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24, chemin de Borde Rouge –Auzeville – CS52627
31326 Castanet Tolosan CEDEX - France

Dernière mise à jour : Mai 2018

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Plantes et Système de cultures Horticoles

Zone de texte éditable et éditée et rééditée

Pierre Franck

Pierre Franck

Dept. Santé des Plantes & Environnement (SPE)

Lab. Plantes et Systèmes de culture Horticoles (PSH)
Team :Conservation Biological Control (CBC)


ORCID : 0000-0002-1904-8325

UR1115 Plantes et Systèmes de culture Horticoles, INRA

Domaine Saint Paul, Site Agroparc,
F-84914 Avignon Cedex 9, France

Tél: (33) 432 72 26 73

Email :



2006-       : Researcher (CR1), Plantes & Systèmes de culture Horticoles, INRA, Avignon.

2002-2005 : Researcher (CR2), Abeille & Environnement, INRA, Avignon.

2000-2001 : Postdoc, Behavioral and Genetics of Social Insect, University of Sydney,

1996-1999 : PhD thesis in Agronomical Sciences, speciality Population & Ecology, Centre de Biologie et de Gestion des Populations, Montpellier.

1995-1996 : National Service, Institut de Médecine Tropicale du Service de Santé des Armées, Marseille.

1994-1995 : Master in Populations Genetics & Evolution, CNRS, Gif-sur-Yvette.


Understanding insect population dynamics and biological interactions in agricultural landscapes is currently central in my researches. I use both demo- and landscape-genetics approaches to shed light on population dynamics. I conduct conjointly researches in community ecology to detect and to analyse trophic interactions. These researches focus on the understanding of ecological processes in apple orchards to enhance moth and aphid pest controls by their natural enemies. The finalized aim of these researches is to develop ecological engineering strategies to improve ecosystem services provided by farmland biodiversity to agriculture and their sustainability.

My phD and post-doctoral researches dealt with questions in evolutionary genetics, notably about the phylogeography and the reproductive biology of social bees.

Key word: population dynamics; population genetics; evolutionary genetics; landscape ecology; community ecology; agro-ecology; conservation biological control; insecticide resistance; Integrative Pest Management;



FRAMEwork (2020-2026) – Management of agricultural landscapes to enhance biodiversity and value chains in territories (H2020, RIA, Graham Begg coord.).

PREPARE (2019-2022) – Territorial levers to reduce pesticide risks and uses: understanding and prediction of landscape agronomic practices on pest regulation (AFB, coord. Sandrine Petit)

Api-Tree(2018-2020) – Strategies based on agrosystemic approaches to control apple pests. (C-IPM, coordination: Aude Alaphilippe).

Agropolis Fondation CAPES project (2013-2017) « Orchard, semi-natural and natural habitat characteristics and agricultural practices that affect the parasitism of the fruit fly Anastrepha fraterculus in apple orchards of the Parana region. A basis for biological control » coordination Lino-Bittencourt Monteiro and Claire Lavigne; partnership: INRA, University of Parana, University Federal of Lavras).

PEERLESS project (2013-2017) – Predictive Ecological Engineering for Landscape Ecosystem Services and Sustainability” (ANR agrobiosphere; coordination Pierre Franck; partenership: INRA, AgroCampusOuest, Université d’Avignon)

SEBIOPAG-PHYTO (2014-2017) « Niveau de régulation biologique et d’usage des produits phytosanitaires le long d’un gradient de pratiques et de paysages : une analyse comparative à partir du réseau SEBIOPAG » (FRB ; coordination Sandrine Petit).

RéPARe (2014-2017) « Régulation Par les Araignées des Ravageurs en vergers » (FRB ; coordination Pierre Franck ; partnership INRA, CTIFL, Sud-Expé, Station expérimental « La Morinière »).

PURE (2011-2015) about « Pest Suppressive Landscapes » (FP7 project; work package coordination Graham Begg, JKI).



In the CBC team : Claire Lavigne, Bertrand Gauffre, Myriam Siegwart, Benoît Sauphanor through PhD and master students co-supervising and several shared research interests…

At Avignon University: Armin Bischoff through projects about the effect of flower on pest biocontrol and student co-supervising.

In the BioSP team at Avignon: Etienne Klein, Julien Papaix, Lionel Roques, Olivier Martin, and Rachid Senoussi through their expertise in spatial statistics and modelling.

At CBGP, Montpellier, with Jean-Marie-Cornuet who supervised my phD, Arnaud Estoup with who I exchanged a supervisor and Jean-Yves Rasplus for his expertise on Hymenoptera systematics.

From the REX consortium : Denis Bourguet, François Delmotte, Thomas Guillemaud, Xavier Reboud, Corinne Vacher, Anne-Sophie Walker through fruitful discussions about the evolution of pesticide and drug resistance.

From the SEBIOPAG network with notably Manuel Plantagenest, Sandrine Petit, Aude Vialate, Stéphanie Aviron and Benoît Ricci.

At the University of Sydney with Ben Oldroyd though a postdoctoral work on stingless bees

At the Universities of Neuchatel and Lausane, Swissland with Ted Turlings and Nadir Alvarez through Violaine Jourdie phD.

From the University of Talca, Chile with Blas Lavandero, Eduardo Fuentes-Contreras and Claudio Ramirez on several population genetic studies.   

From the University of Thessaly, Greece with John Margaritopoulos on insecticide resistance in Cydia pomonella populations.



Ludivine Laffon (2020-2023) – Trophic interactions among the hoverfly and parasitoid communities, the flower plants and the insect pests in the apple orchards (Co-supervising  with Armin Bischoff & Francoise Lescourret, Avignon Université).

Lucas Etienne (2019-2021) – Impact of landscape structure and farming practices on level of pest infestation and pesticide use in perennial cropping systems (Co-supervising  with Adrien Rusch et Claire Lavigne, Avignon Université).

Vitor Barrile Tomazella (2018-2019) – Morphological and molecular identification of parasitoids involved in parasitism of apple insect pests (Co-supervising with Luís Cláudio Silveira) – Federal University of Lavras, Brazil.

Luan Odorizzi (2015-2016) – Arthropod movement from flower strip border to apple orchard and pest biocontrol (co-supervising with Armin Bishoff and Odair Fernandez) – University of Sao Paolo State, Jaboticabal, Brazil.

Marlice Botelho-Costa (2014-2015) – Predation and parasitism of apple insect pests (Co-supervising with Claire Lavigne and César Carvalho) – Federal University of Lavras, Brazil.

Amandine Juhel (2015-2017) – Population dynamics of Meligethes aeneus and its main parasitoid Tersilochus heterocerus in heterogeneous agricultural landscapes (Co-supervising with Muriel Morison and Corentin Barbut, funding INRA and ANR PEERLESS project) – AgroParisTech.

Laurence Albert (2015-2017) – Agroecological infrastructures effecs on pest regulation in apple cider orchards (Co-supervising with Manuel Plantegenest) – AgroCampus Ouest.

Manon Lefebvre (2014-2016) – Biocontrol of insect pests by spiders in apple orchards (co-supervising with Claire Lavigne, funding CTIFL and FRB RéPARe project).

Mariline Maalouly (2011-2013) – Parasitism of codling moth larvae in orchard landscapes (co-supervising with Claire Lavigne) – Université d’Avignon.

Benoît Ricci (2007-2009) – Spatial dynamics of codling moth populations in the Durance Valley, France (co-supervising with Claire Lavigne) – Université d’Avignon.




Basoalto, A., Ramírez, C.C., Lavandero, B., Devotto, L., Curkovic, T., Franck, P., Fuentes-Contreras, E., 2020. Population Genetic Structure of Codling Moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), in Different Localities and Host Plants in Chile. Insects 11.



Araujo, E.S., Monteiro, L.B., Monteiro, R.S., Nishimura, G., Franck, P., Lavigne, C., 2019. Impact of native forest remnants and wild host plants on the abundance of the South American fruit fly, Anastrepha fraterculus in Brazilian apple orchards. Agriculture, Ecosystems & Environment 275, 93-99.

Dainese, M., et al. 2019. A global synthesis reveals biodiversity-mediated benefits for crop production. Sci Adv 5. eaax0121.

Wan, F. et al. 2019. A chromosome-level genome assembly of Cydia pomonella provides insights into chemical ecology and insecticide resistance. Nature Communications 10, 4237.



Santos, L.A.O., Botelho Costa, M., Lavigne, C., Fernandes, O.A., Bischoff, A., Franck, P., 2018. Influence of the margin vegetation on the conservation of aphid biological control in apple orchards. Journal of Insect Conservation 22, 465-474.

Karp, D.S., Chaplin-Kramer, R., Meehan, T.D., Martin, E.A., DeClerck, F., Grab, H., Gratton, C., Hunt, L., Larsen, A.E., Martínez-Salinas, A., O’Rourke, M.E., Rusch, A., Poveda, K., Jonsson, M., Rosenheim, J.A., Schellhorn, N.A., Tscharntke, T., Wratten, S.D., Zhang, W., Iverson, A.L., et al., 2018. Crop pests and predators exhibit inconsistent responses to surrounding landscape composition. Proceedings of the National Academy of Sciences 115, e7863-e7870.

Juhel, A.S., Barbu, C.M., Valantin-Morison, M., Gauffre, B., Leblois, R., Olivares, J., Franck, P., 2018. Limited genetic structure and demographic expansion of the Brassicogethes aeneus populations in France and in Europe. Pest Management Science.



Albert, L., Franck, P., Gilles, Y., Plantegenest, M., 2017. Impact of agroecological infrastructures on the dynamics of Dysaphis plantaginea Passerini (Hemiptera: Aphididae) and its natural enemies in apple orchards in northwestern France. Environemental Entomology 46, 528-537.

Franck, P., Maalouly-Matar, M., Olivares, J., 2017. Molecular tools for the detection and the identification of Hymenoptera parasitoids in Tortricid fruit pests. International Journal of Molecular Sciences 18, 2031.

Lefebvre, M., Papaïx, J., Mollot, G., Deschodt, P., Lavigne, C., Ricard, J.-M., Mandrin, J.-F., Franck, P., 2017. Bayesian inferences of arthropod movements between hedgerows and orchards. Basic and Applied Ecology 21, 76-84.

Lefebvre, M., Franck, P., Olivares, J., Ricard, J.-M., Mandrin, J.-F., Lavigne, C., 2017. Spider predation on rosy apple aphid in conventional, organic and insecticide-free orchards and its impact on aphid populations. Biological Control 104, 57-65.

Juhel, A.S., Barbu, C.M., Franck, P., Roger-Estrade, J., Butier, A., Bazot, M., Valantin-Morison, M., 2017. Characterization of the pollen beetle, Brassicogethes aeneus, dispersal from woodlands to winter oilseed rape fields. PLOS ONE 12, e0183878.

Begg, G.S., Cook, S.M., Dye, R., Ferrante, M., Franck, P., Lavigne, C., Lövei, G.L., Mansion-Vaquie, A., Pell, J.K., Petit, S., Quesada, N., Ricci, B., Wratten, S.D. & Birch, A.N.E. (2017) A functional overview of conservation biological control. Crop Protection.


2016 :

REX [Bourget, D., Delmotte, F., Franck, P., Guillemaud, T., Reboud, X., Vacher, C., Walker, A.-S.] (2016) Combining selective pressures to enhance the durability of disease resistance genes. Frontiers in Plant Science, 7, e1916.

Lefebvre, M., Franck, P., Toubon, J.F., Bouvier, J.C. & Lavigne, C. (2016) The impact of landscape composition on the occurrence of a canopy dwelling spider depends on orchard management. Agriculture Ecosystems & Environment, 215, 20-29.

Roques, L., Walker, E., Franck, P., Soubeyrand, S. & Klein, E.K. (2016) Using genetic data to estimate diffusion rates in heterogeneous landscapes. Journal of Mathematical Biology, 73, 397–422.

2015 :

Maalouly, M., Franck, P. & Lavigne, C. (2015) Temporal dynamics of parasitoid assemblages parasitizing the codling moth. Biological Control, 82, 31-39.

2014 :

Martin, O., Ricci, B., Franck, P., Toubon, J.-F., Senoussi, R. & Lavigne, C. (2014) Modelling codling moth damage as a function of aduts monitoring and crop protection: a non-linear generalised mixed model approach with time varying covariates. Journal of Agricultural, Biological, and Environmental Statistics, 19, 419-436.

Fuentes-Contreras, E., Basoalto, E., Franck, P., Lavandero, B. & Ramírez, C.C. (2014) Measuring local genetic variability in populations of codling moth (Lepidoptera: Tortricidae) across an unmanaged and commercial orchard interface. Environmental Entomology, 43, 520-527.

2013 :

Ricci, B., Franck, P., Valantin-Morison, M., Bohan, D.A. & Lavigne, C. (2013) Do species population parameters and landscape characteristics affect the relationship between local population abundance and surrounding habitat amount? Ecological Complexity, 15, 62-70.

Maalouly, M., Franck, P., Bouvier, J.-C., Toubon, J.-F. & Lavigne, C. (2013) Codling moth parasitism is affected by semi-natural habitats and agricultural practices at orchard and landscape levels Agriculture, Ecosystems & Environment, 169, 33-42.

Monteiro, L.B., Lavigne, C., Ricci, B., Franck, P., Toubon, J.-F., Sauphanor, B., 2013 Predation of codling moth eggs is affected by pest management practices at orchard and landscape levels. Agriculture, Ecosystems & Environment, doi:10.1016/j.agee.2011.1010.1012.

REX Consortium [Bourguet, D., Delmotte F., Franck, P. Guillemaud, T., Reboud, X., Vacher, C.], 2013 Heterogeneity of selection and the evolution of resistance. Trends in Ecology and Evolution, online


2012 :

Boreau de Roincé, C., Lavigne, C., Ricard, J.-M., Franck, P., Bouvier, J.-C., Garcin, A., Symondson, W.O.C., 2012. Predation by generalist predators on the codling moth versus a closely-related emerging pest the oriental fruit moth: a molecular analysis. Agricultural and Forest Entomology 14, 260-269.

Franck, P., Siegwart, M., Olivares, J., Toubon, J.-F., Lavigne, C., 2012. Multiple origins of the sodium channel kdr mutations in codling moth populations. PloS ONE 7, e43543.

Margaritopoulos, J.T., Voudouris, C.C., Olivares, J., Sauphanor, B., Mamuris, Z., Tsitsipis, J.A., Franck, P., 2012. Dispersal ability in codling moth: mark–release–recapture experiments and kinship analysis. Agricultural and Forest Entomology, 399-407.

Voudouris, C.C., Franck, P., Olivares, J., Sauphanor, B., Mamuris, Z., Tsitsipis, J.A., Margaritopoulos, J.T., 2012. Comparing the genetic structure of codling moth Cydia pomonella (L.) from Greece and France: long distance gene-flow in a sedentary pest species. Bulletin of Entomological Research 102, 185-198.


2011 :

Franck, P., Ricci, B., Klein, E.K., Olivares, J., Simon, S., Cornuet, J.-M., Lavigne, C., 2011. Genetic inferences about the population dynamics of codling moth females at a local scale. Genetica 139, 949-960.

Guillemaud, T., Blin, A., Simon, S., Morel, K., Franck, P., 2011. Weak spatial and temporal population genetic structure in the rosy apple aphid, Dysaphis plantaginea, in French apple orchards. PloS ONE 6, e21263.

Lavandero, B., Figueroa, C.C., Franck, P., Mendez, A., 2011. Estimating gene flow between refuges and crops: A case study of the biological control of Eriosoma lanigerum by Aphelinus mali in apple orchards. PloS ONE 6, e26694.

Ricci, B.R.B., Franck, P., Bouvier, J.C., Casado, D., Lavigne, C., 2011. Effects of hedgerow characteristics on intra-orchard distribution of larval codling moth. Agriculture Ecosystems & Environment 140, 395-400.

Voudouris, C.C., Sauphanor, B., Franck, P., Reyes, M., Mamuris, Z., Tsitsipis, J.A., Vontas, J., Margaritopoulos, J.T., 2011. Insecticide resistance status of the codling moth Cydia pomonella (Lepidoptera: Tortricidae) from Greece. Pesticide Biochemistry and Physiology 100, 229-238.


2010 :

Fargette, M., Berthier, K., Richaud, M., Lollier, V., Franck, P., Hernandez, A., Frutos, R., 2010. Crosses prior to parthenogenesis explain the current genetic diversity of tropical plant-parasitic Meloidogyne species (Nematoda: Tylenchida). Infection, Genetics and Evolution 10, 807-814.

Franck, P., Timm, A.E., 2010. Population genetic structure of Cydia pomonella: a review and case study comparing spatiotemporal variation. Journal of Applied Entomology 134, 191-200.

Jourdie, V., Alvarez, N., Molina-Ochoa, J., Williams, T., Bergvinson, D., Benrey, B., Turlings, T.C.J., Franck, P., 2010. Population genetic structure of two primary parasitoids of Spodoptera frugiperda (Lepidopotera), Chelonus insularis and Campoletis sonorensis (Hymenoptera): to what extent is the host plant important? Molecular Ecology 19, 2168-2179.

Lavigne, C., Ricci, B., Franck, P., Senoussi, R., 2010. Spatial analyses of ecological count data: A density map comparison approach. Basic and Applied Ecology 11, 734-742.

REX consortium [Bourguet, D., Delmotte F., Franck, P. Guillemaud, T., Reboud, X., Vacher, C.], 2010. The skill and style to model the evolution of resistance to pesticide and drug. Evolutionary Application 3, 375-390.


2009 :

Brazier, C., Fontaine, S., Micoud, A., Mottet, C., Speich, P., Marmeisse, R., Melayah, D., Frayssinet-Tachet, L., Sauphanor, B., Franck, P., 2009. Resistance of the codling moth (Cydia pomonella) to diflubenzuron and to pyrethrinoids: situation in 2007. In : Association Francaise de Protection des Plantes (Eds.), 8ème Conférence Internationale sur les Ravageurs en Agriculture (Montpellier, 22-23 Octobre 2008). AFPP, Alfortville, pp. 457-465.

Cornuet, J.-M., Excoffier, L., Franck, P., Estoup, A., 2009. Bayesian inference under complex evolutionary scenarios using microsatellite markers: multiple divergence and genetic admixture events in the honey bee, Apis mellifera. In: Mahoney, C.L., Springer, D.A. (Eds.), Genetic Diversity. Nova Sciences Publishers, Hauppauge, pp. 229-246.

Franck, P., Lavigne, C., Monteiro, L., Ricci, B., Sauphanor, B., Toubon, J.-F., 2009. Comment contrôler les insectes ravageurs en vergers? L’example du carpocapse. Biofutur 305, 43-45.

Jourdie, V., Alvarez, N., Turlings, T.C.J., Franck, P., 2009. Isolation and characterization of polymorphic microsatellite loci in two primary parasitoids of the noctuid Spodoptera frugiperda: Chelonus insularis and Campoletis sonorensis (Hymenoptera). Molecular Ecology Resources 9, 171-173.

Reyes, M., Franck, P., Olivares, J., Margaritopoulos, J., Knight, A., Sauphanor, B., 2009. Worldwide variability of insecticide resistance mechanisms in the codling moth, Cydia pomonella L. (Lepidoptera: Tortricidae). Bulletin of Entomological Research 99, 359-369.

Ricci, B., P. Franck, J.-F. Toubon, J.-C. Bouvier, B. Sauphanor, Lavigne, C., 2009. The influence of landscape on insect pest dynamics: a case study in southeastern France. Landscape Ecology 24:337-349.


2004-2008 :




Franck, P., M. Reyes, J. Olivares, and B. Sauphanor. 2007. Genetic architecture in codling moth populations: comparison between microsatellite and insecticide resistance markers. Molecular Ecology 16:3554-3564.

Franck, P., E. C. Cameron, G. Good, J.-Y. Rasplus, and B. P. Oldroyd. 2004. Nest architecture and genetic differentiation in a species complex of Australian stingless bees. Molecular Ecology 13:2317-2331.

Franck, P., Guérin, F., Loiseau, A. and Sauphanor, B. (2005) Isolation and characterization of microsatellite loci in the codling moth Cydia pomonella L. (Lepidoptera, Tortricidae). Molecular Ecology Notes, 5, 99-102.

REX consortium [Bourguet, D., Delmotte F., Franck, P. Guillemaud, T., Reboud, X., Vacher, C.]. 2007. Structure of the scientific community modelling the evolution of resistance. PloS ONE 2:e1275.

Reyes, M., Franck, P., Charlemillot, P.-J., Ioriatti, C., Olivares, J., Pasqualini, E., Sauphanor, B., 2007. Diversity of insecticide resistance mechanisms and spectrum in European populations of the Codling moth, Cydia pomonella. Pest Management Science 63, 890–902.

Sauphanor, B., Franck, P., Lasnier, T., Toubon, J.-F., Beslay, D., T., B., Bouvier, J.-C., Renou, M., 2007. Insecticide resistance may enhance the response to a host-plant volatile kairomone for the codling moth, Cydia pomonella (L.). Naturwissenschaften (2007) 94, 449–458.


2001-2004 :

Badino, G., Celebrano, G., Manino, A. , Franck, P. (2004) Allozyme evidence of recent genetic shifts in honey bee populations of Italy. Journal of Apicultural Research, 43, 147-149.

Cameron, E.C., Franck, P. , Oldroyd, B.P. (2004) Genetic structure of nest aggregations and drone congregations of the southeast Asian stingless bee Trigona collina. Molecular Ecology, 13, 2357-2364.

Clarke, K.E., Rinderer, T.E., Franck, P., Quezada-Euán, J.J.G. and Oldroyd, B.P. (2002) The africanization of the honeybees (Apis mellifera L.) of the Yucatan: a study of a massive hybridization event across time. Evolution, 56, 1462-1474.

Green, C.L., Franck, P. and Oldroyd, B.P. (2001) Characterization of microsatellite loci for Trigona carbonaria, a stingless bee endemic to Australia. Molecular Ecology Notes, 1, 89-92.

Franck, P., L. Garnery, A. Loiseau, B. P. Oldroyd, H. R. Hepburn, M. Solignac, and J.-M. Cornuet. 2001. Genetic diversity of the honeybee in Africa: microsatellite and mitochondrial data. Heredity 86:420-430.

Franck, P., M. Solignac, D. Vautrin, J.-M. Cornuet, G. Koeniger, and N. Koeniger. 2002. Sperm competition and last-male precedence in the honeybee. Animal Behaviour 64:503-509.

Franck, P., Cameron, E.C., Good, G., Rasplus, J.-Y. and Oldroyd, B.P. (2004) Nest architecture and genetic differentiation in a species complex of Australian stingless bees. Molecular Ecology, 13, 2317-2331.

Guérin, F., Franck, P., Loiseau, A., Devaux, M. and Le Cam, B. (2004) Isolation of 21 new polymorphic microsatellite loci in the phytopathogenic fungus Venturia inaequalis. Molecular Ecology Notes, 4, 268-270.

Palmer, K., Oldroyd, B.P., Franck, P. and Hadisoesilo, S. (2001) Very high paternity frequency in Apis nigrocincta. Insectes Sociaux, 48, 327-332.


1996-2000 :

Cornuet, J.-M., Aulagnier, S., Lek, S., Franck, P. , Solignac, M. (1996) Classifying individuals among infra-specific taxa using microsatellite data and neural networks. Comptes rendus des séances de l'Académie des sciences. Sciences de la vie., 319, 1167-1177.

Franck, P., Garnery, L., Solignac, M., Cornuet, J.-M. (1998) The origin of West European subspecies of honeybees (Apis mellifera) : new insights from microsatellite and mitochondrial data. Evolution, 52, 1119-1134.

Franck, P., Coussy, H., Le Conte, Y., Solignac, M., Garnery, L. and Cornuet, J.-M. (1999) Microsatellite analysis of sperm usage in honeybee. Insect Molecular Biology, 8, 419-421.

Franck, P., Garnery, L., Celebrano, G., Solignac, M. and Cornuet, J.-M. (2000) Hybrid origins of the Italian honeybees (Apis mellifera ligustica and A. m.sicula). Molecular Ecology, 9, 907-921.

Franck, P., Garnery, L., Koeninger, N., Crewe, R., Lahner, G. and Solignac, M. (2000) Evolution of extreme polyandry: an estimate of mating frequency in two African subspecies, Apis mellifera monticola and A .m. scutellata. Insectes Sociaux, 47, 364-370.

Garnery, L., Franck, P., Baudry, E., Vautrin, D., Cornuet, J.-M. and Solignac, M. (1998) Genetic biodiversity of the West European honeybee (Apis mellifera mellifera and Apis mellifera iberica). I. Mitochondrial DNA. Genetic, Selection and Evolution, 30, 31-47.

Garnery, L., Franck, P., Baudry, E., Vautrin, D., Cornuet, J.-M. and Solignac, M. (1998) Genetic biodiversity of the West European honeybee (Apis mellifera mellifera and Apis mellifera iberica). II. Microsatellite loci. Genetic, Selection and Evolution, 30, 49-74.

Viard, F., Franck, P., Dubois, M.P., Estoup, A. and Jarne, P. (1998) Variation of microsatellite size homoplasy across electromorphs, loci, and populations in three invertebrate species. Journal of Molecular Evolution, 47, 42-51.