, CITED

B. Agnvall, M. Jöngren, E. Strandberg, and P. Jensen, Heritability and Genetic Correlations of Fear-Related Behaviour in Red Junglefowl???Possible Implications for Early Domestication, PLoS ONE, vol.92, issue.4, p.35162, 2012.
DOI : 10.1371/journal.pone.0035162.t005

C. E. Allen, P. Beldade, B. J. Zwaan, and P. M. Brakefield, Differences in the selection response of serially repeated color pattern characters: Standing variation, development, and evolution, BMC Evolutionary Biology, vol.8, issue.1, p.94, 2008.
DOI : 10.1186/1471-2148-8-94

Y. Aso, D. Sitaraman, T. Ichinose, K. R. Kaun, K. Vogt et al., Mushroom body output neurons encode valence and guide memorybased action selection in Drosophila. eLife 3:e04580, Avargù es-Weber, A., and M. Giurfa. 2013. Conceptual learning by miniature brains, p.20131907, 2014.

J. M. Baeder and B. H. King, Associative Learning of Color by Males of the Parasitoid Wasp Nasonia vitripennis (Hymenoptera: Pteromalidae), Journal of Insect Behavior, vol.17, issue.2, 2004.
DOI : 10.1023/B:JOIR.0000028570.93123.dc

, J. Insect Behav, vol.17, pp.201-213

R. D. Barrett and D. Schluter, Adaptation from standing genetic variation, Trends in Ecology & Evolution, vol.23, issue.1, pp.38-44, 2008.
DOI : 10.1016/j.tree.2007.09.008

P. Beldade, K. Koops, and P. M. Brakefield, Developmental constraints versus flexibility in morphological evolution, Nature, vol.22, issue.6883, pp.844-847, 2002.
DOI : 10.1002/1521-1878(200012)22:12<1075::AID-BIES5>3.0.CO;2-M

R. A. Boulton and D. M. Shuker, The costs and benefits of multiple mating in a mostly monandrous wasp, Evolution, vol.264, issue.4, pp.939-949, 2015.
DOI : 10.1098/rspb.1997.0010

C. Brandes and R. Menzel, Common mechanisms in proboscis extension conditioning and visual learning revealed by genetic selection in honeybees (Apis mellifera capensis), Journal of Comparative Physiology A, vol.166, issue.4, pp.545-552, 1990.
DOI : 10.1007/BF00192025

M. K. Burke, J. P. Dunham, P. Shahrestani, K. R. Thornton, M. R. Rose et al., Genome-wide analysis of a long-term evolution experiment with Drosophila, Nature, vol.10, issue.7315, p.587, 2010.
DOI : 10.1554/05-273.1

J. G. Burns, J. Foucaud, and F. Mery, Costs of memory: lessons from 'mini' brains, Proceedings of the Royal Society B: Biological Sciences, vol.278, issue.1702, pp.923-929, 2011.
DOI : 10.1098/rspb.2010.1337

A. Cardona, S. Saalfeld, J. Schindelin, I. Arganda-carreras, S. Preibisch et al., TrakEM2 Software for Neural Circuit Reconstruction, PLoS ONE, vol.25, issue.6, p.38011, 2012.
DOI : 10.1371/journal.pone.0038011.s010

J. Cho, N. Yu, J. Choi, S. Sim, S. J. Kang et al.,

D. I. Kim, V. N. Choi, and . Kim, Multiple repressive mechanisms in the hippocampus during memory formation, Science, vol.350, pp.82-87, 2015.

M. Costa-mattioli, W. S. Sossin, E. Klann, and N. Sonenberg, Translational Control of Long-Lasting Synaptic Plasticity and Memory, Neuron, vol.61, issue.1, pp.10-26, 2009.
DOI : 10.1016/j.neuron.2008.10.055

J. Dubnau and T. Tully, : New Insights for Learning and Memory, Annual Review of Neuroscience, vol.21, issue.1, pp.407-444, 1998.
DOI : 10.1146/annurev.neuro.21.1.407

A. S. Dunlap and D. W. Stephens, Experimental evolution of prepared learning, Proc. Natl. Acad. Sci. USA, pp.11750-11755, 2014.
DOI : 10.1037/a0017759

J. Ellers and M. Liefting, Extending the integrated phenotype: covariance and correlation in plasticity of behavioural traits, Current Opinion in Insect Science, vol.9, pp.31-35, 2015.
DOI : 10.1016/j.cois.2015.05.013

S. E. Fahrbach, S. M. Farris, J. P. Sullivan, and G. Robinson, Limits on volume changes in the mushroom bodies of the honey bee brain, Journal of Neurobiology, vol.26, issue.2, pp.141-151, 2003.
DOI : 10.1002/neu.480260111

M. J. Fitzpatrick, Y. Ben-shahar, H. M. Smid, L. E. Vet, G. E. Robinson et al., Candidate genes for behavioural ecology, Trends in Ecology & Evolution, vol.20, issue.2, pp.96-104, 2005.
DOI : 10.1016/j.tree.2004.11.017

S. Ginsburg and E. Jablonka, The evolution of associative learning: A factor in the Cambrian explosion, Journal of Theoretical Biology, vol.266, issue.1, pp.11-20, 2010.
DOI : 10.1016/j.jtbi.2010.06.017

M. Giurfa, Cognition with few neurons: higher-order learning in insects, Trends in Neurosciences, vol.36, issue.5, pp.285-294, 2013.
DOI : 10.1016/j.tins.2012.12.011

W. Gronenberg and G. O. López-riquelme, Multisensory convergence in the mushroom bodies of ants and bees, Acta Biologica Hungarica, vol.55, issue.1-4, pp.31-37, 2004.
DOI : 10.1556/ABiol.55.2004.1-4.5

J. Groothuis and H. M. Smid, <b><i>Nasonia</i></b> Parasitic Wasps Escape from Haller's Rule by Diphasic, Partially Isometric Brain-Body Size Scaling and Selective Neuropil Adaptations, Brain, Behavior and Evolution, vol.90, issue.3, pp.243-254, 2017.
DOI : 10.1086/405402
URL : http://www.karger.com/Article/Pdf/480421

A. Haverkamp and H. M. Smid, Octopamine-like immunoreactive neurons in the brain and subesophageal ganglion of the parasitic wasps Nasonia vitripennis and N. giraulti, Cell and Tissue Research, vol.23, issue.2, pp.313-329, 2014.
DOI : 10.1016/j.cub.2013.09.056

M. L. Head, C. A. Hinde, A. J. Moore, and N. J. Royle, Correlated evolution in parental care in females but not males in response to selection on paternity assurance behaviour, Ecology Letters, vol.105, issue.7, pp.803-810, 2014.
DOI : 10.1073/pnas.0803146105

S. D. Healy, I. E. Bacon, O. Haggis, A. P. Harris, and L. A. Kelley, Explanations for variation in cognitive ability: Behavioural ecology meets comparative cognition, Behavioural Processes, vol.80, issue.3, pp.288-294, 2009.
DOI : 10.1016/j.beproc.2008.10.002

C. L. Hitchcock and D. F. Sherry, Long-term memory for cache sites in the black-capped chickadee, Animal Behaviour, vol.40, issue.4, pp.701-712, 1990.
DOI : 10.1016/S0003-3472(05)80699-2

K. M. Hoedjes, H. M. Smid, E. G. Schijlen, L. E. Vet, and J. J. Van-vugt, Learning-induced gene expression in the heads of two Nasonia species that differ in long-term memory formation, BMC Genomics, vol.29, issue.1, pp.1-13, 2015.
DOI : 10.1038/nbt.1883

K. M. Hoedjes, H. M. Smid, L. E. Vet, and J. H. Werren, Introgression study reveals two quantitative trait loci involved in interspecific variation in memory retention among Nasonia wasp species, Heredity, vol.2009, issue.6, pp.542-550, 2014.
DOI : 10.1126/science.1178028
URL : http://www.nature.com/hdy/journal/v113/n6/pdf/hdy201466a.pdf

K. M. Hoedjes, J. L. Steidle, J. H. Werren, L. E. Vet, and H. M. Smid, parasitic wasps, Genes, Brain and Behavior, vol.42, issue.7, pp.879-887, 2012.
DOI : 10.1086/405402

B. Hollis and T. Kawecki, Male cognitive performance declines in the absence of sexual selection, Proceedings of the Royal Society B: Biological Sciences, vol.17, issue.12, p.20132873, 2014.
DOI : 10.1016/S0168-9525(01)02526-4

K. Honjo and K. Furukubo-tokunaga, Distinctive Neuronal Networks and Biochemical Pathways for Appetitive and Aversive Memory in Drosophila Larvae, Journal of Neuroscience, vol.29, issue.3, pp.852-862, 2009.
DOI : 10.1523/JNEUROSCI.1315-08.2009
URL : http://www.jneurosci.org/content/jneuro/29/3/852.full.pdf

E. R. Kandel, The Molecular Biology of Memory Storage: A Dialogue Between Genes and Synapses, Science, vol.294, issue.5544, pp.1030-1038, 2001.
DOI : 10.1126/science.1067020

A. Kotrschal, B. Rogell, A. Bundsen, B. Svensson, S. Zajitschek et al., Artificial Selection on Relative Brain Size in the Guppy Reveals Costs and Benefits of Evolving a Larger Brain, Current Biology, vol.23, issue.2, pp.168-171, 2013.
DOI : 10.1016/j.cub.2012.11.058

M. A. Krause, Evolutionary perspectives on learning: conceptual and methodological issues in the study of adaptive specializations, Animal Cognition, vol.9, issue.2, pp.807-820, 2015.
DOI : 10.1037/h0058825

E. M. Macphail and J. J. Bolhuis, The evolution of intelligence: adaptive specializations versus general process, Biological Reviews of the Cambridge Philosophical Society, vol.49, issue.3, pp.341-364, 2001.
DOI : 10.1037/0097-7403.12.4.412

C. Margulies, T. Tully, and J. Dubnau, Deconstructing Memory in Drosophila, Current Biology, vol.15, issue.17, pp.700-713, 2005.
DOI : 10.1016/j.cub.2005.08.024
URL : https://doi.org/10.1016/j.cub.2005.08.024

J. A. Mather, Navigation by spatial memory and use of visual landmarks in octopuses, Journal of Comparative Physiology A, vol.35, issue.4, pp.491-497, 1991.
DOI : 10.1007/BF00199609

R. Menzel, The insect mushroom body, an experience-dependent recoding device, Journal of Physiology-Paris, vol.108, issue.2-3, pp.84-95, 2014.
DOI : 10.1016/j.jphysparis.2014.07.004

F. Mery and T. J. Kawecki, Experimental evolution of learning ability in fruit flies, Proc. Natl. Acad. Sci. USA 99, pp.14274-14279, 2002.
DOI : 10.1038/nm0402-349

?. , A fitness cost of learning ability in Drosophila melanogaster, Proc. R Soc. Lond. B Biol. Sci, vol.270, pp.2465-2469, 2003.

F. Mery, J. Pont, T. Preat, and T. J. Kawecki, Physiological and Biochemical Zoology, vol.80, issue.4, pp.399-405, 2007.
DOI : 10.1086/518014

M. Mizunami and Y. Matsumoto, Roles of Octopamine and Dopamine Neurons for Mediating Appetitive and Aversive Signals in Pavlovian Conditioning in Crickets, Frontiers in Physiology, vol.412, p.1027, 2017.
DOI : 10.1038/35083500

H. Muller and L. Chittka, Diff??rences Interindividuelles en termes d'Apprentissage des Couleurs, Formes et Odeurs chez le Bourdon (Hymenoptera: Apidae: Bombus terrestris), Entomologia Generalis, vol.34, issue.1-2, pp.1-8, 2012.
DOI : 10.1127/entom.gen/34/2012/1

R. A. Murphy, E. Mondragon, and V. A. Murphy, Rule Learning by Rats, Science, vol.274, issue.5294, pp.1849-1851, 2008.
DOI : 10.1126/science.274.5294.1926
URL : http://openaccess.city.ac.uk/22060/1/Murphy-Mondragon-Murphy_2008.pdf

Y. Nakatani, Y. Matsumoto, Y. Mori, D. Hirashima, H. Nishino et al., Why the carrot is more effective than the stick: Different dynamics of punishment memory and reward memory and its possible biological basis, Neurobiology of Learning and Memory, vol.92, issue.3, pp.370-380, 2009.
DOI : 10.1016/j.nlm.2009.05.003

L. C. Odling-smee, J. W. Boughman, and V. A. Braithwaite, Sympatric species of threespine stickleback differ in their performance in a spatial learning task, Behavioral Ecology and Sociobiology, vol.4, issue.12, pp.1935-1945, 2008.
DOI : 10.1046/j.1467-2979.2003.00125.x

S. E. Oliai and B. H. King, Associative learning in response to color in the parasitoid wasp Nasonia vitripennis (Hymenoptera: Pteromalidae), 2000.

, J. Insect Behav, vol.13, pp.55-69

D. R. Papaj and L. E. Vet, Odor learning and foraging success in the parasitoid,Leptopilina heterotoma, Journal of Chemical Ecology, vol.35, issue.11, pp.3137-3150, 1990.
DOI : 10.1007/BF00979616

M. R. Papini, Pattern and process in the evolution of learning., Psychological Review, vol.109, issue.1, pp.186-201, 2002.
DOI : 10.1037/0033-295X.109.1.186

J. L. Pitman, S. Dasgupta, M. J. Krashes, B. Leung, P. N. Perrat et al., There are many ways to train a fly, Fly, vol.3, issue.1, pp.3-9, 2009.
DOI : 10.4161/fly.3.1.7726

R. Team, R: A language and environment for statistical computing . R Foundation for Statistical Computing, 2018.

N. Raine, T. Ings, O. Ramos-rodriguez, and L. Chittka, Unterschiede im Lernverhalten zwischen Kolonien einer freilebenden Britischen Hummelpopulation (Hymenoptera: Apidae: Bombus terrestris audax), Entomologia Generalis, vol.28, issue.4, pp.241-256, 2006.
DOI : 10.1127/entom.gen/28/2006/241

N. E. Raine and L. Chittka, The correlation of learning speed and natural foraging success in bumble-bees, Proceedings of the Royal Society B: Biological Sciences, vol.34, issue.19, pp.803-808, 2008.
DOI : 10.1242/jeb.00570

R. A. Rescorla, Behavioral Studies of Pavlovian Conditioning, Annual Review of Neuroscience, vol.11, issue.1, pp.329-352, 1988.
DOI : 10.1146/annurev.ne.11.030188.001553

B. D. Roitberg, J. Sircom, C. A. Roitberg, J. J. Van-alphen, and M. Mangel, Life expectancy and reproduction, Nature, vol.364, issue.6433, p.108, 1993.
DOI : 10.1038/364108a0

C. Rowe and S. D. Healy, Measuring variation in cognition, Behavioral Ecology, vol.16, issue.7, pp.1287-1292, 2014.
DOI : 10.1007/s10071-012-0559-x

E. Santini, T. N. Huynh, and E. Klann, Mechanisms of Translation Control Underlying Long-Lasting Synaptic Plasticity and the Consolidation of Long-Term Memory, Progress in molecular biology and translational science, pp.131-167, 2014.
DOI : 10.1016/B978-0-12-420170-5.00005-2

R. Scheiner, J. Erber, and R. Page-jr, Tactile learning and the individual evaluation of the reward in honey bees ( Apis mellifera L.), Journal of Comparative Physiology A: Sensory, Neural, and Behavioral Physiology, vol.185, issue.1, pp.1-10, 1999.
DOI : 10.1007/s003590050360

J. Schindelin, I. Arganda-carreras, E. Frise, V. Kaynig, M. Longair et al., Fiji: an open-source platform for biological-image analysis, Nature Methods, vol.27, issue.7, pp.676-682, 2012.
DOI : 10.1093/bioinformatics/btr390

M. J. Sheehan and E. A. Tibbetts, Specialized Face Learning Is Associated with Individual Recognition in Paper Wasps, Science, vol.3, issue.3, pp.1272-1275, 2011.
DOI : 10.1007/s100710000075

S. J. Shettleworth, Cognition, evolution, and behavior, 2010.

K. K. Siwicki and L. Ladewski, Associative learning and memory in Drosophila: beyond olfactory conditioning, Behavioural Processes, vol.64, issue.2, pp.225-238, 2003.
DOI : 10.1016/S0376-6357(03)00137-2

E. C. Snell-rood, An overview of the evolutionary causes and consequences of behavioural plasticity, Animal Behaviour, vol.85, issue.5, pp.1004-1011, 2013.
DOI : 10.1016/j.anbehav.2012.12.031

E. C. Snell-rood, D. R. Papaj, and W. Gronenberg, Brain Size: A Global or Induced Cost of Learning?, Brain, Behavior and Evolution, vol.73, issue.2, pp.111-128, 2009.
DOI : 10.1159/000213647

K. Steck, B. S. Hansson, and M. Knaden, Desert ants benefit from combining visual and olfactory landmarks, Journal of Experimental Biology, vol.214, issue.8, pp.1307-1312, 2011.
DOI : 10.1242/jeb.053579
URL : http://jeb.biologists.org/content/214/8/1307.full.pdf

S. Steinberg, M. Dicke, L. E. Vet, and R. Wanningen, Response of the braconid parasitoid Cotesia (=Apanteles) glomerata to volatile infochemicals: effects of bioassay set-up, parasitoid age and experience and barometric flux, Entomologia Experimentalis et Applicata, vol.2, issue.2, pp.163-175, 1992.
DOI : 10.1146/annurev.en.21.010176.000545

D. W. Stephens, Learning and Behavioral Ecology: Incomplete Information and Environmental Predictability, Insect Learning, pp.195-218, 1993.
DOI : 10.1007/978-1-4615-2814-2_8

T. Tully, Discovery of genes involved with learning and memory: An experimental synthesis of Hirschian and Benzerian perspectives, Proc. Natl. Acad. Sci. USA 93, pp.13460-13467, 1996.
DOI : 10.1016/0896-6273(95)90174-4

T. Tully, T. Preat, S. Boynton, and M. D. Vecchio, Genetic dissection of consolidated memory in Drosophila, Cell, vol.79, issue.1, pp.35-47, 1994.
DOI : 10.1016/0092-8674(94)90398-0

T. Tully and W. G. Quinn, Classical conditioning and retention in normal and mutantDrosophila melanogaster, Journal of Comparative Physiology A, vol.17, issue.2, pp.263-277, 1985.
DOI : 10.1093/icb/17.3.707

L. Van-de-zande, S. Ferber, A. De-haan, L. W. Beukeboom, J. Van-heerwaarden et al., outbred laboratory population for genetic analysis, Molecular Ecology Resources, vol.169, issue.3, pp.578-587, 2014.
DOI : 10.1534/genetics.104.032573

E. Van-der-woude and H. M. Smid, parasitic wasps, Journal of Comparative Neurology, vol.506, issue.6, pp.1876-1891, 2015.
DOI : 10.1002/cne.21548

J. J. Van-vugt, K. M. Hoedjes, H. C. Van-de-geest, E. W. Schijlen, L. E. Vet et al., Differentially expressed genes linked to natural variation in long-term memory formation in Cotesia parasitic wasps, Frontiers in Behavioral Neuroscience, vol.20, p.255, 2015.
DOI : 10.1101/lm.026575.112

K. Vogt, C. Schnaitmann, K. V. Dylla, S. Knapek, Y. Aso et al., Author response, eLife, vol.33, p.2395, 2014.
DOI : 10.7554/eLife.02395.019

S. Waddell, Dopamine reveals neural circuit mechanisms of fly memory, Trends in Neurosciences, vol.33, issue.10, pp.457-464, 2010.
DOI : 10.1016/j.tins.2010.07.001

?. , Neural plasticity: dopamine tunes the mushroom body output network, Curr. Biol, vol.26, pp.109-112, 2016.

J. H. Werren and D. W. Loehlin, The Parasitoid Wasp Nasonia: An Emerging Model System with Haploid Male Genetics, Cold Spring Harbor Protocols, vol.2009, issue.10, pp.1-11, 2009.
DOI : 10.1101/pdb.emo134
URL : http://europepmc.org/articles/pmc2916733?pdf=render

M. K. Zwoinska, M. I. Lind, M. Cortazar-chinarro, M. Ramsden, and A. A. Maklakov, Selection on learning performance results in the correlated evolution of sexual dimorphism in life history, Evolution, vol.48, issue.2, pp.342-357, 2016.
DOI : 10.1016/j.exger.2013.09.008