels (trawling, creeling, van Veen grabs, Niskin bottles)- Scuba diving work (specimen collection, tagging, density survey, distance measurement, coring, photography, sound recording, video recording)- Sampling and various measurements in intertidal environments

Laboratory work: - Museum measurements (National Museum of Natural History, Smithsonian Institution, Washington, DC, USA; Australian Museum of Natural History, Sydney, Australia)- Behavioural observations (design of arena for observation of agonistic interactions using video recording facilities, high speed video recording) - Force measurements of crustacean claws (design and conception of measuring apparatus with pressure sensor, use of piezoelectric impact force sensor) - Marine European faunal identification (crustaceans, molluscs, polychaetes, fish, zooplankton) - Flume measurement (current velocity within and around vegetal simulated structure)

Analytical work:Geometric morphometric (followed two Geometric morphometric courses), Phylogenetic comparative methods (followed the anthrotree workshop on comparative methods), Univariate and multivariate analysis, MySQL database, R programming, image analysis, capture re-capture analyses in Mark

UNIVERSITY SERVICE AND PROFESSIONAL SOCIETY MEMBERSHIPS

2010 Co-coordinator of the Behavior and Morphology discussion group at the University of Massachusetts, Amherst.

2005-2007 Research student representative on the Learning and Research Sub-Committees of the Management Committee at the University Marine Biological Station, Millport.

Association for the Study of Animal Behaviour, European Society for Evolutionary Biology, Society for Experimental Biology, Society for Integrative and Comparative Biology

ADDITIONAL INFORMATION

Language: French: native language, English: written, read, spoken, Spanish: read

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Academic qualifications: 2010: French qualification for “maître de conférence” (equivalent lecturer) university and museum in section 67 (population biology) and 68 (organism biology).

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1.3. LIST OF PUBLICATIONSFournir une liste de « publications choisies » du candidat destinée à éclairer le comité d’évaluation ; préciser le mode de sélection retenu pour établir cette liste ; les publications les plus significatives feront l’objet d’une référence complète avec le titre de l’article, son nombre de pages et éventuellement son facteur d’impact ; indiquer également le nombre total de publications dans les revues avec comité de lecture.List of significant publications ordered by impact factor. They are relevant for the project because they address modularity analyses and crustacean morphological quantification.

IF: 5.429 Claverie, T., Chan, E. & Patek, S. N. (2011) Modularity and scaling in fast movements: power amplification in mantis shrimps, Evolution 65, 443-461

Number of pages: 19

IF: 4.857 Claverie, T. & Smith, I. P. (2007) Functional significance of an unusual chela dimorphism in a marine decapod: specialisation as a weapon?, Proceedings of the Royal Society B 274, 3033-3038.

Number of pages: 6

IF: 1.380 Claverie, T. & Smith, I. P. (2010) Allometry and sexual dimorphism in the chela shape of the squat lobster, Munida rugosa. Aquatic Biology 8, 179-187.

Number of pages: 9

IF: 0.995 Claverie, T. & Smith, I. P. (2009) Morphological maturity and allometric growth in the squat lobster Munida rugosa, Journal of the Marine Biological Association of the UK 89, 1189-1194.

Number of pages: 6

Total number of peer reviewed articles: 9

2. CONTEXT AND RELEVANCE TO THE CALL (6 pages maximum) Présentation générale du problème qu’il est proposé de traiter dans le projet et du cadre de travail (recherche fondamentale, recherche industrielle ou développement expérimental). (0,5 page maximum)

What drives the evolution of biodiversity? The concentration of diversity in particular species-rich habitats is well known and comparatively well understood. By stark contrast, the concentration of biodiversity into particular branches of the tree of life is much less studied. Why do some groups contain tens of thousands of species, while their sister clades contain only a handful [30, 45]?

All species (with the exception of sibling species) differ morphologically. Hence, morphological diversity – or “disparity” [10, 53] – is a correlate of species diversity. The ability to evolve into novel morphologies is therefore integral to the ability to diversify. But what governs morphological plasticity in evolutionary terms? A strong candidate at the phenotypic level is modularity. A

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highly modular body is one composed of many units upon which selection appears to be able to operate independently: not only to different degrees but also in different directions. This contrasts with a highly integrated morphology, where change in one region necessitates simultaneous changes in others. Modular organisation in organisms have been hypothesised to enhance evolvability [57], however very few empirical studies have tested this hypothesis [24, 43] and even less have evaluated its role in the potential for clade diversification [18, 24, 62]. Therefore at a time where understanding the processes generating and influencing biodiversity is critical to human knowledge, the role of modularity in generating biodiversity can not be ignored. This fundamental research proposes to investigate the link between phenotypic modularity and morphological disparity within and across clades.

2.1. RELEVANCE OF THE PROPOSAL(2 pages maximum)Préciser :

Le positionnement du projet par rapport au contexte développé précédemment : vis-à-vis des projets et recherches concurrents, complémentaires ou antérieurs, des brevets et standards…

- Le positionnement du projet aux niveaux européen et international.The micro and macroevolutionary implication of the discoveries made in

developmental biology have just begun and this project will bring empirical evidence for a novel and provocative theory. The modular organization of living organisms have been hypothesized and commented for many years but it is only in the past 20 years with the advancement made in the field of evolution and development (Evo-Devo) that modularity was rigorously quantified and demonstrated [14, 32, 57-59]. Nowadays sufficient knowledge and techniques have been acquired to investigate the evolutionary consequences of the discoveries made in the field of Evo-Devo [6]. A provocative idea was that modularity enhances evolvability by reducing the pleiotropic effect of the genome on phenotypic expression [57]. This idea have been well discussed and tested theoretically but empirical studies are lacking [18, 24, 29, 31, 43, 54, 57-59, 62]. Therefore, one of the principal aims of the proposed project is to provide empirical information to test whether modularity and evolvability are related.

Among the many model organisms available to address the role of modularity in diversification few were used in previous studies; a limitation which will be considered in this project. All research being pursued to understand evolution of modularity or its consequence on evolvability is mainly made on mammal skull or mandibles [18, 24, 43, 47]. Furthermore, mammal skulls are extremely integrated (poorly modular) in the first place [28] and for this reason; it is possible that such model might be best suited to address the issue of integration on evolvability. The present study would provide a new model to approach similar questions. Arthropods (the model organisms of this study) are known to be highly modular and one of the most diverse organism present on the planet [11, 13, 45, 61]. Therefore the data that would arise from the present project would be beneficial to the scientific community that investigates similar questions. Finally this study would provide the morphometrics tools to promote arthropods as great models to understand the evolution of modularity.

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The theoretical framework surrounding the question of the proposed research is of wide international interest and teams around the world investigate the evolutionary consequences of discoveries made in developmental biology. The question that modularity might enhance evolvability was first proposed by teams in USA [57, 58]. This idea promoted many discussions on the international scene but remained mainly theoretic [29, 31, 54, 59]. Nowadays the interest on developmental mechanisms to understand patterns of biodiversity became one of the driving interests for many international teams [1, 10, 11, 18, 27]. It is only recently, with the advancement of knowledge and techniques that researchers from South America, Northern America and United Kingdom started to investigate empirical evidence for this theory [18, 24, 43, 62]. Therefore this project will address questions which largely interest the European and broader international scientific community.

2.2. PRESENTATION OF THE LABORATORYPrésenter le laboratoire d’accueil, ses axes et activités de recherche et les moyens qu’il met à disposition du candidat. (4 pages maximum)Welcoming laboratory: Biogeosciences (UMR CNRS / University of Burgundy)

The Biogeosciences Joint Research Unit is a research community currently comprising 80 members, including researchers from the CNRS (French National Centre for Scientific Research), faculty from the University of Burgundy and the EPHE (Ecole Pratique de Hautes Etudes), post-doctoral fellows, doctoral students, and technical staff. This multidisciplinary lab, combining Earth and Life Sciences, is comprised of three research groups. While each group of 20-25 members has its own research themes, research at the interface between all three groups is promoted and encouraged.

1. ECO/EVO (Ecology and Evolution), led by Thierry Rigaud. The main aim for this group is to understand how living organisms adapt to environmental changes, focusing more specifically on the biotic environment, chiefly through the study of cooperation and conflict between mating partners. Host-parasite relationships, immuno-ecology and mating systems are the main topics studied in this group.

2. FED (Form, Evolution and Diversity), led by Sophie Montuire. Fossil and modern biodiversity are key research themes, with a focus on the understanding of how biodiversity appears, diversifies, and is either maintained or lost, from the level of the individual up to the clade, at several spatiotemporal scales of analysis, using various parameters, such as morphology, taxonomy, and genetics.

3. SEDS (Systems, Environments and Dynamics of Sediments), led by Jean-François Deconinck. This group focuses on the analysis of sedimentary systems, with various axes: the transfer of matter in past and current systems; Jurassic/Cretaceous palaeoclimates; the formation and diagenesis of carbonate reservoirs; interactions between the stratigraphic record and the deformation of continental lithosphere.



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Such well-defined groups ensure a continuum of competence and research questions within the Biogeosciences lab. Collaboration between the ECO/EVO and FED groups focuses on the general theme of biodiversity and evolutionary mechanisms, while the FED and SEDS groups collaborate on sedimentary systems, their mode of operation and the information they provide (e.g. ancient climates).

Welcoming group: FED (Form, Evolution and Diversity)

The main research questions of the FED group concern biodiversity, its implementation, its maintenance and the diversification mechanisms considered from the level of the individual up to the clade. The group integrates various spatiotemporal scales of analysis, and uses several parameters to describe biodiversity (morphology, taxonomy and genetics). Three main axes are particularly considered: evolution and development, differentiation and phylogeny, and biodiversity dynamics.

1. Evolution and developmentThe Evolution and Development axis focuses on the study of

developmental processes at various taxonomical scales. Some projects at the macroevolutionary scale concentrate on the role of ontogenetic modifications on morphological expression and macroevolutionary dynamics. Other projects concentrate on the study of modularity and morphological integration at the clade scale, or at a finer scale. The third group of projects is at the microevolutionary scale, focusing on the study of mechanisms controlling development stability. The morphometric approach, shared by all these projects, has earned international recognition for the laboratory. The genetic facet can also be integrated through external collaboration. The EVO/DEVO approach is of particular interest for research questions at a macroevolutionary scale. The emergence of large-scale organisational plans, the study of developmental constraints, the structuring of morphological space or the use of the temporal dimension of development in the understanding of divergences between species (heterochronism), are themes which may now be re-examined in the light of new morphometric approaches. More recently EVO/DEVO has also endeavoured to answer questions about microevolution. The characterisation of phenotypic variation and the understanding of the processes that determine its origin and control are the focus of this research. Various projects centred on three major aspects of development are included under this heading: (1) the study of ontogenetic processes- Biomineralisation and ontogenesis(model organisms: bivalves, crustaceans)- Morphological space and ontogenetic disparities(model organisms: extant and fossil cephalopods), (2) the evolution of modularity and of morphological integration(model organisms: echinoderms, cephalopods, rodents) and (3) the study of developmental stability control(model organisms: sea urchins, ground beetles, fruit flies).

2. Differentiation and phylogenyPhylogenetic approaches, combined with ecological, geographical or

behavioural information, allow the general causes of differentiation to be studied. The task of obtaining well-resolved phylogenies that are operational



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therefore seems primordial. These diverse phylogenetic approaches are necessary to identify general tendencies and to consider all the processes leading to specific diversity among clades (the biodiversity dynamics axis).The validity of studies dealing with differentiation of populations, speciation and phylogenetic relationships is heavily dependent on the quality and quantity of morphological or genetic data. Defining a character or a number of informative characters is a prerequisite to the study of the evolutionary history of taxa, whether individuals within populations, subspecies or species. Estimating levels of differentiation (e.g. morphological, genetic, karyological) between individuals or populations remains a key problem. A general appreciation of the mechanisms that induce differentiation, or, on the contrary, create homogeneity in populations, is important for the understanding of how diversity (genetic or morphological) is structured.Two groups of projects are identified under this heading: (1) Differentiation and speciation mechanisms (model organisms: rodents, sea urchins) and (2) Phylogenetic reconstructions(model organisms: echinoderms, cephalopods).

3. Biodiversity DynamicsThe aim of this research axis is to describe and analyse biodiversity

dynamics on various spatiotemporal scales. More specifically, this will link the structuring of diversity (morphological, genetic, or specific) with palaeoenvironmental variations at local, regional and global scales. The contribution of the temporal dimension to the study of evolutionary processes will then allow the study of biodiversity fluctuations in periods of crises (extinctions, migrations) and reconquest. Two major themes are included under this heading: (1) Biomineralisation (model organisms: molluscs). The general aim will be to classify in families the proteins of calcifying matrixes, depending on their primary structure but also their function in mineralisation, and to propose a coherent model of carbonate biomineralisation. (2) Biodiversity over time (model organisms: rodents, molluscs, echinoderms).Projects will seek to test various models of community response to environmental change and will examine 1) collections of Jurassic and Cretaceous invertebrates, 2) biodiversity and global change through the example of Antarctic echinoids, and 3) the impact of climate change on biodiversity and also on morphological and genetic evolutionary rhythms in voles.

Facilities usable by applicant:Morphometric measurements and data processing:

Measuring microscope Nikon MM-60 (precision 1µm) Geometric computer Quadra-Chek 200 Metronics: 3D digitizer (x, y, z) Polhemus 3Space digitizer: 3D digitizer (x, y, z) Epson Perfection 1670 scanner: picture scanner 1 digital calliper with USB data output: distance measurements 7 digital callipers: distance measurements Winwedge software: transfers coordinates into excel Hyperspace 1.5 software: transfers coordinates into excel Optimas 6.5 software: coordinate acquisition, distance measurements

outline extraction from digital pictures. Matlab 6.5: programming software



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Aquarium lab facility: 25 m2 wet laboratory with controlled temperature and light condition

3. SCIENTIFIC AND TECHNICAL DESCRIPTION(8 pages maximum)

2.3. STATE OF THE ART

Décrire le contexte et les enjeux scientifiques dans lequel se situe le projet en présentant un état de l’art national et international dressant l’état des connaissances sur le sujet. Faire apparaître d’éventuels résultats préliminaires. Inclure les références bibliographiques nécessaires en annexe 7.1.

Can modularity – the division of an organism into structural, genetic or functional units – influence biodiversity? Modularity theoretically enhances the level of evolvability [10, 57, 58]. Evolvability is the capacity that living organisms have to produce novelty (new phenotypes) upon which natural selection can act [29]. If a species (or even a clade) has a great capacity to evolve, it will produce important variation in phenotypes and will be able to rapidly adapt to changing environment. Therefore highly evolvable species (or clade) should become more phenotypically variable than poorly evolvable ones [20]. By extension if modularity enhances evolvability, it should also enhance diversity of phenotypes or disparity [24, 62].

Modularity is a universal property of living organisms and should be considered as a hierarchical organisation of phenotypic trait that represent modules with various degrees of independence among them, rather than a segmentation of the organisms in truly independent structural blocks [34, 41, 60]. Many studies have highlighted the universal occurrence of modularity in taxa (e.g., mammals [63], arthropods [61], birds [39], bryozoans [12], molluscs [48]), but rigorous anatomical localisation of module was impaired by the fact that modules are never fully independent from each other [14, 33, 34, 41]. Therefore a consensus was made on defining a phenotypic module by a strong covariation among traits within the module but a weak covariation of traits among modules [14, 32]. Furthermore, a second level of complexity is the hierarchical nature of modularity. Large anatomical modules such as human arm can be subdivided in internal modules such as the hand and forearm which are more correlated together than they are with other modules within for example the leg [3, 4, 7, 41, 63]. However, little studies have examined this hierarchical degree of modularity, and it is not clear whether internal module of a limb are always more correlated together than they are with other trait elsewhere in the body [3, 7].

Past studies and present research focuses on determining the modular organisation of organisms by relating genes expression to phenotypic traits and their function, but little attention is given to the role of modularity in enhancing evolvability. So far studies have particularly investigated the relationship between genetic expression and developmental modularity in the context of genotype-phenotype mapping [4, 14, 35, 46, 57, 58], some studies have quantified the stability of modularity during evolution [22, 42, 43, 47, 50], and little have attempted to measure the relationship between modularity and evolvability [18, 24, 43, 62]. Modularity of mammals skulls appear to be stable



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in closely related clades and variable across distant ones [22, 42]. Furthermore, three studies on mammal skulls and one study on insect development suggest that modularity enhance evolvability [18, 24, 43, 62]. It was particularly shown that a strong integration of traits within one module was reducing the morphological disparity of that same module [24]. Finally, the consequence of hierarchical organisation of modules on evolvability is yet awaited to be addressed.

All the studies that investigated the role of phenotypic modularity on diversification have focused on trait covariation within mammal skulls [18, 24, 43], but other organisms such as arthropods are potentially great model for such studies. Mammal skulls, because of their important integration of traits [28], might represent limits to study the effect of modularity on diversification even if background knowledge on this system makes it highly appealing. Other model organisms could be arthropods. They are the most diverse group of metazoans, with approximately one million species described (all vertebrates represent just 60,000 species by comparison) and have an amazing array of morphological adaptations [1, 5, 13, 21, 61]. Furthermore, developmental studies demonstrated that arthropods are highly modular: different body segments belong to different developmental modules, and podomeres (leg segments) or groups of podomeres in the same limb also belong to different modules [9, 11, 37, 49, 61]. Finally, the enormous diversity of arthropods has frequently been attributed to their modular organisation but the theory remain to be tested [9, 61].

2.4. PROGRESS BEYOND THE STATE OF THE ART

Décrire les objectifs scientifiques/techniques du projet.

Présenter l’avancée scientifique attendue. Préciser l’originalité et le caractère ambitieux du projet.

Détailler les verrous scientifiques et techniques à lever par la réalisation du projet.

Décrire éventuellement le ou les produits finaux développés à l’issue du projet montrant le caractère innovant du projet.

Présenter les résultats escomptés en proposant si possible des critères de réussite et d’évaluation adaptés au type de projet, permettant d’évaluer les résultats en fin de projet.

Le cas échéant (programmes exigeant la pluridisciplinarité), démontrer l'articulation entre les disciplines scientifiques.Aim of the proposalEstablish the relationship between organism modularity and morphological disparity

To address such aim, it is first necessary to choose the model that will be used in this project. Arthropods, perhaps the most iconic segmented organisms, are great models to study the relationship between modularity and diversity because they were shown to be highly modular as well as extremely morphologically and speciose diverse [1, 5, 13, 21, 61]. Among arthropods, crustaceans are the most developmentally and morphologically variable promising to offer great diversity in patterns of modularity and morphological disparity [9, 21, 49, 61]. Finally among crustacean, thoracic segments of decapods (e.g. crab, shrimps and lobster) are the most relevant trait to study



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because their function is known, they are extremely variables and a large body of literature focused on their variation and the reason behind it [e.g. 5, 8, 16, 40, 49].

The aim of this project requires following a series of steps. The first step is to understand the modular structure of the thoracic segment of crustacean decapods and its stability through evolution. Secondly, modularity of decapods can be studied at two levels which are among body segments and within the legs (nested within body segments). It is therefore necessary to test the relationship between hierarchical arrangement of modularity and morphological disparity because such association could influence the interpretation of subsequent results. For example if body segments are highly integrated (high developmental covariation), the degree of modularity within legs will be comparable among body segments and overall disparity could depend entirely on the lower modularity degree (within legs). Conversely high modularity among body segments could lead to a variation in the degree of within leg modularity which could enhance the overall organism disparity. Finally, in the last step of this investigation, two levels of morphological variation need to be addressed. The first level, within population, is related to plasticity or population genetic structure. This level represents the raw variation among organisms that selection will act upon. It will produce a direct measurement of the effect of modularity in promoting novelty. The second level of variation, across species, is related to interspecific genetic differences which is the result of long lasting selection on a continuously changing pool of novelty. This level will inform the true consequences of modular organization for organism evolvability. Therefore the investigation will be organized in four points:

Defining the modular organization of the model organism Measure the stability of this modular organization during evolution Measure of the relationship between the modular structure (hierarchy of

modular dependence) and disparity Establish the connection between modularity and disparity in a population

and clade levelDetailed description of the objectives:

1. Defining the modular organization of the model organismQuestionWhat is the modular organization of the five thoracic segments holding the appendages in crustacean decapods?HypothesisTwo levels of modularity are awaited: each of the thoracic segments should display a great level of independence and each appendage should hold two modules a distal one and a proximal one.Objective and contribution to scientific advancement

The goal of this section will be to locate the different module that composes the thoracic region of crustacean decapods. Localization of modules and quantification of the degree of independence among them will be carried out using specially developed morphometric approaches [32]. Furthermore morphological plasticity of appendages will be investigated in carefully designed developmental experiments to detect the location of different modules. The results of the two approaches will subsequently be compared to validate the



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modular structure of the model organisms. Such measurement will allow connecting past work on genetic expression to phenotypic variation [9, 49]. Furthermore the results of this work will allow connecting previous work on appendages function to offer a better insight into the evolution of arthropod appendages.Justification

The modular organization of arthropod is likely to be determined by the location of few specific genes expression called hox and Dll genes [11]. Body segments are truly genetically modular and their identity is determined by the expression of homeotic genes, common to all arthropods and known as hox genes [49]. Any alteration of the expression of these genes could cause a change in the identity of particular segments as it was shown in Drosophila by, for example, growing wings on segments not supposed to have them [11]. The second sets of genes, the Dll genes, are also common in all arthropods and are responsible for the development of all the distal section of thoracic appendages. A non expression of these genes would result in the obliteration of the distal part of the appendages [9, 21, 49, 61]. Developmental organization of segment and appendages is generally more complex, but the location of developmental module is likely to be identified by the expression of these two set of genes. Therefore, I hypothesize that each body segment is an independent (or semi-independent) module as well as the proximal vs distal region of each appendages. Furthermore the degree of modularity within each leg shall be nested within the modularity of each segment.

Phenotypic plasticity will be used to locate modules. One of the mechanisms that produce phenotypic plasticity is the variation in the tissue development of some particular trait during ontogeny [60]. Localizing anatomical variation due to plasticity can be used to identify unity of developmental tissues or modules. The morphology of crustacean appendages is extremely plastic [25, 56]. Furthermore, the function of the different appendages in many species of crustacean decapods was well studied and determined [e.g. 17, 36, 40]. Therefore it is possible to associate a function with each of the modules defined previously and alter that function during development to control plastic variation. Variation can subsequently be quantified and particular modules of interest can be located. For example the morphology of crab claw was shown to be altered by diet manipulation or claw immobilization during development [56]. Therefore alteration of particular appendage function will be use to verify the location of particular modules.

2. Measure the stability of this modular organization during evolutionQuestionHow stable is the modular organization in crustacean decapods?HypothesisThe existence and location of the previously defined modules should be stable across crustacean decapods; however the degree of independence among these modules should be variable across clades.Objective and contribution to scientific advancement

The goal of this section is to quantify the stability (or instability) of the modules location and their relative degree of independence across decapods. Morphometric methods will be used to quantify modularity within at least five



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species chosen to represent all decapods. Subsequently, published phylogenies will be used to map and reconstruct these variable to understand the evolutionary history of modularity in decapods. This study will represent the first attempt to reconstruct the evolution of modularity within a clade and will bring fundamental information on the evolution of modularity which is not yet understood [59]. Justification

Evidence from previous work suggests that the modular organization of anatomical structures is stable within clade, but the degree of connection among these modules vary. The organization of modularity in mammal skulls was shown to be relatively stable in closely related clade (marsupial and placental) but vary to some degree for distant ones (monotremes such as platypus) [23]. However the level of integration among and within these modules was more variables [23, 24, 50]. Hox and Dll genes expression determine the modular development of decapods and were showed to occur in all arthropods [49, 61]. Only some crustacean mandibles appendages (which are not part of the thoracic segments) were shown not to express Dll genes [49, 61]. Therefore, the location of the different module as defined previously should remain relatively stable across decapods but the degree by which these modules are connected might be relatively variable.

3. Measure of the relationship between the modular structure (hierarchy of modular dependence) and disparity

QuestionIs the degree of modularity at one hierarchical level related to the degree of modularity at a lower hierarchical level and how is it related to morphological disparity?HypothesisThe degree of independence among the body segments modules is related to the variation of the degree of independence between the leg modules. Also the overall disparity of organisms is related to the degree of independence among modules from the two hierarchical levels.Objective and contribution to scientific advancement

The goal of this section is to quantify how the hierarchical modular organization can influence morphological disparity. In this section detailed disparity measurements for each module will be done using Foote [19] method and will be compared across modules. However, disparity measurements are sensible to the number of measured variables used to quantify morphology [19]. Decapod morphologies are extremely variable and in order to compare disparity among modules, novel methods will be developed to quantify every appendages morphology with the same number of variables. These measurements will be performed in several species that differs in their degree of modularity. This investigation will be the first attempt to show how hierarchical modularity in organisms can affect their morphology. This study will connect the variation in hierarchical level of modularity to the variation in morphological disparity. The results will inform if the same degree of disparity might be reached by controlling the independence at different level of modularity (body segment vs within appendages). Therefore, this study could bring new insights in the understanding of convergent evolution and in the concept of “many-to-one



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mapping” showing how different pathway can be used to reach the same results [2, 26]. Justification

Hierarchical modularity was found in mice mandible and butterfly wing and highlight an underlying complex organization. Mice mandible [4] and eyespot on butterfly wings [3, 7] were showed to have a hierarchical modular structure mainly due to variation in the timing and location of genes expression during tissues development. In arthropods the types of appendage that will be developed on a particular segment depend on homeotic genes such as hox genes. Once the identity of the segment is determined, the expression of Dll genes controls the development of the appendage distal region and therefore determines its morphology [11]. Although development of arthropod limbs is more complicated than this, the hierarchical interaction in these two type of genes expression allow us to hypothesize a hierarchical structure in the degree of independence among modules.

How modularity is organized could have an important effect on morphological disparity and its interpretation. Morphological disparity is a measure of the diversity in phenotypes and directly relate to the amount of novelty present in a species or a clade [19, 53]. The abundance of novel phenotype in a species can be influenced by various parameters but is originally controlled by the capacity of the species to evolve. Modularity was hypothesized to enhance evolvability [57]; however the influence of the hierarchical organization of modularity on evolvability never got addressed. If the overall species disparity is directly linked to the degree of modularity, high appendage modularity and low body segment modularity could give the same level of disparity than low appendage modularity and high body segment modularity. Therefore in this study it was important to consider the relative effect of both level of modularity on disparity for subsequent interpretation of results.

4. Establish the connection between modularity and disparity at a population and clade level

QuestionHow does the degree of modularity relate to morphological disparity at a population level (variation due to genetic diversity and phenotypic plasticity) and at a clade level (variation due to genetic evolution)?HypothesisThe degree of modularity in decapods is directly related to the degree of morphological disparity within a population and within a clade; however the degree of correlation between modularity and disparity in the population is higher than within a clade.Objective and contribution to scientific advancement

The objective of this section is to measure the relationship that exists between the degree of modularity and the level of morphological disparity. A second objective is to measure how this relationship varies within a population and within a clade. This study will require a quantification of the morphological variation within the population of several species and also a quantification of the morphology of a representative selection of species for several clades (related to species considered for populations). From these measurements, the degree of modularity and the level of morphological disparity will be quantified. The



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results from this section will show whether a link exist between modularity and evolvability which has been well established theoretically but lack of any data. Furthermore results from the relationship between modularity and evolvability at a population level and at a clade level will highlight if selection might override modularity in controlling morphological disparity [24]. Finally results from such a study have been requested in many publications but remain poorly addressed [24, 57, 59].Justification

The study of the relationship between modularity and disparity at a population level and at a clade level are both relevant to measure short and long term effect of modularity on evolvability. In populations, phenotypic variations are mainly due to genetic differences and plasticity; also selection of particular phenotype might remain minimal [60]. Interspecific variations are likely to be dominated by genetic differences upon which various selections might have occurred for thousand of years. If modularity enhances evolvability, at the population level, it will increase novelty and therefore enhance disparity [57]. At an interspecific level, modularity will have continuously enhanced novelty giving greater pool of variation on which selection acted upon. This might result in the enhancement of clade diversity and disparity [57, 62]. However depending on the nature of selections (directional, stabilizing or disruptive) the relationship between modularity and disparity might become larger or smaller [24].

SummaryOverall this project will allow the evaluation of the relationship that might

exist between organism modularity and evolvability, a provocative suggestion well established theoretically but poorly tested empirically. This project proposes to use morphological disparity as a proxy for evolvability and will use morphometric method to quantify modularity. Furthermore forced plastic variation during organism development will be use to confirm results from morphometric assessments. Difficulty related to the complexity of developmental organization will be addressed by quantifying of modularity at different hierachical levels [3, 4, 7]. Finally the effect of modularity on evolvability will be tested at two levels: the short term production of variation within populations and the long term enhancement of diversity across species within clade.

4. SCIENTIFIC AND TECHNICAL OBJECTIVES / PROJECT ORGANIZATION

(8 pages maximum)

2.5. SCIENTIFIC PROGRAMME, PROJECT STRUCTURE (2 pages maximum)Présentez le programme scientifique et justifiez la décomposition en tâches du programme de travail en cohérence avec les objectifs poursuivis. Utilisez un diagramme pour présenter les liens entre les différentes tâches (organigramme technique)



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Les tâches représentent les grandes phases du projet. Elles sont en nombre limité.

N'oubliez pas les activités et actions correspondant à la dissémination et à la valorisation.The Scientific program will be organized in four tasks:

Task1: Development of a novel method to quantify decapods morphology Task2: Developmental Alteration of particular modules in one species of

decapod Task3: Measure specimen morphology from different population of

decapods Task4: Measure the morphology of species within several clades

Each of these tasks will be use to address several objectives.

Fig. 1: Organigram representing the connections among tasks and objectives and the usage of tasks to address the different objectives.

1. Task1: Development of a novel method to quantify decapods morphology.

This project requires quantifying the morphology of each leg elements (podomere) of decapods for two different purposes. The first purpose is to quantify the morphological variation of each segment with enough precision for locating modules and determining their degree of independence among adjacent ones. Although this first purpose require precise measurements, no particularly novel method is required for such quantification since coordinates of point or various linear measurements can be used. The second purpose is to quantify morphological disparity among modules. The quantification of morphological disparity for a species is equivalent to a measure of the volume, in a morphological space (space that represents every possible morphologies), that this species occupy [19]. However the structure of the morphological space depends on the number of variables used to measure a particular structure. For example the morphological space required to represent the shape of a three dimensional object with orthogonal faces such as a cube will have three dimensions (length, width and height). However the morphological space required for representing all possible parallepiped require one more dimension to characterize the angle of one of the lateral faces relative to the base of the object. The shape of the podomere of decapods legs can be extremely variable going from a cylinder shape to a nail shape or claw shape. Therefore a novel method is necessary to quantify accurately their shapes in a manner that they all



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belong to the same morphological space (same number of variables) so comparable values of disparity can subsequently be obtained.

This novel morphological quantification will particularly be needed to fulfill the third and fourth objectives which require quantifying disparity within population and within clades of decapods (c.f. Fig. 1).

2. Task 2: Developmental Alteration of particular modules in one species of decapod

The different modules within the thoracic segments of the decapods measured in this project will be located using morphometric methods that rely on phenotypic variations. Therefore in order to validate the presence, the location and the independence of the different modules, some developmental measures are necessary. In this task I will use phenotypic plasticity to control morphological variation under various developmental constraints. Subsequently I will quantify the resultant morphological variation to identify which anatomical regions were affected.

The set of experiments proposed in this task will be particularly needed to fulfill the first objective of this proposal which aim at defining the modular organization of the thoracic region of the crustacean decapods (Fig. 1).

3. Task 3: Measure specimen morphology from different populations of decapods

Quantifying the morphological variation of the thoracic appendages of the specimen within a population will be performed for at least five different species. This task will require the collection of specimen directly on the field in quantity sufficiently large to estimate the population variability. Furthermore these measurements will be used to quantify modularity (location of module and degree of independence among them) and to quantify disparity.

This task will be fundamental to accomplish all the objectives of this project.

4. Task 4: Measure the morphology of species within several cladesThe same clades represented by the species measured in task 3 will be

selected for this task. The morphology of numerous species representing each clade will be measured from preserved specimen in museums. The morphology of the species from at least three clades (and up to ten) with different degree of modularity (established from task 3) will be measured.

This task will be fundamental to accomplish the second part of the fourth objective which is one of the major questions of this proposal (Fig. 1).

2.6. PROJECT MANAGEMENT (2 pages maximum)Préciser les aspects organisationnels du projet.

The project will be organized in three main data collection activities described in task 2, 3 and 4. Two of these activities require sampling specimen of crustacean in their natural environment. The marine biological station of Roscoff was selected for these activities because the fauna diversity in Roscoff is



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one of the greatest available nationally, the field sites are easy to access and the station possesses all the facility required for such project (boats and wet labs).

The first activity, the controlled development of crustacean (Task 2), will require animal collection and hiring a technician for assistance with the project. Specimens will be collected on the shore and brought back to the biogeoscience laboratory aquarium facilities in coolers via roadway. Thereafter, they will be grown in this facility and regularly fed with prey collected from the same shore brought back and frozen. One person will be recruited to assist with daily maintenance of specimen, and conduct the project. Job will be advertised by standard scientific advertisement and interviews will be organized.

The second activity, the quantification of morphology within population (Task 3), will require animal collection via Roscoff marine station facilities (boats) and from nearby sea shore. Once collected, animals will be measured in the station wet laboratory using a portable 3D digitizer. It is anticipated that a total of four month at the marine station in Roscoff will be necessary to sample and measure specimen required for this project.

The third activity, the interspecific quantification of morphology (Task 4), will require trips to the national museum of natural history collections in Paris. Specimen will be measured on site using the portable 3D digitizer and it is expected that this activity will also require up to four month at the museum in Paris.

2.7. DESCRIPTION OF THE TASKS(Idéalement 1 ou 2 pages par tâche)Pour chaque tâche, décrire :

- les objectifs de la tâche et éventuels indicateurs de succès,- le responsable de la tâche, - le programme détaillé des travaux par tâche,- les livrables de la tâche,- la description des méthodes et des choix techniques et de la manière dont les solutions seront

apportées,- les risques de la tâche et les solutions de repli envisagées.

4.3.1 TASK 1Development of a novel method to quantify decapods morphology

Objectives: The objective of this task is to develop a method to quantify every

podomere morphology using the same number of variables.Person responsible of the task:

Thomas ClaverieDetails:

This task will require obtaining three dimensional (3D) coordinates of points representing the surface of different podomeres and develop programming codes to fit a quadratic polynomial surface on these coordinates. Codes will be developed in the statistical software R [51].Deliverables:

The deliverables of this task will be a program or a series of codes which will be able to transform a set of 3D coordinates into multivariate variables usable to quantify disparity.Description of the methods:



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The series of articulating podomeres in a decapod leg can be modelled as a series of cylinders, elliptic cylinders, truncated ellipsoid and cones with articulation hinges between them (Fig. 2). The surfaces of these solids can all be modelled in three dimensions using the same quadratic equation with different parameters substituted:

Where a, b, c, f, g, h, p, q and d are parameters that can be adjusted to model either an ellipsoid (d = -1 and f, g, h, p, q and r = 0), an elliptic cylinder (the same as the ellipsoid but with c = 0) or a cone (f, g, h, p, q, r and d = 0 and c < 0). Therefore different morphologies can be measured with the same number of variables, a task hardly achievable with more traditional methods. Similar approaches have been used in iconic morphospace (space defined by morphological variables) studies, such as those modelling gastropod shells using logarithmic spirals [52].

Using reverse engineering procedures, coordinates of points on each of these segments will be measured with a 3D digitizer (immersion microscribe MX, Fig. 2). Quadratic surfaces will be subsequently fitted to these coordinates using a least squares procedures implemented by a custom made program (in R). As a result, the morphology of each of these podomeres will be quantified by the quadratic parameters. Furthermore, other variables will be measured to quantify the relative positions of hinges and features mounted on the podomeres such as the fixed fingers of claws (Fig. 2). Such variables could be the angle (α) subtended by the proximal and distal set of hinges, and the ratios H and C that represent the location of a fixed finger in the claw (Fig. 2). In order to separate size from shape, measurements will be standardised to the overall size of each specimen. Finally, grouping the variables from different podomere will be used to quantify segment morphology or overall anatomy.

Fig. 2: 3D coordinates were measured on medial leg podomere (top), on claws (middle), and on tips of the legs (lower) using a 3D digitizer (microscribe MX, white point on picture represent locations of measured coordinates). Trial calculations were used to estimate quadratic parameters from the 3D coordinates (X, Y and Z represent ratios from the quadratic parameters that



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represent respectively morphological changes in the x, y and z directions). The angle (α) made between the proximal and distal set of hinges and the ratios H and C giving the location of the fixed finger on claws were also calculated. The error term is the standard error calculated from three independent measurements. Risks and alternative solutions:

The risk associated with this task is that the fit of quadratic surfaces on complicated podomeres morphologies might lead to low level of accuracy. If such event occurred and would be such that it would alter disparity measurements, linear measurements such as length, width and height of the podomeres (which were used in numerous studies) will be calculated from 3D coordinates. Such measurements will give a less precise representation of podomeres shape but could be sufficient to measure some values of disparity.

4.3.2 TASK 2Developmental alteration of particular modules in one species of decapod

Objectives: The objective of this task is to measure the morphological variations due

to phenotypic plasticity in the development of decapods appendages under functional constraints which target particular modules.Person responsible of the task: Hired technicianDetails:

Specimen from one commonly occurring species of decapod (Carcinus maenas) will be raised from an early juvenile stage to a young adult size for few months under particular functional constraints targeting each time one particular module. Movement of legs, hardness of prey items and locomotors stimulation will be modified among treatments to promote radical changes in appendages development due to plasticity. Such perturbations have previously been measured to significantly affect morphology [25, 56]. Post experimental measurements will indicate which region of the appendage gets altered highlighting the location of the developmental modules. These results will subsequently be compared with the morphometric investigation of modularity on the natural population of Carcinus maenas (done in task 3). Deliverables:

This task will produce data that will indicate the location of the various developmental modules present among the appendages of one species of decapod. The final delivered information will be the assignment of each podomere to a particular module based on their morphological differences among treatments. Description of the methods:

Carcinus maenas uses its first pair of appendages (the claw) for feeding and the four other appendages to walk or climb. Furthermore the distal region of the appendage is supposedly a different module than the proximal region. Therefore, two type of functional alteration will be performed to alter either the distal or proximal module. The first type of disturbance will be to glue the movable finger of the claw (dactylus) to the non movable one (pollex) in a way the claw is not anymore functional. Animals will be fed hard shelled prey



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(mussels: Mytilus edulis) and prey shell will be broken up for glued specimen. This manipulation was previously showed to affect significantly the morphology of the claw [56]. The second alteration will concern walking legs. The tip of the leg is heavily used for griping the substrate while the proximal region of the leg to lift the animal body. Therefore the proximal part of the leg will be hold in one position by being glued to a rigid stick also glued to the body. This manipulation would alter the mobility of the proximal part of the leg therefore altering its function while allowing de distal part of the leg to grip the substrate freely. Furthermore, animal surrounding will be set up to enhance climbing and walking activity (disposition of rocks). Finally crustacean molt as they grow, therefore after each molts, functionally altered appendages will be glued again.

Five parallel experiments (one experiment per appendages) will run with each having 10 specimens with one pair of functionally altered appendage (treatment) and 10 specimens without any functionally altered appendages (control). A paired comparison of the morphology of each appendages podomere will subsequently be performed between treatments and controls groups to estimate the location of the functionally altered modules.Risks and alternative solutions:

The risks associated with this task are that the specimen morphology does not respond to appendage alteration (although such modifications were previously observed [56]). Alternative solution would be to repeat experiment with a different species therefore prior to launch the experiment; pilot study with small sample size will be performed using different species.

4.3.3 TASK 3Measure specimen morphology from different populations of decapods

Objectives: The objective of this task is to quantify the appendage morphology of

different population of decapods, characterize the modularity and its stability and quantify the disparity for each species.Person responsible of the task:


The modular organization and disparity of at least five species of decapods (and up to 10) will be examined. Approximately 50 specimens from a unique population for each species will be directly collected on the field. Previous studies showed that approximately 50 specimens per species are necessary to estimate the degree of modularity [15, 64]. Subsequently each individual will be measured using the method described in task 1: 3D coordinate for each podomeres will be used to quantify modularity and quadratic equation parameter will be used to quantify disparity. Furthermore, published phylogeny will be used along with likelihood based methods to reconstruct the evolutionary history of modularity [44, 55].Deliverables:

This task will permit the acquisition of morphometric data (3D coordinates and quadratic parameters for each podomere) usable to quantify the shape variability within populations for at least five species. Furthermore, location of the different module and the degree of independence among them will be quantified along with morphological disparity. The deliverable data will



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be the modules location (Podomere will be ascribed a value indicating their belonging to particular modules), the degree of modular independence (matrix of the degree of covariation among modules per species), and the disparity of each species (disparity values calculated for each module and for the overall species). Finally the stability of modularity in decapods will be represented by the reconstructed evolution of these variables on a phylogeny.Description of the methods:

Specimen will be collected partly from the rocky shore around the marine biological station of Roscoff and partly from the benthic area using the marine station facility (research vessel). Species such as Carcinus maenas, Maja squinado, Necora puber, Cancer pagurus and Xantho incisus are abundant species in this environment and will be sampled without many difficulties. The specimen will be brought back in the lab (in the Roscoff marine station) and 3D coordinates for each podomere will be recorded in duplicate using a 3D digitizer. These coordinates will subsequently be used to quantify modularity using previously developed geometric morphometric methods [32, 33, 38, 41]. Disparity will be quantified using Foote [19] methods based on the quadratic function parameters calculated using method describe in task 1. Finally parameters defining modularity will be mapped on a phylogeny [44] and evolutionary history of modularity will be reconstructed using likelihood based methods [55]Risks and alternative solutions:

The risk encountered in this task will be difficulty to collect large number of specimens from the benthic environment due to weather condition while on field trips. As a replacement, numerous other species are available on the sea shore but are less abundant, much smaller (which would complicate greatly the measurements) and less interesting with respect to their disparity. Alternatively specimen could be acquired from roughly the same geographic area from specimen supply service from the Roscoff marine station or commercial fish market but there is a risk that specimen diverge significantly in their genetic diversity.

4.3.4 TASK 4Measure the morphology of species within several clades

Objectives: The objective of this task is to quantify the appendage morphology of

different species of decapods belonging to the same clade as the species previously investigated, quantify modularity across species for each clade and quantify the disparity for each clade.Person responsible of the task:


The morphology of at least 20 species for at least three clade representing species considered in task 3 (and showing evidence of differences in the degree of modularity) will be measured from museum collections. Clades selected will represent individual families (used to regroup closely related species) and I anticipate that the Majidae, Xanthidae and Portunidae will be interesting clades to consider due to their variety in morphological variation. Morphological



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measurement will be used to quantify disparity and interspecific modularity (also called evolutionary modularity [32, 47]) for each clade.Deliverables:

This task will provide similar morphological data than previous task but from a larger array of species. This task will also provide data reflecting modularity quantification and disparity measurements as previously described (i.e. task 3) for at least three and up to 10 families of decapods. Description of the methods:

Specimen will be measured from the collection of the museum of natural history in Paris. This museum was selected due to the large size of its collection offering sufficient numbers of species to quantify the disparity of the three proposed families. Two to three specimens will be measured for each species. 3D coordinates will be used to quantify the modularity across species as it was done in various studies of mammal skulls and mandible [18, 22, 47, 50]. As in task 3, 3D coordinates will be used to obtain quadratic equation parameters for each podomere of the measured specimen. Subsequently these parameters will be used to quantification family disparity using Foote [19] method. Finally modularity parameters will be compared to disparity values in order to establish the relationship between modularity and disparity.Risks and alternative solutions:

The risk associated with this task is that numerous preserved specimens might be damaged leaving too little species available in a considered family. Alternatively other family could be considered or a visit to other museum such as the museum of natural history in London could help complement sample within a family.

2.8. TASKS SCHEDULE, DELIVERABLES AND MILESTONES(3 pages maximum)Présenter sous forme graphique un échéancier des différentes tâches et leurs dépendances (diagramme de Gantt par exemple).

Présenter un tableau synthétique de l'ensemble des livrables du projet (numéro de tâche, date, intitulé, responsable).

Préciser de façon synthétique les jalons scientifiques et/ou techniques, les principaux points de rendez-vous, les points bloquants ou aléas qui risquent de remettre en cause l'aboutissement du projet ainsi que les réunions de projet prévues.Timeline of investigation in month from the start date anticipated in September 2011

Activity\Month

1-4 5-8 9-12 13-16

17-20

21-24

25-28

29-32

33-36

Task 1Task 2 PilotTask 3Task 4Data analysisDissemination



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Summary of the deliverables

Deliverables Task Date due

Responsible

Program to quantify quadratic parameters Task 1 January 2012

Thomas Claverie

Anatomical location of modules from altered development

Task 2 May 2013

Recruited technician

Morphometric data from at least one decapod population and location of modules

Task 3 June 2012

Thomas Claverie

Morphometric data from several population of decapods and quantified modularity/disparity

Task 3 September 2013

Thomas Claverie

Morphometric data on at least 3 clades of decapods

Task 4 December 2013

Thomas Claverie

Interspecific modularity quantification and disparity values calculated

Task 4 February 2013

Thomas Claverie

Landmark dates

Event Date Final decision and plan on how disparity will be quantified and

evaluation of the progress on growth. Recruit technician to perform task 2

January 2012

First year evaluation: evaluation of the success for the growth experiment and decision on the final number and identity of species to samples based on measurement experience, data variability and modularity variations

September 2012

Final results of growth experiment due and end of the technician employment

May 2013

Second year evaluation: data collection finished or closed to be finish, data analysis already well advanced

September 2013

End of the project: final evaluation on the productivity based on publication and data generated. Proposition and outline of future project required

September 2014

5. DISSEMINATION AND EXPLOITATION OF RESULTS (1 à 2 pages)Présenter les stratégies de valorisation des résultats :

- la communication scientifique;- la communication auprès du grand public;- la valorisation des résultats attendus;- les retombées scientifiques, techniques, industrielles, économiques …- autres retombées (normalisation, information des pouvoirs publics, ...)- les échéances et la nature des retombées technico- économiques attendues- l’incidence éventuelle sur l’emploi, la création d’activités nouvelles, …

Présenter les grandes lignes des modes de protection et d’exploitation des résultats



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Results from this project will be disseminated to the science community via conferences and scientific publication but also to the general public via the constitution of website and blogs and via outreach in public lectures. The main beneficiaries of this project are anticipated to be the academic community working on developmental biology, evolutionary biology, ecology, comparative anatomy, functional morphology, morphometric and carcinology. Therefore these potential beneficiaries will be targeted in my networking efforts.

Effort to disseminate results from this project and network with interested colleague will be done via seminar and conference presentation. I will actively pursue opportunities to present my work in seminars at French and overseas institutions. I will also present results at several high profile international conferences in order to reach as wide as possible an audience. These will include the 14th congress of the European Society for Evolutionary Biology in 2013. By this time, I will have finished the measurements for quantifying the modularity organisation within populations and its relationship to disparity, and will be able to present these results to a broad audience of evolutionary biologists. In 2014, I will attend the World Conference on Marine Biodiversity which offers an ideal platform to reach systematists, carcinologists, conservationists and policy makers and I will present the results of this project. Finally each year I will present the advancement of my research at the annual Evolutionary Biology Meeting of Marseilles.

The research will be published in a minimum of three high impact journals, in addition to those focussed on more restricted questions. Furthermore these articles will have an open access to promote the dissemination to community without or limited literature access. The first one would present results on the evolution of modularity in decapods, and will propose hypotheses for what drove differences in morphological diversity among clades. This will be targeted to PLoS Biology, Evolution or the Journal of Evolutionary Biology. The second article will present analyses of the morphological disparity among clades and will be submitted to Systematic Biology, Evolution or Proceedings of the Royal Society of London B. The last core article will synthesise data from the whole project by comparing levels of modularity and amounts of morphological disparity. Ultimately, this will allow us to test whether modularity promotes evolvability. This could be attractive to Science, Nature or Proceedings of the National Academy of Sciences of the United States of America.

In order to disseminate the knowledge gained in this project to the broadest public possible, I will make good usage of internet and public outreach initiatives. First I will deposit published data on online depository such as British Oceanographic Data Centre. I will also produce a dedicated set of webpages promoting this research, tailored to specialists (repositories for data and software/codes targeting open access freeware such as R to promote exchanges), evolutionary biologists (abstracts, summaries, pdfs and press releases), interested lay people (accessible summaries of the evolutionary findings). The latter will promote the public understanding of science in a field that is frequently misrepresented and misunderstood. I have excellent experience of disseminating information in this way (http://www.ocf.berkeley.edu/~claverie). Furthermore, I will also produce a blog, which I previously used to good effect on a previous project


http://www.ocf.berkeley.edu/~claverie


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Project MEvo-DivA


(http://stomatopodhunters.blogspot.com). This is much less formal, and will encourage interaction and public engagement. Furthermore, I will use the opportunity of working in the national museum of natural history in Paris to actively propose my participation in the outreach events of the museum such as in “les rendez vous du museum”. I will also propose to participate to the outreach events from the University of Burgundy and organized by the Biogeoscience laboratory.

3. SCIENTIFIC JUSTIFICATION FOR THE MOBILISATION OF THE RESOURCES

On présentera ici la justification scientifique et technique des moyens demandés dans le document de soumission A.

3.1. EQUIPMENTPréciser la nature des équipements* et justifier le choix des équipements

Si nécessaire, préciser la part de financement demandé sur le projet et si les achats envisagés doivent être complétés par d’autres sources de financement. Si tel est le cas, indiquer le montant et l’origine de ces financements complémentaires.

*Un devis sera demandé si le projet est retenu pour financement.Five types of equipments will be necessary in this project: a 3D digitizer with an associated laptop, two computers for daily usage, aquariums and a freezer.

3D digitizer: the most appropriate equipment is the Immersion Microscribe MX with accessories. Although the host laboratory possesses 3D digitizing apparatus, none of them are portable (which is an obligatory condition) or appropriate to perform the necessary measurements. Accessories such as calibrating tolls, carrying box and triggering paddle are necessary Cost: 10 106 €

Laptop: a sturdy laptop is necessary to register 3D coordinates from the digitizer. This laptop has to be robust enough for extended work in wet lab environment and museum. A suitable model is a Lenovo thinkpad T series.Cost: 1 210 €

Computers: two desktop computers are necessary for the daily usage of the postdoctorant and the technician.Cost: 1000 €

Aquariums: The biogeoscience laboratory have aquarium facilities, but particular aquariums specific for this project are necessary (Task 2). Five 200 L tanks equipped to receive salt water specimens and sufficient dehydrated salt to renew water for a year.Cost: 3 580 €

Freezer: one upright 250 L freezer to keep frozen food and occasional collected specimen is necessary because no freezers will be available in the biogeoscience department to keep such material.Cost: 400 €

Overall cost of the section: 16 296 €


http://stomatopodhunters.blogspot.com/


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Project MEvo-DivA


3.2. PERSONNEL COSTSLe personnel non permanent (doctorant, post-doctorants, CDD..) financé sur le projet devra être justifié.

Fournir les profils des postes à pourvoir pour les personnels à recruter (1/2 page maximum par type de poste)

Pour le doctorant, préciser si la demande de bourse de thèse est prévue ou en cours, en préciser la nature et la part de financement imputable au projet.

Two persons will be employed on this project: Thomas Claverie as a postdoctorant and a technician for a year.

The postdoctorant will be the principal investigator of this project and will be employed full time for 36 month. He will perform most of the measurements, data analyses and results dissemination. Salary: 2 834 € per month

A full time technician will be hired for the year 2012 in order to assist the postdoctorant with animal maintenance and data analyses. This position is necessary because while field data are being collected, one person will have to be continuously present in the laboratory to maintain aquariums, feed animals and restore manipulation of specimen after each molt (see task 2). Furthermore, the type of analysis necessary in this project require time consuming data manipulation which will be accessible for a qualified technician. Salary: 2 017 € per monthTotal personal cost: 126 200 €

3.3. SUBCONTRACTINGPréciser :

- la nature des prestations

- le type de prestataire.Trawling from the RV Neomysis from the marine biological station in Roscoff will be used to collect benthic species. 30 half day trips are anticipated to be necessary.Cost: 1 280 €

3.4. TRAVELPréciser :

- les missions liées aux travaux d’acquisition sur le terrain (campagnes de mesures…)

- les missions relevant de colloques, congrès…Field work at the marine biological station in Roscoff:

Return trip Dijon-Roscoff with rental vehicle to collect specimen for developmental experiment (274 €)

4 return trips Dijon-Roscoff by train (960 €) Rental of laboratory space at the marine biological station in Roscoff for 4

month (1 600 €) Cost for 4 month accommodation within the marine station structure

(2 960 €)Measurements at the national museum of natural history in Paris:

4 return trips Dijon-Paris by train (280 €) Accommodation for 4 month (4 000 €)

Venue to conferences: Two major international conferences (2 900 €)



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Two Evolutionary Biology Meeting at Marseille (1 380 €)Overall cost of travels: 14 354 €

3.5. EXPENSES FOR INWARD BILLING (COSTS JUSTIFIED BY INTERNAL PROCEDURES OF INVOICING)

Préciser la nature des prestationsAnticipated cost for daily activity while in the Biogeoscience laboratory (e.g. printing paper). Cost: 500 €

3.6. OTHER WORKING COSTSToute dépense significative relevant de ce poste devra être justifiée.Cost incurring publication:

I estimate that of the major papers we intend to publish, approximately one per year will incur significant page charges and will represent a significant cost for open access. I have therefore budgeted 4 500 € within the term of the fellowship (3 papers @ 1,500 €).

Other costs: Ziplock bags to store collected preys (50 €) Polystyrene boxes to bring back specimen for developmental experiment

from Roscoff (40 €).Total cost: 4 590 €

4. ANNEXES

4.1. REFERENCESInclure la liste des références bibliographiques utilisées dans la partie « Etat de l’art ».1. Adamowicz, S.J., A. Purvis, and M.A. Wills, 2008, Increasing

morphological complexity as a major evolutionary trend in the Crustacea. Proc. Natl. Acad. Sci. U. S. A., 105: 4786-4791.

2. Alfaro, M.E., D.I. Bolnick, and P.C. Wainwright, 2005, Evolutionary consequences of many-to-one mapping of jaw morphology to mechanics in labrid fishes. Am. Nat., 165: E140-E154.

3. Allen, C.E., 2008, The "Eyespot Module" and eyespots as modules: development, evolution, and integration of a complex phenotype. J. Exp. Zool. Part B, 310B: 179-190.

4. Atchley, W.R. and B.K. Hall, 1991, A model for development and evolution of complex morphological structures. Biol. Rev., 66: 101-157.

5. Boxshall, G.A., 2004, The evolution of arthropod limbs. Biol. Rev., 79: 253-300.

6. Breuker, C.J., V. Debat, and C.P. Klingenberg, 2006, Functional evo-devo. Trends Ecol. Evol., 21: 488-492.

7. Breuker, C.J., et al., 2007, Integration of wings and their eyespots in the Speckled Wood butterfly Pararge aegeria. J. Exp. Zool. Part B, 308B: 454-463.

8. Brown, S.C., S.R. Cassuto, and R.W. Loos, 1979, Biomechanics of chelipeds in some decapod crustaceans. J. Zool., 188: 143-159.



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9. Browne, W.E. and N.H. Patel, 2000, Molecular genetics of crustacean feeding appendage development and diversification. Semin. Cell Dev. Biol., 11: 427-435.

10. Carroll, S.B., 2001, Chance and necessity: the evolution of morphological complexity and diversity. Nature, 409: 1102-1109.

11. Carroll, S.B., J.K. Grenier, and S.D. Weatherbee, 2005, From DNA to diversity: molecular genetics and the evolution of animal design. 2nd ed. Oxford: Blackwell Publishing. 258.

12. Carter, M.C., D.P. Gordon, and J.P.A. Gardner, 2010, Polymorphism and variation in modular animals: morphometric and density analyses of bryozoan avicularia. Mar. Ecol. Prog. Ser., 399: 117-130.

13. Chapman, A.D., 2009, Numbers of living species in Australia and the world. 2nd ed. Canberra, Australia. 81.

14. Cheverud, J.M., 1982, Phenotypic, genetic, and environmental morphological integration in the cranium. Evolution, 36: 499-516.

15. Claverie, T., E. Chan, and S.N. Patek, 2011, Modularity and scaling in fast movements: power amplification in mantis shrimp. Evolution, 65: 443-461.

16. Claverie, T. and I.P. Smith, 2007, Functional significance of an unusual chela dimorphism in a marine decapod: specialization as a weapon? Proc. R. Soc. Lond. B, 274: 3033-3038.

17. Crane, J., 1975, Fiddler crabs of the world (Ocypodidae: genus Uca). Princeton, New Jersey: Princeton University Press. 736.

18. Drake, A.G. and C.P. Klingenberg, 2010, Large-scale diversification of skull shape in domestic dogs: disparity and modularity. Am. Nat., 175: 289-301.

19. Foote, M., 1993, Contributions of individual taxa to overall morphological disparity. Paleobiology, 19: 403-419.

20. Gerhart, J. and M. Kirschner, 1997, Cells, Embryos, and Evolution: Toward a cellular and developmental understanding of phenotypic variation and evolutionary adaptability. Malden, MA: Blackwell Science. 656.

21. Giorgianni, M. and N.H. Patel. Conquering land, air and water: the evolution and development of arthropod appendages. in Evolving form and function: fossils and development. 2005. New Haven, CT.

22. Goswami, A., 2006, Cranial modularity shifts during mammalian evolution. Am. Nat., 168: 270-280.

23. Goswami, A., 2007, Cranial modularity and sequence heterochrony in mammals. Evolution & Development, 9: 290-298.

24. Goswami, A. and P.D. Polly, 2010, The influence of modularity on cranial morphological disparity in carnivora and primates (Mammalia). PLoS One, 5: e9517-e9517.

25. Govind, C.K. and J. Pearce, 1989, Critical period for determining claw asymmetry in developing lobsters. J. Exp. Zool., 249: 31-35.

26. Gregory, J.T., 1951, Convergent evolution: the jaws of Hesperornis and the mosasaurs. Evolution, 5: 345-354.

27. Hallgrímsson, B. and B.K. Hall, eds. 2005, Variation: a central concept in biology. Elsevier Academic Press: Amsterdam. 592.

28. Hallgrímsson, B., et al., 2007, Epigenetic interactions and the structure of phenotypic variation in the cranium. Evolution & Development, 9: 76-91.



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29. Hansen, T.F., 2003, Is modularity necessary for evolvability? Remarks on the relationship between pleiotropy and evolvability. BioSystems, 69: 83-94.

30. Hendry, A.P., 2009, Speciation. Nature, 458: 162-164.31. Kemp, T.S., 2007, The concept of correlated progression as the basis of a

model for the evolutionary origin of major new taxa. Proc. R. Soc. Lond. B, 274: 1667-1673.

32. Klingenberg, C.P., 2008, Morphological integration and developmental modularity. Annu. Rev. Ecol. Evol. Syst., 39: 115-132.

33. Klingenberg, C.P., 2009, Morphometric integration and modularity in configurations of landmarks: tools for evaluating a priori hypotheses. Evolution & Development, 11: 405-421.

34. Klingenberg, C.P., M. Katharina, and J.C. Auffray, 2003, Developmental integration in a complex morphological structure: how distinct are the modules in the mouse mandible? Evolution & Development, 5: 522-531.

35. Klingenberg, C.P., L.J. Leamy, and J.M. Cheverud, 2004, Integration and modularity of quantitative trait locus effects on geometric shape in the mouse mandible. Genetics, 166: 1909-1921.

36. Lee, S.Y., 1995, Cheliped size and structure: the evolution of a multifunctional decapod organ. J. Exp. Mar. Biol. Ecol., 193: 161-176.

37. Liubicich, D.M., et al., 2009, Knockdown of Parhyale Ultrabithorax recapitulates evolutionary changes in crustacean appendage morphology. Proc. Natl. Acad. Sci. U. S. A., 106: 13892-13896.

38. Magwene, P.M., 2001, New tools for studying integration and modularity. Evolution, 55: 1734-1745.

39. Marcus, L.F., 1990, Traditional morphometrics, in Proceedings of the michigan morphometrics workshop, F.J. Rohlf and F.L. Bookstein, Editors. The University of Michigan Museum of Zoology: Ann Arbor, Michigan. p. 380.

40. Mariappan, P., C. Balasundaram, and B. Schmitz, 2000, Decapod crustacean chelipeds: an overview. J. Biosci., 25: 301-313.

41. Marquez, E., 2008, A statistical framework for testing modularity in multidimensional data. Evolution, 62: 2688-2708.

42. Marroig, G. and J.M. Cheverud, 2001, A comparison of phenotypic variation and covariation patterns and the role of phylogeny, ecology, and ontogeny during cranial evolution of new world monkeys. Evolution, 55: 2576-2600.

43. Marroig, G., et al., 2009, The evolution of modularity in the mammalian skull II: evolutionary consequences. Evol. Biol., 36: 136-148.

44. Martin, J.W., K.A. Crandall, and D.L. Felder, eds. 2009, Decapod crustacean phylogenetics. Boca Raton, FL. 616.

45. Mayhew, P.J., 2007, Why are there so many insect species? Perspectives from fossils and phylogenis. Biol. Rev., 82: 425-454.

46. Mezey, J.G., J.M. Cheverud, and G.P. Wagner, 2000, Is the genotype-phenotype map modular? A statistical approach using mouse quantitative trait loci data. Genetics, 156: 305-311.

47. Monteiro, L.R., V. Bonato, and S.F. dos Reis, 2005, Evolutionary integration and morphological diversification in complex morphological



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structures: mandible shape divergence in spiny rats (Rodentia, Echimyidae). Evolution & Development, 7: 429-439.

48. Page, L., 2005, Development of foregut and proboscis in the Buccinid Neogastropod Nassarius mendicus: evolutionary opportunity exploited by a developmental module. J. Morphol., 264: 327-338.

49. Pangamiban, G., et al., 1995, The development of crustacean limbs and the evolution of arthropods. Science, 270: 1363-1366.

50. Porto, A., et al., 2009, The evolution of modularity in the mammalian skull I: morphological integration patterns and magnitudes. Evol. Biol., 34: 118-135.

51. R Development Core Team, R: A language and environment for statistical computing. 2009, R Foundation for Statistical Computing: Vienna, Austria.

52. Raup, D.M., 1966, Geometric analysis of shell coiling: general problems. J. Paleontology, 40: 1178-1190.

53. Roy, K. and M. Foote, 1997, Morphological approaches to measuring biodiversity. Trends Ecol. Evol., 12: 277-281.

54. Schlosser, G., 2002, Modularity and the units of evolution. Theor. Biosci., 121: 1-80.

55. Schluter, D., et al., 1997, Likelihood of ancestor states in adaptive radiation. Evolution, 51: 1699-1711.

56. Smith, L.D. and A.R. Palmer, 1994, Effects of manipulated diet on size and performance of brachyuran crab claws. Science, 264: 710-712.

57. Wagner, G. and L. Altenberg, 1996, Perspective: complex adaptations and the evolution of evolvability. Evolution, 50: 967-976.

58. Wagner, G.P., 1996, Homologues, natural kinds and the evolution of modularity. Amer. Zool., 36: 36-43.

59. Wagner, G.P., M. Pavlicev, and J.M. Cheverud, 2007, The road to modularity. Nature Reviews Genetics, 8: 921-931.

60. West-Eberhard, M.J., 2003, Developmental Plasticity and Evolution. New York, USA: Oxford University Press.

61. Williams, T.A. and L.M. Nagy, 2001, Developmental modularity and the evolutionary diversification of arthropod limbs. J. Exp. Zool. Part B, 291: 241-257.

62. Yang, S.A., 2001, Modularity, evolvability, and adaptative radiations: a comparison of the hemi- and holometabolous insects. Evolution & Development, 3: 59-72.

63. Young, N.M. and B. Hallgrimsson, 2005, Serial homology and the evolution of mammalian limb covariation structure. Evolution, 59: 2691-2704.

64. Zelditch, M.L., A.R. Wood, and D.L. Swiderski, 2009, Building developmental integration into functional systems: function-induced integration of mandibular shape. Evol. Biol., 36: 71-87.



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4.2. INVOLVEMENT OF THE CANDIDATE IN OTHER CONTRACTSMentionner ici les projets en cours d’évaluation soit au sein de programmes de l’ANR, soit auprès d’organismes, de fondations, à l’Union Européenne, etc. que ce soit comme coordinateur ou comme partenaire. Pour chacun, donner le nom de l’appel à projets, le titre du projet et le nom du coordinateur.I am not involved in other contracts

Part. Person-months

Programme NameFunding body

Funding amount

Funding bodyFunding amount

Project Title Name of co-ordinator

Start and Finish dates

N°

4.3. ADDITIONAL ADMINISTRATIVE DOCUMENT PROGRAMME RETOUR POST-DOCTORANTS 2011

PROJECT ACRONYM : Candidate (project leader)

Gender Firstname Lastname

Tél. Tél. portable E-mail

Date of birth Nationality Place of birth

Personal address

Date and place of PhD submission

Starting date and place of last post-doc position abroad (out of France)

Closing date and place of last post-doc abroad (out of France)



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Project MEvo-DivA


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