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Associate Professor - Head of the team EVOFONCT

Muséum National d’Histoire naturelle
Institut de Systématique, Evolution, Biodiversité
Sorbonne Universités
Equipe Evolution fonctionnelle et Systématique (EVOFONCT)
57 rue Cuvier - CP50
F- 75005 Paris, France
☏ : (+33) 1 40 79 81 26
Fax : (+33) 1 40 79 56 79
Email : legendre’at’

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My research deals with the evolution of behavior in insects. I reconstruct phylogenies before using them as a historical reference to test evolutionary hypotheses. Following this phylogenetic approach, I mainly investigate the evolution of social behavior in termites and cockroaches (Insecta : Dictyoptera).

I am also the curator of Isoptera (termites), Embioptera, Ephemeroptera, Plecoptera, Psocoptera, Trichoptera and Zoraptera in Paris Museum


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Soldiers and workers in Singapore

Given its paradoxical dimension with regard to natural selection, social behavior has long been of central interest in biology. Indeed, in the “Origin of species”, Darwin already mentioned how sterile castes of insect societies were a “special difficulty, which at first appeared to [him] insuperable”.

Living in a group brings diverse advantages and constraints to individuals. If advantages exceed drawbacks, natural selection can readily explain group living origin. Few exaggerated forms of social behavior like altruism, however, seem hardly interpretable in the only light of natural selection. Thus, social behavior evolution still remains a tremendous challenge, which is taken up using different approaches : ethology, population genetics, phylogeny, genomics…

My work aims at studying the evolution of social behavior in a phylogenetic framework.

1. Social evolution and evolutionary trends

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Social categories
From Wilson (1971)

Social behavior evolution has always been seen as a progressive, irreversible and complexity-increasing path, with eusociality as the ultimate step. Even though some exceptions to this linear trend have been shown, ‘laws’ of specialization and of irreversibility are still closely tied to the evolution of social behavior.

Classification of social categories illustrates these evolutionary trends, each category being a more complex step than the previous one. Complexity increase with the acquisition of one or several key features, which would make individuals better adapted to their environment and thus would be selectively advantageous.

These evolutionary trends can be tested in a phylogenetic framework : Comparing expected patterns under a null hypothesis (e.g., evolution is linear) with the observed patterns enable us to evaluate how adequate to social evolution of our model organisms this hypothesis is.

2. Evolutionary forces and modalities of social evolution

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The aptive triangle
From Gould (2002)

In social behavior classification, a switch from one category to another is associated with the acquisition of at least one key feature. The acquisition of such key features has been explained mostly in a functional context. In other words, key features are seen as adaptations.

This scenario overemphasizes the importance of ecological pressures in social behavior evolution to the detriment of other pressures. Yet, alternatives and complements to natural selection (e.g., phylogenetic inertia) may play a significant role in social evolution.

Phylogenetic analyses and behavioral observations can help to highlight the relative importance of these different forces.

3. Phylogenetic analysis of behavior
Studying the evolution of behavior in a phylogenetic framework requires the availability of phylogenetic hypotheses. Consequently, the first step of most of my research projects is to build a phylogenetic hypothesis using molecules and/or morphology. Non-traditional characters like behavioral ones are also integrated in phylogenetic matrices. Such process allows a better definition of behaviors observed and also a more thorough investigation of social behavior evolution hypotheses. But it might also require adapting or developing methodology to reach an adequate phylogenetic framework.

Therefore, building new tools for the phylogenetic analysis of behavior is part of my research projects.


- Colinet, C., Exbrayat, M., Robillard, T. and Legendre, F.. 2016. Species richness and composition in cockroaches (Dictyoptera : Blattodea) along an elevational gradient in Papua New Guinea. In : Insects of Mount Wilhelm, Papua New Guinea (Robillard, T., Legendre, F., Villemant C. and Leponce M., Eds), 206 : 129-140.

- Leponce, M., Novotny, V., Pascal, O., Robillard, T., Legendre, F., Villemant, C. et al. 2016. Land module of Our Planet Reviewed, Papua New Guinea : aims, methods and first taxonomical results. In : Insects of Mount Wilhelm, Papua New Guinea (Robillard, T., Legendre, F., Villemant C. and Leponce M., Eds), 206 : 11-48.

- Robillard, T., Dong, J., Legendre, F. and Agauvoa S. 2016. The brachypterous Lebinthini crickets from Papua New Guinea, with description of two genera and four new species (Orthoptera : Gryllidae : Eneopterinae). In : Insects of Mount Wilhelm, Papua New Guinea (Robillard, T., Legendre, F., Villemant C. and Leponce M., Eds), 206 : 149-202.

- Robillard, T., Legendre, F., Villemant C. and Leponce M. 2016 Insects of Mount Wilhelm, Papua New Guinea. Muséum national d’Histoire naturelle ; Mémoires du Muséum national d’Histoire naturelle 209 : 573 pp.

- Goutte S., Dubois A., Howard S.D., Marquez R., Rowley J.J.L., Dehling J.M., Grandcolas P., Xiong R., Legendre F. 2016. Environmental constraints and call evolution in torrent dwelling frogs. Evolution 70 : 811–826.

- Legendre, F., Nel, A., Svenson, G. J., Robillard, T., Pellens, R., & Grandcolas, P. (2015). Phylogeny of Dictyoptera : Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence. Plos One, 10 (7) : e0130127.

- Chintauan-Marquier*, I.C., Legendre*, F., Hugel, S., Robillard, T., Grandcolas, P., Nel, A., Zuccon, D. and Desutter-Grandcolas, L. (2015). Laying the foundations of evolutionary and systematic studies in crickets (Insecta, Orthoptera) : a multilocus phylogenetic analysis. Cladistics 32 : 54-81. * = equal contributions.

- Legendre, F., Deleporte, P., Depraetere, M., Gasc, A., Pellens, R., Grandcolas, P. (2015). Dyadic behavioural interactions in cockroaches (Blaberidae) : ecomorphological and evolutionary implications. Behaviour 152(9) : 1229-1256.

- Caesar, M., Roy, R., Legendre, F., Grandcolas, P. and Pellens, R. (2015). Catalogue of Dictyoptera from Syria and neighbouring countries (Lebanon, Turkey, Iraq and Jordan). Zootaxa 3948 : 71-92

- Picart, L., Forget, P.-M., D’Haese, C.A., Daugeron, C. Beni, S., Bounzel, R., Kergresse, E., Legendre, F., Murienne, J. and Guilbert, E. (2014). The cafotrop method : an imporved rope-climbing method for access and movement in the canopy. Ecotropica 20 : 45-52

- Grandcolas, P., Nattier, R., Pellens, R. and Legendre, F. (2014) Diversity and distribution of the genus Rothisilpha (Dictyoptera, Blattidae) in New Caledonia : Evidence from new microendemic species. In : Zoologia Neocedonica 8. Biodiversity studies in New Caledonia (Guilbert, E., Robillard, T., Jourdan, H and Grandcolas, P. (Eds), Muséum national d’Histoire naturelle, Paris : 299-308

- Nel, A., Prokop, J., Grandcolas, P., Garrouste, R., Lapeyrie, J., Legendre, F., Anisyutkin, L., and Kirejtshuk, A. G. (2014). The beetle-like Palaeozoic and Mesozoic roachoids of the so-called “umenocoleoid” lineage (Dictyoptera : Ponopterixidae fam. nov.). Comptes Rendus Palevol. 13 : 545-554

- Coty, D., Aria, C., Garrouste, R., Wils, P., Legendre, F. and Nel, A. (2014) The first ant-termite syninclusion in amber with CT-scan analysis of taphonomy. PLoS ONE 9 : e104410

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BJLS 2014

- Legendre, F., D’Haese, C. A., Deleporte P., Pellens, R., Whiting M. F., Schliep K. and Grandcolas, P. (2014) The evolution of social behaviour in Blaberid cockroaches with diverse habitats and social systems : phylogenetic analysis of behavioural sequences. Biological Journal of the Linnean Society 111 : 58-77

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- Goutte, S., Dubois A. and Legendre, F.. (2013) The importance of ambient sound level to characterise anuran habitat. PLoS ONE 10(8):e78020

- Legendre, F. and Grandcolas, P. Ordre Dictyoptera : Blattodea. In : Les Insectes du monde - classification - clés pour la reconnaissance des principales familles, (Aberlenc H-P. and Delvare, G., Eds) ; (in press)

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Evol & Dvpt 2013

- Legendre, F., Whiting, M. F. and Grandcolas, P. (2013) Phylogenetic analyses of termite post-embryonic sequences illuminate caste and developmental pathway evolution. Evolution & Development 15 : 146-157

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Animal Behaviour 2012

- Legendre, F., Marting, P.R. and Cocroft, R.B. (2012) Competitive masking of vibrational signals during mate searching in a treehopper. Animal Behaviour 83 : 361-368

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Cladistics 2011

- Grandcolas, P., Nattier, R., Legendre, F. and Pellens, R. (2011) Mapping extrinsic traits such as extinction risks or modelled bioclimatic niches on phylogenies : does it make sense at all ? Cladistics 27 : 181-185

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Syst Entomol 2010

- Legendre, F., Robillard, T., Song, H., Whiting, M. F. and Desutter-Grandcolas (2010). One hundred years of instability in Ensiferan relationships. Systematic Entomology 35 : 475-488

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BJLS 2008

- Legendre, F., Robillard, T., Desutter-Grandcolas, L., Whiting, M.F. and Grandcolas, P. (2008). Phylogenetic analysis of non-stereotyped behavioral sequences with a successive event-pairing method. Biological Journal of the Linnean Society 94 : 853-867

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QRB 2008

- Legendre, F. (2008). Cockroaches : Ecology, Behavior, and Natural History (by Bell, W. J., Roth, L. M. et Nalepa, C. A.) Book Review. The Quarterly Review of Biology 83 : 222

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MPE 2008

- Legendre, F., Whiting, M. F., Bordereau, C., Cancello, E. M., Evans, T. and Grandcolas, P. (2008). The phylogeny of termites (Dictyoptera : Isoptera) based on mitochondrial and nuclear genes : implications for the evolution of the worker and pseudergate castes, and foraging behaviors. Molecular Phylogenetics and Evolution 48 : 615-627

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JIB 2008

- Legendre, F., Pellens, R. and Grandcolas, P. (2008). A comparison of behavioral interactions in solitary and presocial Zetoborinae cockroaches (Blattaria, Blaberidae). Journal of Insect Behavior 21 : 351-365

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Cladistics 2007

- Desutter-Grandcolas, L., Legendre, F., Grandcolas, P., Robillard, T. and Murienne, J. (2007). Distinguishing between convergence and parallelism is central to comparative biology : a reply to Williams and Ebach. Cladistics 23 : 90-94

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MPE 2007

- Pellens, R., D’Haese, C. A., Bellés, X., Piulachs, M-D., Legendre, F., Wheeler, W. and Grandcolas, P. (2007). The evolutionary transition from subsocial to eusocial behavior in Dictyoptera : phylogenetic evidence for modification of the "shift-in-dependent-care" hypothesis with a new subsocial cockroach. Molecular Phylogenetics and Evolution 43 : 616-626

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Stud Neo Fauna 2007

- Pellens, R., Legendre, F. and Grandcolas, P. (2007). Phylogenetic analysis of social behavior evolution in [Zetoborinae + Blaberinae + Gyninae + Diplopterinae] cockroaches : an update with the study of endemic radiations from the Atlantic forest. Studies in Neotropical Fauna 42 : 25-31

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Cladistics 2006

- Robillard, T., Legendre, F., Desutter-Grandcolas, L. and Grandcolas, P. (2006). Phylogenetic analysis and alignment of behavioral sequences by direct optimization. Cladistics 22 : 602-633

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Cladistics 2005

- Desutter-Grandcolas, L., Legendre, F., Grandcolas, P., Robillard, T. and Murienne, J. (2005). Convergence and parallelism : is a new life ahead of old concepts ? Cladistics 21:51-61

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Cladistics 2005

- Murienne, J., Grandcolas, P., Piulachs, M. D., Bellés, X., D’Haese, C., Legendre, F., Pellens, R. and Guilbert, E. (2005). Evolution on a shaky piece of Gondwana : is local endemism recent in New Caledonia ? Cladistics 21 : 2-7

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Zoosystema 2005

- Grandcolas, P., Legendre, F., Park, Y. C., Bellés, X., Murienne, J. and Pellens, R. The genus Cryptocercus in East Asia : distribution and new species (Insecta, Dictyoptera, Blattaria, Polyphagidae). Zoosystema 27 : 725-732

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Cladistics 2004

- Grandcolas, P., Guilbert, E., Robillard, T., D’Haese, C. A., Murienne, J. and Legendre, F. (2004). Mapping characters on a tree with or without the outgroups. Cladistics 20 : 579-582


Graduate students

  • Dominic Evangelista (NSF post-doctoral fellow, 2016-2018) : Phylogenomics and diversification in Blaberoidea. Co-advisor with Akito Kawahara.
  • Julien Troudet (October 2014 - ) : Latitudinal Diveristy Gradient : the message of systematics through the biodiveristy databases - co-direction with Régine Vignes-Lebbe.
  • Sandra Goutte (PhD fellow, September 2010 - June 2014) : Adaptations to noisy torrent environments in frogs : combining morphological, eco-ethological and phylogenetic approaches - co-direction with Alain Dubois.
    Undergraduate students
  • France Thouzé (Master 2 Labex BCDiv- co-direction with Philippe Lopez)
  • Augustin Lafond (Master 2 - co-direction with Tony Robillard)
  • Laëtitia Labore (Master 1)
  • Delia Dupré (Master 1)
  • Mylène Exbrayat (Licence 3)
  • Camille Colinet (Licence 2)
  • Lauriane Pindrais (BTS 2nd year)
  • François guichard (BTS 2nd year)
  • Gabin Colombini (BTS 2nd year)
  • Quentin Lefèvre (BTS 2nd year)
  • François Moreau (BTS 2nd year)

Regular classes :

  • Phylogenetic analysis of behavioral sequences (Doctoral school of MNHN)
  • Information sources in phylogenetics : behavioral data (Master 1 of UPMC - Paris VI)
  • Phylogenetic tests of evolutionary hypotheses (Master 1 of UPMC - Paris VI)
  • Probabilistic phylogenetic inference (Master 1 and 2 of UPMC - Paris VI)
  • Phylogenetic reconstruction : why and how ? (Formation of High School teachers)

Ponctual interventions :

  • Non-traditional characters in phylogenetics (Doctoral school of MNHN)
  • Classification of living things (Guest lecture in Paul Eluard Middle School, Montreuil)


- New Caledonia 2016. Our Planet Reviewed. Study of cockroach communitues and micro-endemism.
- French Guiana 2015. Our Planet Reviewed. Collecting Orthopteroid Insects in Mitaraka.

- Philippines 2014. Collecting Blaberidae and Blattidae cockroaches.

- United States 2013. Collecting in Virginia (Jefferson National Forest).

- Papua New Guinea 2012. Our Planet Reviewed, IBISCA Niugini. Collecting Orthopteroid Insects along an altitudinal transect on Mount Wilhelm.

- New-Caledonia 2010. Rapid Assessment Program (Conservation International) on Mount Panié. In charge of Orthopteroïd Insects.

- Indonesia 2010. Study of cockroach communities with emphasis on Perisphaerus sp. (Blaberidae : Perisphaeriinae).

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Perisphaerus flexicollis

- Singapore 2009. Collecting cockroaches with a special emphasis on the subsocial species Perisphaerus flexicollis (Blaberidae : Perisphaeriinae).

Perisphaerus flexicollis exhibits subsocial behaviours like nymph-guarding by females. Also, first instar nymphs probably feed upon nourishment produced by the mother.

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Tylopelta americana

- United States 2008. Field observations and collecting of Tylopelta gibbera (Hemiptera : Membracidae) in Missouri.

Like other treehoppers studied so far, Tylopelta americana communicates through vibrations transmitted via its host plant (Desmodium sp.)
I investigated male localisation behaviour of a mating partner and male-male competition in such behaviour.

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Thanatophyllum akinetum
Female with its brood

- French Guyana 2006. Study of cockroach communities on Kaw Mountain.

Thanatophyllum akinetum is a solitary Zetoborinae living in the litter, whereas Paradicta rotunda is a gregarious Blaberinae living in treeholes.


  • Action Transversale du MNHN (20,000 €) - 2011-2015.
  • Lavoisier Grant (18,000 €), Ministère des Affaires Etrangères, December 2007 - December 2008.
  • Rosen Price (500 $), 3rd best student oral presentation, Hennig XXV, Oaxaca (Mexico), 2006.
  • Marie Stopes Student Travel Award (500 $) in 2005, 2006 and 2007.
  • Travel Grant from Paris Doctoral School ED 392 (1,000 €) in 2006 and 2007
  • PhD Grant, Ministère de l’Education nationale, de la Recherche et des Technologies, October 2004 - September 2007.
  • Master 2 Grant, Ministère de l’Education nationale, de la Recherche et des Technologies, October 2003 - June 2004.

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