Mathieu GAREL - Personal Home Page

Research | Contact Information | Publications | Thesis


I am a research engineer working within the Office National de la Chasse et de la Faune Sauvage. Most of my work is focused on ecology and evolution of life history traits in large herbivores; with a special interest for mountain ungulates and wildlife management.

Contact Information

ONCFS - CNERA Faune de Montagne
5 allée de Bethléem
Z.I. Mayencin
38610 Gières

Phone: +33 (0)4 76 59 32 07
Fax: +33 (0)4 76 89 33 74


Peer Reviewed Articles

60. Akl, T. , Bourgoin, G., Souq, M.-L., Appolinaire, J., Poirel, M.-T., Gibert, P., Abi Rizk, G., Garel, M. & Zenner, L. 2019. Detection of Tick-Borne Pathogens in Questing Ixodes ricinus of the French Pyrenees and first identification of Rickettsia monacensis in France. Parasite, in press  

59. Portanier, E., Garel, M., Devillard, S., Maillard, D., Poissant, J., Galan, M., Benabed, S., Poirel, M.-T., Duhayer, J., Itty, C., Bourgoin, G. 2019. Both candidate gene and neutral genetic diversity correlate with parasite resistance in female Mediterranean mouflon. BMC Ecology, 19, 1-14    Download pdf file

58.- Espunyes, J., Bartolomé, J., Garel, M., Gálvez-Cerón, A., Fernández-Aguilar, X., Colom-Cadena, A., Calleja, J. A., Gassó, D., Jarque, L., Lavín, S., Marco, I. and Serrano, E. 2019. Seasonal diet composition of Pyrenean chamois is mainly shaped by primary production waves. PLoS ONE, 14, 1-23  Download pdf file

57.- Munden, R., Börger, L., Wilson, R. P., Redcliffe, J., Loison, A., Garel, M. & Potts, J. R. 2018. Making sense of ultrahigh-resolution movement data: A new algorithm for inferring sites of interest. Ecology and Evolution, DOI: 10.1002/ece3.4721  Download pdf file

56.- Lambert, S., Ezanno, P, Garel, M. and Gilot-Fromont, E. 2018. Demographic stochasticity drives epidemiological patterns in wildlife with implications for diseases and population management. Scientific Reports, 8, 16846  Download pdf file

55.- Portanier, E., Larroque, J., Garel, M., Marchand, P., Maillard, D., Bourgoin, G. & Devillard, S. 2018. Landscape genetics matches with behavioral ecology and brings new insight on the functional connectivity in Mediterranean mouflon. Landscape Ecology, 33, 1069-1085  Download pdf file

54.- Peron, G., Duparc, A., Garel, M., Marchand, P., Morellet, N., Said, S. & Loison, A. 2018. Circadian periodicity in space use by ungulates of temperate regions: how much, when, and why? Journal of Animal Ecology, 87, 1299-1308  

53.- Thuiller, W., Guéguen, M., Bison, M., Duparc, A., Garel, M., Loison, A., Renaud, J. & Poggiato, G. 2018. Combining point-process and landscape vegetation models to predict large herbivore distributions in space and time -- a case study of Rupicapra rupicapra. Diversity and Distributions, 24, 352-362  Download pdf file

52.- Bourgoin, G., Marchand, P., Hewison, A.J.M., Ruckstuhl, K.E. & Garel, M. Social behaviour as a predominant driver of sexual, age-dependent and reproductive segregation in Mediterranean mouflon. 2018. Animal Behaviour, 136, 87-100  Download pdf file

51.- Gilot-Fromont, E., Garel, M., Gibert, P., Lambert, S., Menaut, P., Bonetti, B., Game, Y., Reynaud, G. & Foulché, K. 2018. Self-clearance of pestivirus in a Pyrenean chamois population. Journal of Wildlife Diseases, 54, 335-341  Download pdf file

50.- Portanier, E., Garel, M., Devillard, S., Marchand, P., Andru, J., Maillard, D. & Bourgoin, G. 2017. Introduction history overrides social factors in explaining genetic structure of females in Mediterranean mouflon. Ecology and Evolution, 7, 9580-9591  Download pdf file

49.- Villamuelas, M., Serrano, E., Fernandez, N., López-Olvera, J. R., Garel, M., Santos, J., Espunyes, J., Parra-Aguado, M. A., Ramanzin, M., Fernández-Aguilar, X., Colom-Cadena, A., Marco, I., Lavín, S., Bartolomé, J. & Albanell, E. 2017. Predicting herbivore faecal nitrogen using a multispecies near-infrared reflectance spectroscopy calibration. PLoS ONE, 12, e0176635  Download pdf file

48.- Richard, Q., Toïgo, C., Appolinaire, J., Loison, A., & Garel, M. 2017. From gestation to weaning: Combining robust design and multi-event models unveils cost of lactation in a large herbivore. Journal of Animal Ecology, 86, 1497-1509  Download pdf file

47.- Oliver-Guimera, A., Martinez-Carrasco, C., Tvarijonativiciute, A., Rocío Ruiz de Ybáñez, M., Martínez-Guijosa, J., Ramón López-Olvera, J., Fernández-Aguilar, X., Colom-Cadena, A., Mentaberre, G., Velarde-Nieto, R., Gassó-Garcia, D., Garel, M., Lavín, S., Rossi, L. & Serrano, E. 2017. The physiological cost of male-biased parasitism in a nearly monomorphic mammal. Parasite & Vectors, 10, 200  Download pdf file

46.- Marchand, P., Garel, M., Bourgoin, G., Duparc, A., Dubray, D., Maillard, D. & Loison, A. 2017. Combining familiarity and landscape features helps break down the barriers between movements and home ranges in a non-territorial large herbivore. Journal of Animal Ecology, 86, 371--383   Download pdf file

45.- Hamel, S., Gaillard, J.-M., Yoccoz, N.G., Albon, S., Côté, S.D., Craine, J. M., Festa-Bianchet, M., Garel, M., Lee, P., Moss, C., Nussey, D. H., Pelletier, F., Stien, A. & Tveraa, T. 2016. Cohort variation in individual body mass dissipates with age in large herbivores. Ecological Monographs, in press   Download pdf file

44.- Forsyth, D. M., Garel, M. & Mcleod, S. R. 2016. Estimating age and age class of harvested hog deer from eye lens mass using frequentist and Bayesian methods. Wildlife Biology, 22, 137-143   Download pdf file

43.- Kourkgy, C., Garel, M., Appolinaire, J., Loison, A. & Toïgo, C. 2016. Onset of autumn shapes the timing of birth in Pyrenean chamois more than onset of spring. Journal of Animal Ecology, 85, 581-590   Download pdf file

42.- Villamuelas, M., Fernández, N., Albanell, E., Gálvez-Cerón, A., Bartolomé, J., Mentaberre, G., López-Olvera, J. R., Fernández-Aguilar, X., Colom-Cadena, A., López-Martín, J. M., Garel, M., Marco, I. & Serrano, E. 2016. The Enhanced Vegetation Index (EVI) as a proxy for diet quality and composition in a mountain ungulate. Ecological Indicators, 61, 658-666  Download pdf file

41.- Serrano, E., Colom-Cadena, A., Gilot-Fromont, E., Garel, M., Cabezón, O., Velarde, R., Fernández-Sirera, L., Fernández-Aguilar, X., Rosell, R., Lavín, S. & Marco, I. 2015. Border Disease Virus: an exceptional driver of chamois populations among other threats. Frontiers in Microbiology, 6, 1307   Download pdf file

40.- Bleu, J., Herfindal, I., Loison, A., Kwak, A. M. G., Garel, M., Toïgo, C., Rempfler, T., Filli, F. & Sæther, B. E. S. 2015. Age-specific survival and annual variation in survival of female chamois differ between populations. Oecologia, 179, 1091-1098  Download pdf file

39.- Beaunée, G., Gilot-Fromont, E., Garel, M. & Ezanno, P. 2015. A novel epidemiological model to better understand and predict the observed seasonal spread of Pestivirus in Pyrenean chamois populations. Veterinary Research, 46, 86   Download pdf file

38.- Guerrini, M., Forcina, G., Panayides, R., Lorenzini, R., Garel, M., Anayiotos, P., Kassinis, N. & Barbanera, F. 2015. Molecular DNA identity of the mouflon of Cyprus (Ovis orientalis ophion, Bovidae): Near Eastern origin and divergence from Western Mediterranean conspecific populations. Systematics & Biodiversity, 13, 472-483,   Download pdf file

37.- Martínez-Guijosa, J., Martínez-Carrasco, C., López-Olvera, J. R., Fernández-Aguilar, X., Colom-Cadena, A., Cabezón, O., Mentaberre, G., Ferrer, D., Velarde, R., Gassó, D., Garel, M., Rossi, L., Lavín, S. & Serrano, E. 2015. Male-biased gastrointestinal parasitism in a monomorphic mountain ungulate. Parasites & Vectors, 8, 165, DOI 10.1186/s13071-015-0774-9   Download pdf file

36.- Gálvez-Cerón, A., Gassó, D., López-Olvera, J. R., Mentaberre, G., Bartolomé, J., Marco, I., Ferrer, D., Rossi, L., Garel, M., Lavín, S., Clauss, M. & Serrano, E. 2015. Gastrointestinal nematodes and dietary fibre: Two factors to consider when using FN for wildlife nutrition monitoring. Ecological Indicators, 52, 161-169   Download pdf file

35.- Marchand, P., Garel, M., Bourgoin, G., Dubray, D., Maillard, D. & Loison, A. 2015. Coupling scale-specific habitat selection and activity reveals sex-specific food/cover trade-offs in a large herbivore. Animal Behaviour, 102, 169-187   Download pdf file

34.- Marchand, P., Garel, M., Bourgoin, G., Dubray, D., Maillard, D. & Loison, A. 2015. Sex-specific adjustments in habitat selection contribute to buffer mouflon against summer conditions. Behavioral Ecology, 26, 472-482   Download pdf file

33.- Marchand, P., Garel, M., Bourgoin, G., Dubray, D., Maillard, D. & Loison, A. 2014. Impacts of tourism and hunting on a large herbivore's spatio-temporal behavior in and around a French protected area. Biological Conservation, 177, 1-11   Download pdf file

32.- Garel, M., Gaillard, J.-M., Delorme, D. & Van Laere, G. 2014. Eruption patterns of permanent front teeth as an indicator of performance in roe deer. Ecological Indicators, 45, 300-307   Download pdf file

31.- Marchand, P., Garel, M., Bourgoin, G., Michel, P., Maillard, D. & Loison, A. 2014. Habitat-related variation in carcass mass of a large herbivore revealed by combining hunting and GPS data. Journal of Wildlife Management, 78, 657-670   Download pdf file

30.- Darmon, G., Bourgoin, G., Marchand, P., Garel, M., Dubray, D., Jullien, J.-M. & Loison, A. 2014. Do ecologically close species shift their daily activities when in sympatry? A test on chamois in presence of mouflon. Biological Journal of the Linnean Society, 111, 621-626   Download pdf file

29.- Benoist, S. , Garel, M., Cugnasse, J.-M. & Blanchard, P. 2013. Human disturbances, habitat characteristics and social environment generate sex-specific responses in vigilance of Mediterranean mouflon. PLoS ONE, 8, e82960   Download pdf file

28.- Douhard, M. , Bonenfant, C., Gaillard, J.-M., Hamann, J.-L., Garel, M., Michallet, J. & Klein, F. 2013. Roaring counts are not suitable for the monitoring of red deer Cervus elaphus population abundance. Wildlife Biology, 19, 94-101   Download pdf file

27.- Marchand, P., Redjadj, C., Garel, M., Cugnasse, J.-M., Maillard, D. & Loison, A. 2013. Are mouflon Ovis gmelini musimon really grazers? A review of variations in diet composition. Mammal Review, 43, 275-291   Download pdf file

26.- Garel, M., Forsyth, D. M., Loison, A., Dubray, D., Jullien, J.-M., Tustin, K. G., Maillard, D. & Gaillard, J.-M. 2011. Age-related male reproductive effort in two mountain ungulates of contrasting sexual size dimorphism. Canadian Journal of Zoology, 89, 929-937   Download pdf file

25.- Bourgoin, G., Garel, M., Blanchard, P., Dubray, D., Maillard, D. & Gaillard, J.-M. 2011. Daily responses of mouflon activity to summer climatic conditions. Canadian Journal of Zoology, 89, 765-773   Download pdf file

24.- Garel, M., Gaillard, J.-M., Jullien, J.-M., Dubray, D., Maillard, D. & Loison, A. 2011. Population abundance and early spring conditions determine variation in body mass of juvenile chamois. Journal of Mammalogy, 95, 1112-1117   Download pdf file

23.- Cazau, M., Garel, M. & Maillard, D. 2011. Responses of heather moorland and mediterranean mouflon foraging to prescribed-burning and cutting. Journal of Wildlife Management, 75, 967-972   Download pdf file

22.- Hibert, F., Maillard, D., Fritz, H., Garel, M., Abdou, H. N. & Winterton, P. 2010. Ageing of ungulate pellets in semi-arid landscapes: how the shade of colour can refine pellet-group counts. European Journal of Wildlife Research, 57, 495-503   Download pdf file

21.- Garel, M., Gaillard, J.-M., Chevrier, T., Michallet, J., Delorme, D. & Van Laere, G. 2010. Testing reliability of body size measurements using hind foot length in roe deer. Journal of Wildlife Management, 74, 1382-1386   Download pdf file

20.- Garel, M., Bonenfant, C., Hamann, J.-L., Klein, F. & Gaillard, J.-M. 2010. Are abundance indices derived from spotlight counts reliable to monitor red deer Cervus elaphus populations? Wildlife Biology, 16, 77-84   Download pdf file

19.- Bourgoin, G., Garel, M., Dubray, D., Maillard, D., & Gaillard, J.-M. 2009. What determines global positioning system fix success when monitoring free-ranging mouflon? European Journal of Wildlife Research, 55, 603-613   Download pdf file

18.- Hamel, S., Garel, M., Festa-Bianchet, M., Gaillard, J.-M. & Côté, S. D. 2009. Spring NDVI correlates with fecal crude protein in mountain ungulates. Journal of Applied Ecology, 46, 582-589   Download pdf file

17.- Santin-Janin, H., Garel, M., Chapuis, J.-L. & Pontier, D. 2009. Assessing the performance of NDVI as a proxy for plant biomass using non-linear models: a case study on the Kerguelen archipelago. Polar Biology, 32, 861-871   Download pdf file

16.- Bonenfant, C., Gaillard, J.-M., Coulson, T. H., Festa-Bianchet, M., Loison, A., Garel, M., Loe, L. E., Blanchard, P., Pettorelli, N., Owen-Smith, N., Du Toit, J. & Duncan, P. 2009. Empirical evidences of density-dependence in populations of large herbivores. Advances in Ecological Research, 41, 313-357   Download pdf file

15.- Garel, M., Loison, A., Jullien, J.-M., Dubray, D., Maillard, D. & Gaillard J.-M. 2009. Sex-specific growth in alpine chamois. Journal of Mammalogy, 90, 954-960   Download pdf file

14.- Garel, M., Solberg, E. J., Sæther, B.-E., Grotan, V., Tufto, J. & Heim M. 2009. Age, size and spatio-temporal variation in ovulation patterns of a seasonal breeder, the Norwegian moose (Alces alces). American Naturalist, 173, 89-104   Download pdf file

13.- Bonenfant, C., Pelletier, F., Garel, M. & Bergeron, P. 2009. Age-dependent relationship between horn growth and survival in wild sheep. Journal of Animal Ecology, 78, 161-171   Download pdf file

12.- Bourgoin, G., Garel, M., Van Moorter, B., Dubray, D., Maillard, D., Marty, E. & Gaillard, J.-M. 2008. Determinants of seasonal variation in activity patterns of mouflon. Canadian Journal of Zoology, 86, 1410-1418   Download pdf file

11.- Solberg, E. J., Garel, M., Heim, M., Grotan, V. & Sæther, B.-E. 2008. Lack of compensatory body growth in a high performance moose Alces alces population. Oecologia, 158, 485-498   Download pdf file

10.- Solberg, E., Heim, M., Grotan, V., Sæther, B.-E. & Garel, M. 2007. Annual variation in maternal age and calving date generates cohort effects in moose (Alces alces) body mass. Oecologia, 154, 259-271   Download pdf file

9.- Garel, M., Cugnasse, J.-M., Maillard, D., Gaillard, J.-M, Hewison, A.J.M. & Dubray, D. 2007. Selective harvesting and habitat loss produce long-term life-history changes in a mouflon population. Ecological Applications, 17, 1607-1618   Download pdf file

8.- Garel, M., Cugnasse, J.-M., Hewison, A. J. M. & Maillard, D. 2006. Errors in age determination of mouflon in the field. Wildlife Society Bulletin, 34, 300-306   Download pdf file

7.- Cugnasse, J.-M. & Garel, M. 2006. Capturer les jeunes ongulés à l'aide d'un rapace : un test de faisabilité chez le mouflon. Mammalia, 70, 313-314   Download pdf file

6.- Garel, M., Solberg, E. J., Sæther, B.-E., Herfindal, I. & Hogda, K.-A. 2006. The length of growing season and adult sex-ratio shape sexual size dimorphism in moose. Ecology, 87, 745-758   Download pdf file

5.- Garel, M., Cugnasse, J.-M., Gaillard, J.-M., Loison, A., Gibert, P., Douvre, P. & Dubray, D. 2005. Reproductive output of female mouflon (Ovis gmelini musimon x Ovis sp.) : a comparative analysis. Journal of Zoology, 266, 65-71   Download pdf file

4.- Garel, M., Cugnasse, J.-M., Gaillard, J.-M., Loison, A., Santosa, Y. & Maublanc, M.-L. 2005. Effect of observer experience on the monitoring of a mouflon population. Acta Theriologica, 50, 109-114   Download pdf file

3.- Garel, M., Cugnasse, J.-M., Loison, A., Gaillard, J.-M., Vuiton, C. & Maillard, D. 2005. Monitoring the abundance of mouflon in south France. European Journal of Wildlife Research, 51, 69-76   Download pdf file

2.- Garel, M., Cugnasse, J.-M. & Maillard, D. 2005. Reliability of mouflon aging using morphological characteristics from observations in the field. Wildlife Biology, 11, 229-236   Download pdf file

1.- Garel, M., Loison, A., Gaillard, J.-M., Cugnasse, J.-M. & Maillard, D. 2004. The effects of a severe drought on mouflon lamb survival. Proceedings of the Royal Society of London Biological Sciences (Suppl.), 271, S471-S473   Download pdf file


Garel, M. 2006. Conséquences de la chasse et des contraintes environnementales sur la démographie des populations d'ongulés. L'exemple du mouflon méditerranéen (Ovis gmelini musimon x Ovis sp.) en France et de l'élan (Alces alces) en Norvège.   Download pdf file


Several vertebrate populations have to face indirect and direct human pressures. Our aim here is to show how these pressures influence ungulate population dynamics beyond environmental factors commonly accounted for, such as habitat quality or climate. We present two case studies : Mediterranean mouflon (Ovis gmelini musimon x Ovis sp.) in France and moose (Alces alces) in Norway. We analyzed a long term study (30 years) of a mouflon population located in southern France (Hérault). Our study on moose was based on a transversal and comparative analysis of several populations inhabiting contrasted environments in Norway along a latitudinal gradient.

Our study on mouflon highlighted the advantages and limits of commonly used survey methods for ungulate population monitoring. We then showed that the dynamics of this population was influenced by (1) habitat closure, resulting from the reduction of pastoral activity, leading to the use of sub-optimal resources by mouflon, (2) selective hunting on trophy males, (3) the recent history of mouflon from which our population originated, (4) summer drought. These processes influence the dynamics of the mouflon through (1) a decrease in phenotypic quality as measured by body mass and trophy size (processes 1 and 2), (2) variation in reproductive output of females (processes 3 and 4), (3) and variation in lamb summer survival (process 4).

In moose, we showed that sexual size dimorphism (SSD) increases with increasing environmental seasonality, probably caused by higher resource quality when the growing season was short and intense. Selective hunting against males also shapes SSD by de- creasing the proportion of adult males in the population. A low proportion of adult males could lead to a higher proportion of young males involved in reproduction, which in turn may have a negative impact on their body growth.

Based on two cases studies at different spatial scales, our results highlighted the ex- istence of specific biological characteristics in harvested populations. In addition to the environmental constraints under which these populations evolve, we demonstrated that man can have a strong impact on their demography.