Emmanuel Fara

ImageEmmanuel FARA, Ph.D.

Assistant Professor in Palaeontology (with Habilitation)

Laboratoire Biogéosciences (UMR CNRS / uB 6282)

UFR Sciences de la Vie, de la Terre et de l'Environnement (SVTE)

Université de Bourgogne

6 boulevard Gabriel

21000 Dijon


Tél : +33 380-39-39-70

Fax : +33 380-39-63-87

Email : This email address is being protected from spambots. You need JavaScript enabled to view it.


What are the processes structuring biodiversity at the local, regional, and global scale? How do biological communities respond to environmental changes and what are the associated kinetics?

My research activities revolve around these questions and they concern two major axes:

  • The first one focuses on the evolution and structure of local and regional communities in relation to palaeoenvironment. Both the temporal and spatial components of the patterns are investigated using fossil assemblages of invertebrates and vertebrates.
  • The second axis is a long term project with an ambitious aim: to define macroevolutionary patterns and processes for Phanerozoïc diversifications. It consists in the development of new datasets and algorithms for testing current models of diversification and recovery dynamics.

The general objective is to identify the mechanisms driving biodiversity and extinction through time and to define the spatio-temporal scales at which these processes occur. As such, my research is well-anchored into macroevolution and macroecology, two disciplines for which the fossil record is an ideal working area.

Studied models


Tethyan ammonites from the Pliensbachian (Early Jurassic). Collaboration with Jean-Louis Dommergues, Christian Meister and Pascal Neige.

Marine benthic macrofaunas from the Lutetian (Eocene) of the Paris Basin. Collaboration with Didier Merle (MNHN) et Pascal Neige (uB).

Ichthyofauna of the Early Cretaceous Santana Formation, northeastern Brazil.

Mammal assemblages from the Mio-Pliocene of Africa. Collaboration with MPFT research group (Mission Paléoanthropologique Franco-Tchadienne).

Datasets on fossil tetrapods and participation to the Paleobiology Database

Main results

Quality of the fossil record

  • Re-evaluation of the « Lazarus effect », an important marcoevolutionary notion at the interface between biological processes and the quality of the fossil record (Fara 2001)
  • Demonstration of the good quality of the fossil record for Cretaceous continental tetrapods (Fara & Benton 2000, Fara 2002, Fara 2004).

Global diversification dynamics

  • Documentation of mega-biases in the current global diversification paradigms for tetrapods (Alroy et al. 2003, and MS submitted), and participation to the Paleobiology Database.
  • Evidence for a strong extinction selectivity among tetrapod families during the K/T biotic crisis (Fara 2000).
  • Proposal of a new method for assessing the minimal species diversity of taxonomic groups with poor fossil records (Fara 2004).
  • First quantitative estimates of the diversity dynamics for lissamphibians and lepidosaurs in the Jurassic-Eocene interval (Ph.D. thesis and Fara, 2004)
  • First quantitative estimate of the fossil record and diversity of gonorynchiform fishes (Fara et al 2007, Fara et al. 2009)

Community evolution

  • Evidence for a local environemental control upon the structure of Pliocene mammal communities in Chad (Fara et al., 2005a).
  • Evidence for the structural evolution of Cretaceous bony fish communities (Santana Formation, Araripe Basin, northeastern Brazil) during the opening of the South Atlantic Ocean (Fara et al., 2005b)


  • Revision of a phytosaurs species (Archosauria) and challenge of its use as a biochronological indicator for the Triassic (Fara & Hungerbühler 2000)
  • Description of the first occurrence of thalattosuchian crocodiles in the Jurassic of Tunisia (Fara et al. 2002)
  • Discovery of the oldest fossil snake from Brazil, an anilioïd from the Late Cretaceous (Zaher et al., 2003).



Articles in journals and book chapters

  • Hardy C., Fara E., Laffont R., Dommergues J.-L., Meister C. & Neige P. (2012) - Deep-time phylogenetic clustering of extinctions in an evolutionarily dynamic clade (Early Jurassic ammonites) -   PLoS ONE 7(5): e37977. doi:10.1371/journal.pone.0037977  [PDF]
  • Moreau J.-D., Gand G., Fara E. & Michelin A. (2012) - Biometric and morphometric approaches on Lower Hettangian dinosaur footprints from the Rodez Strait (Aveyron, France) - C. R. Palevol 11 (4): 231-239. [PDF]
  • Le Fur S., Fara E. & Vignaud P. (2011) - Effect of simulated faunal impoverishment and mixture on the ecological structure of modern mammal faunas: implications for the reconstruction of Mio-Pliocene African palaeoenvironments - Palaeogeography Palaeoclimatology Palaeoecology 305:295-309. [PDF]
  • Escarguel, G., Fara E., Brayard, A. & Legendre, S. (2011) – Biodiversity is not (and never has been) a bed of roses ! – Comptes Rendus Biologies 334: 351-359. [PDF]
  • Dera G., Neige P., Dommergues J.-L., Fara E., Laffont R. & Pellenard P. (2010) High-resolution dynamics of Early Jurassic marine extinctions: the case of Pliensbachian-Toarcian ammonites (Cephalopoda) - Journal of the Geological Society of London 167: 21-33. [PDF]
  • Tabuce R, Marivaux L, Lebrun R., Adaci M., Bensalah M., Fabre P-H., Fara E., Gomes Rodrigues H., Hautier L., Jaeger J-J., Lazzari V., Mebrouk F., Peigné S., Sudre J, Tafforeau P., Valentin X. & Mahboubi M. (2009) - Anthropoid versus strepsirhine status of the African Eocene primates Algeripithecus and Azibius: craniodental evidence - Proceedings of the Royal Society B (2009) 276: 4087-4094. Published online at doi:10.1098/rspb.2009.1339. [PDF]
  • Fara E., Gayet M. & Taverne L. (2009) Chapter 6: "The Fossil Record of Gonorynchiforms". In: Poyato-Ariza, F.J., Grande, T., & Diogo, R. (Eds.) - Teleostean Fish Biology; A Comprehensive Examination of Major Taxa.  Vol. 1: Gonorhynchiformes and Ostariophysan Relationships: a Comprehensive Review. Science Publishers Inc., Enfield, New Hampshire, USA, pp. 173-226. [PDF]
  • Dommergues J-L., Fara E. & Meister C. (2009). Ammonite diversity and its palaeobiogeographical structure during the early Pliensbachian (Jurassic) in the western Tethys and adjacent areas - Palaeogeography Palaeoclimatology Palaeoecology 280: 64-77. [PDF]
  • Le Fur S., Fara E., Mackaye H.T., Vignaud P. & Brunet M. (2009). The mammal assemblage of the hominid site TM266 (Late Miocene, Chad Basin): ecological structure and paleoenvironmental implications - Naturwissenschaften 96: 565-574. [PDF]
  • Viriot L., Vignaud P., Boisserie J.-R., Fara E., Guy F., Lihoreau F., Lehman T., Schuster M., Blondel C., Duringer Likius A., Mackaye H.T., Otero O.& Brunet M. (2008) Late Miocene to Early Pliocene Chadian vertebrate faunas: palaeoenvironmental and palaeobiogeographical implications - The Geology of East Libya 3, 303-308. [PDF]
  • Saraiva A.Á.F., Hessel M.H., Guerra N.C. & Fara E. (2007) Concreções calcárias da Formação Santana, Bacia do Araripe: uma proposta de classificação Estudos Geológicos 17 (1): 40-57. [PDF]
  • Fara E., Gayet M. & Taverne L. (2007) Les Gonorynchiformes fossiles: distribution et diversité – Cybium 31  (2): 115-122. [PDF]
  • Pinton A., Fara E. & Otero O. (2006) - Spine anatomy reveals the diversity of catfish through time: a case study of Synodontis (Siluriformes) -  Naturwissenschaften 93: 22-26. [PDF]
  • Fara E., Likius A., Mackaye H.T., Vignaud, P. & Brunet, M. (2005) - Pliocene large-mammal assemblages from northern Chad: sampling and ecological structure - Naturwissenschaften 92: 537-541. [PDF]
  • Fara E., Saraiva A., Campos D.A., Moreira J.K.R., Siebra D.C. & Kellner A.W.A. (2005) - Controlled excavations in the Romualdo Member of the Santana Formation (Araripe Basin, Lower Cretaceous, northeastern Brazil): stratigraphic, palaeoenvironmental and palaeoecological implications - Palaeogeography Palaeoclimatology Palaeoecology 218 (1-2): 145-160. [PDF]
  • Fara E. & Langer M.C. (2004) - Estimates of phylogeny and biochronology - Revista Brasileira de Paleontologia 7(3) : 301-309. [PDF]
  • Fara E. (2004) - Estimating minimum global species diversity for groups with a poor fossil record: a case study of Late Jurassic-Eocene lissamphibians - Palaeogeography Palaeoclimatology Palaeoecology, 207 (1-2): 59-82. [PDF]
  • Fara E. (2002) - Sea-level variations and the quality of the continental fossil record -  Journal of the Geological Society of London, 159: 489-491. [PDF]
  • Fara E., Ouja M., Buffetaut E. & Srarfi D. (2002) - First occurrences of thalattosuchian crocodiles in the Middle and Upper Jurassic of Tunisia - Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 2002 (8): 465-476. [PDF]
  • Fara E. (2001) - What are Lazarus taxa? - Geological Journal, 36: 291-303. [PDF]
  • Fara E. & Hungerbühler A. (2000) - "Paleorhinus magnoculus" from the Upper Triassic of Morocco: a juvenile primitive phytosaur (Archosauria) - Comptes-Rendus de l'Académie des Sciences, Paris IIa, 331 (12): 831-836. [PDF]
  • Fara E. & Benton M.J. (2000) - The fossil record of Cretaceous tetrapods - Palaios, 15 (2): 161-165. [PDF]
  • Fara E. (2000) - Diversity of Callovian-Ypresian (Middle Jurassic-Eocene) tetrapod families and selectivity of extinctions at the K/T boundary - Geobios, 33(3): 387-396. [PDF]

Published abstracts

  • Fara, E. & Saraiva, A. (2009) - Stratigraphy and vertebrate diversity of the Santana Formation (Lower Cretaceous, Brazil): two centuries of misappreciation - Journal of Vertebrate Paleontology, 29 (3): 94A.
  • Alroy J., Carrano M., Clyde W., Fara E., Head J., Hunter J., Uhen M. & Whateley R. (2005) - Ecological spectra through the Cretaceous and Cenozoic tetrapod fossil record - Journal of Vertebrate Paleontology, 25 (3): 31A.
  • Zaher H.D., Langer M.C., Fara E., Carvalho I.S., Arruda J.T. (2003) - A mais antiga serpente (anilioidea) brasileira: Cretaceo Superior do Grupo Bauru, General Salgado, SP - Paleontologia em Destaque 44: 50-51.
  • Alroy J., Behrensmeyer K., Carrano M., Clyde W., Fara E., Fortelius M., Head J., Hunter J., Uhen M. & Wang X. (2003) - The 5% Project: just how good is the fossil record of tetrapods? - Journal of Vertebrate Paleontology, 23 (3): 29A
  • Fara E. & Langer M.C. (2000) - Using cladograms for biostratigraphic inferences - Journal of Vertebrate Paleontology, 20 (3): 40A.

Popularization articles

  • Fara E. (2006) - A Chapada do Araripe: riqueza e perspectiva paleontológica - Publicaçao Especial da Fundaçao Araripe, Crato, Brazil.


  • Fara E. 2011. “ From Macroevolution to macroecology: multi-scale analysis of fossil biodiversity"  – Habilitation Thesis (HDR), University of Burgundy [in French and English; 266 pages]

  • Fara E. 2001. “ Macroevolutionary patterns and the quality of the fossil record: from data to processes” – Ph. D. Thesis, University of Bristol [in English; 318 pages; 87 figures; 1 CD-Rom]

  • Fara E. 1998. “Les familles de tétrapodes au cours du Crétacé: diversité, paléoécologie, et registre fossile” – Master Thesis, Université de Montpellier II , France  [in French, 75 pages]

Current Fieldwork

* Santana Formation (Lower Cretaceous), Ceará State, Brazil.

* Baurú Group  (Upper Cretaceous), São Paulo State, Brazil

* Lutetian outcrops from the Paris Basin.

* Mio-Pliocene sites from the Djourab erg, Chad (org : MPFT).



As an Assitant Professor, Université de Bourgogne

Date: since January 2006

Site: Université de Bourgogne, Dijon, France

Level: From the first year to the fifth year (BSc and MSc levels)

Contents: Palaeontology, Geology, Ecology, Macroecology, Evolution, Embryology.

As a Demonstrator and Lecturer

Date: 1999-2006

Site: University of Bristol (UK), University of Mahasarakham (Thaïland), Universidade Federal do Pernambuco, Recife (Brazil), Université de Clermont-Ferrand (France), Université Montpellier II (France), Université de Poitiers (France).

Levels: the first year to the fifth year (BSc and MSc levels).

Contents: Palaeontology, Geology, Palaeoceanography, Systematics, Evolution, Database, Population genetics.


Selected conferences

  • Conference on the history of biodiversity [in French], International Conference Centre of Cluny (ENSAM), France (october 2008).
  • Conference on the evolution of biodiversity and mass extinctions [in French], La Cité des Sciences, Paris (June 2007). You can download it here.
  • Series of conferences entitled « Recent Advances and Methods in Palaeobiology », University of Mahasarakham, Thaïland (June 2004)

Documentaries and exhibits

  • Annual participations to the science popularization days.
  • Participation to the documentary « Toumaï, le nouvel ancêtre », by Pierre Stine (Gédéon Programmes). Co-production France Télévision-BBC-NHK-TSR-RTB.
  • Co-organizer of the exhibit "Ciências da Terra, Ciências da Vida - Chapada do Araripe", São Paulo, Brazil [August 30 - October 31, 2004] and scientific supervision of the catalogue ("Ciências da Terra, Ciências da Vida - Chapada do Araripe", São Paulo, Brazil, edited by the Fundação A.A. Penteado/MAB/FAAP, 2004).
  • Guided visits of various palaeontological excavation sites and exhibits.


2011: Habilitation to conduct and supervise research 

Since 2006 : Full-time Assistant Professor, Université de Bourgogne, Dijon, France

2003 – 2005: CNRS Post-doc on mammal communities, Université de Poitiers, France

2003 (July–Sept.): Visiting Fellow, Universidade Regional do Cariri –URCA (Brazil)

2002 - 2003: Lecturer, Université de Clermont-Ferrand (France) and Mahasarakham University (Thaïland).

2001 - 2002: self-funded Post-doc, Université Montpellier I, France.


1998-2001: Ph.D. in Palaeobiology, University of Bristol (U.K.)

Title: “Macroevolutionary patterns and the quality of the fossil record: from data to processes”, Supervisor: Pr. Michael. J. Benton

1998: M.Sc. in Palaeontology, Université Montpellier II (France)

1997: [National Service]

1995-1996: BSc and Graduation in Natural Sciences, Université Lyon I (France)