Gábor Herczeg

MSc in Applied Zoology (2001, Szent István University)
PhD in Ecology (2006, Eötvös Loránd University)
DSc (2016, Hungarian Academy of Sciences)
Habilitation (2016, Eötvös Loránd University)

Research interests

  • Adaptation and speciation in cave environments.
  • Evolution of behavioural consistency (animal personalities, behavioural syndromes).
  • Adaptive divergence in nine-spined sticklebacks (Pungitius pungitius): the effects of predation and competition.
  • Sexual selection in European green lizards (Lacerta viridis): visual and chemical signalling.
  • Adaptive divergence in common frogs (Rana temporaria) along a latitudinal gradient.


Zsebők, S., Herczeg, G., Blázi, Gy., Laczi, M., Nagy, G., Török, J., Garamszegi, L.z. (2018). Minimum spanning tree as a new, robust repertoire size comparison method: simulation and test on birdsong. Behavioral Ecology and Sociobiology 72:48.
Horváth, G., Martín, J., López, P., Garamszegi, L.Z., Herczeg, G. (2017). Food and vitamin D3 availability affects lizard personalities: an experiment. Behavioural Ecology and Sociobiology 71:27.
Jablonszky, M., Szász, E., Markó, G., Török, J., Herczeg, G., Garamszegi, L.Z. (2017). Escape ability and risk-taking behaviour in a Hungarian population of the collared flycatcher (Ficedula albicollis). Behavioral Ecology and Sociobiology 71:54.
Pérez-Moreno, J.L., Balázs, G., Wilkins, B., Herczeg, G., Bracken-Grissom, H.D. (2017). The role of isolation on contrasting phylogeographic patterns in two cave crustaceans. BMC Evolutionary Biology 17:247.
Horváth, G., Mészáros, B., Urszán, T.J., Bajer, K., Molnár, O., Garamszegi, L.Z., Herczeg, G. (2017). Environment-dependence of behavioural consistency in adult male European green lizards (Lacerta viridis). PLoS ONE 12(11):e0187657.
Kovács, T., Herczeg, G., Hettyey, A. (2017). Responses in the diet composition of the Common frog (Rana temporaria) to the stochastic gradiation of autumnal moth (Epirrita autumnata) larvae. Acta Zoologica Academiae Scientiarum Hungaricae 63:115-122.
Zsebők, S., Herczeg, G., Blázi, Gy., Laczi, M., Nagy, G., Szász, E., Markó, G., Török, J., Garamszegi, L. Z. (2017). Short- and long-term repeatability and pseudo-repeatability of bird song: sensitivity of signals to varying environments. Behavioral Ecology and Sociobiology 71:154.
Li, Z., Guo, B., Yang, J., Herczeg, G., Gonda, A., Balázs, G., Shikano, T., Calboli, F.C.F., Merilä, J. (2017). Deciphering the genomic architecture of the stickleback brain with a novel multi-locus gene-mapping approach. Molecular Ecology 26:1557-1575.
Vaskuti, É., Zsebők, S., Herczeg, G., Blázi, Gy., Laczi, M., Nagy, G., Török, J., Garamszegi, L.Z. (2016). A kulturális evolúció nyomai az örvös légykapó (Ficedula albicollis) énekében. Állattani Közlemények 101:25-41.
Herczeg, G., Ab Ghani, N.I., Merilä, J. (2016). On plasticity of aggression: influence of past and present predation risk, social environment and sex. Behavioral Ecology and Sociobiology 70:179-187.
Horváth, G., Martín, J., López, P., Garamszegi, L.Z., Bertók, P., Herczeg, G. (2016). Blood parasite infection intensity covaries with risk-taking personality in male Carpetan rock lizards (Iberolacerta cyreni). Ethology 122:355-363.
Molnár, O., Bajer, K., Szövényi, G., Török, J., Herczeg, G. (2016). Space use strategies and nuptial colorin European green lizards. Herpetologica 72:40-46.
Balázs, G., Lewarne, B., Herczeg, G. (2015). In situ underwater tagging of aquatic organisms: a test using the cave-dwelling olm, Proteus anguinus. Annales Zoologici Fennici 52:160-166.
Kopena, R., Herczeg, G., López, P., Martín, J. (2015). Escape strategy of Screiber’s green lizard (Lacerta schreiberi) is determined by the environment but not season or sex. Behaviour 152:1527-1542.
Garamszegi, L.Z., Zagalska-Neubauer, M. Canal, D., Markó, G., Szász, E., Zsebők, S., Szöllősi, E., Herczeg, G., Török, J. (2015). Malaria parasites, immune challenge, MHC variability, and predator avoidance in a passerine bird. Behavioral Ecology 26:1292-1302. (DOI:10.1093/beheco/arv077)
Bajer, K., Horváth, G., Molnár, O., Török, J., Garamszegi, L.Z., Herczeg, G. (2015). European green lizard (Lacerta viridis) personalities: linking behavioural types to ecologically relevant traits at different ontogenetic stages. Behavioural Processes 111:67-74. (DOI 10.1016/j.beproc.2014.11.020)
Garamszegi, L.Z., Markó, G., Szász, E., Zsebők, S., Azcárate, M., Herczeg, G., Török, J. (2015). Among-year variation in the repeatability, within- and between-individual, and phenotypic correlations of behaviors in a natural population. Behavioral Ecology and Sociobiology 69:2005–2017. (DOI 10.1007/s00265-015-2012-z)
Urszán, T.J., Garamszegi, L.Z., Nagy, G., Hettyey, A., Török, J., Herczeg, G. (2015). No personality without experience? A test on Rana dalmatina tadpoles. Ecology and Evolution 5:5847-5856. (DOI: 10.1002/ece3.1804) (open access)
Herczeg, G., Gonda, A., Balázs, G., Noreikiene, K., Merilä, J. (2015). Experimental evidence for sex-specific plasticity in adult brain. Frontiers in Zoology 12:38.
Urszán, T.J., Török, J., Hettyey, A., Garamszegi, L.Z., Herczeg, G. (2015). Behavioural consistency and life history of Rana dalmatina tadpoles. Oecologia 178:129-140. (DOI: 10.1007/s00442-014-3207-0)
Noreikiene, K., Herczeg, G., Gonda, A., Balázs, G., Husby, A., Merilä, J. (2015). Quantitative genetic analysis of brain size variation in sticklebacks: support for the mosaic model of evolution. Proceedings of the Royal Society B 282:20151008.
Saarikivi, J., Herczeg, G. (2014). Do hole-nesting passerine birds fare well at artificial suburban forest edges? Annales Zoologici Fennici 51:488-494
Laine, V.N., Herczeg, G., Shikano, T., Vilkki, J., Merilä, J. (2014). QTL analysis of behavior in Nine-spined sticklebacks (Pungitius pungitius). Behavior Genetics 44:77-88.
Garamszegi, L.Z., Mueller, J.C., Markó, G., Szász, E., Zsebők, S., Herczeg, G., Eens, M., Török, J. (2014). The relationship between DRD4 polymorphisms and phenotypic correlations of behaviors in the collared flycatcher. Ecology and Evolution 4:1466-1479. (open access)
Karhunen, M., Ovaskainen, O., Herczeg, G., Merilä, J. (2014). Bringing habitat information into statistical tests of local adaptation in quantitative traits: a case staudy of nine-spined sticklebacks. Evolution 68:559-568.
Herczeg, G., Välimäki, K., Gonda, A., Merilä, J. (2014). Evidence for sex-specific selection in brain: a case study of the nine-spined stickleback. Journal of Evolutionary Biology 27:1604-1612
Garamszegi, L.Z., Markó, G., Herczeg, G. (2013). A meta-analysis of correlated behaviors with implications for behavioral syndromes: relationships between particular behavioral traits. Behavioral Ecology 24:1068–1080.
Shikano, T., Laine, V.N., Herczeg, G., Vilkki, J., Merilä, J. (2013). Genetic architecture of parallel pelvic reduction in ninespine sticklebacks. Genes, Genomes, Genetics 3:1833-1842.
Gonda, A., Herczeg, G., Merilä, J. (2013). Evolutionary ecology of intraspecific brain size variation: a review. Ecology and Evolution 3:2751–2764.
Herczeg, G., Ab Ghani, N.I., Merilä, J. (2013). Evolution of stickleback feeding behaviour: genetics of population divergence at different ontogenetic stages. Journal of Evolutionary Biology 26:955-962.
Ab Ghani, N.I., Herczeg, G., Leinonen, T., Merilä, J. (2013). Evidence for genetic differentiation in timing of maturation among nine-spined stickleback populations. Journal of Evolutionary Biology 26:775-782.
Aikio, S., Herczeg, G., Kuparinen, A., Merilä, J. (2013). Optimal growth strategies under divergent predation pressure. Journal of Fish Biology 82:318–331.
Loehr, J., Herczeg, G., Leinonen, A., Gonda, A., Van Dongen, S., Merilä, J. (2013). Asymmetry in threespine stickleback lateral plates. Journal of Zoology 289:279-284.
Laine, V.N., Shikano, T., Herczeg, G., Vilkki, J., Merilä, J. (2013). Quantitative trait loci for growth and body size in the nine-spine stickleback Pungitus pungitius L. Molecular Ecology 22:5861-5876.
Bruneaux, M., Johnston, S.E., Herczeg, G., Merilä, J., Primmer, C.R., Vasemägi, A. (2013). Molecular evolutionary and population genomic analysis of the nine-spined stickleback using a modified RAD tag approach. Molecular Ecology 22:565-582.
Molnár, O., Bajer, K., Mészáros, B., Török, J., Herczeg, G. (2013). Negative correlation between nuptial throat colour and blood parasite load in male European green lizards supports the Hamilton–Zuk hypothesis. Naturwissenschaften 100:551-558.
Markó, G., Azcárate, M., Hegyi, G., Herczeg, G., Laczi, M., Nagy, G., Senar, J.C., Török, J., Garamszegi, L.Z. (2013). Behavioural responses to handling stress in the Great Tit: within-individual consistency and the effect of age, sex and body condition. Ornis Hungarica 21:12-25.
Ab Ghani, N.I., Herczeg, G., Merilä, J. (2012). Body size divergence in nine-spined sticklebacks: disentangling additive genetic and maternal effects. Biological Journal of the Linnean Society 107:521-528.
Saarinen, P.K., Pahlberg, J., Herczeg, G., Viljanen, M., Karjalainen, M.E., Shikano, T., Merilä, J., Donner, K. (2012). Spectral tuning by selective chromophore uptake in rods and cones of eight populations of nine-spined stickleback (Pungitius pungitius). Journal of Experimental Biology 215:2760-2773.
Välimäki, K., Herczeg, G., Merilä, J. (2012). Morphological antipredation defences in the nine-spined stickeleback: consititutive, induced or both? Biological Journal of the Linnean Society 107:854-866.
Molnár, O., Bajer, K., Török, J., Herczeg, G. (2012). Individual quality and nuptial throat colour in male European green lizards Journal of Zoology 287:233-239.(DOI:10.1111/j.1469-7998.2012.00916.x)
Välimäki, K., Herczeg, G. (2012). Ontogenetic and evolutionary effects of predation and competition on nine-spined stickleback (Pungitius pungitius) body size. Journal of Animal Ecology 81: 859-867.
Laine, V., Primmer, C., Herczeg, G., Merilä, J., Shikano, T. (2012). Isolation and characterization of 13 new nine-spined stickleback, Pungitius pungitius, microsatellites located nearby candidate genes for behavioural variation. Annales Zoologici Fennici 49:123-128.
Herczeg, G., Garamszegi, L.Z. (2012). Individual deviation from behavioural correlations: a simple approach to study the evolution of behavioural syndromes. Behavioural Ecology and Sociobiology 66:161-169.
Garamszegi, L.Z., Herczeg, G. (2012). Behavioural syndromes, syndrome deviation and the within- and between-individual components of phenotypic correlations: when reality does not meet statistics. Behavioural Ecology and Sociobiology 66:1651-1658.
Gonda, A., Välimäki, K., Herczeg, G., Merilä, J. (2012). Brain development and predation: plastic responses depend on evolutionary history. Biology Letters 8:249-252.
Trokovic, N., Herczeg, G., Ab Ghani, N.I., Shikano, T., Merilä, J. (2012). High levels of fluctuating asymmetry in isolated stickleback populations. BMC Evolutionary Biology 12:115.
Herczeg, G., Gonda, A., Kuparinen, A., Merilä, J. (2012). Contrasting growth strategies of pond versus marine populations of nine-spined stickleback (Pungitius pungitius): a combined effect of predation and competition? Evolutionary Ecology 26:109-122.
Garamszegi, L.Z., Markó, G., Herczeg, G. (2012). A meta-analysis of correlated behaviours with implications for behavioural syndromes: mean effect size, publication bias, phylogenetic effects and the role of mediator variables. Evolutionary Ecology 26:1213-1235.
Leinonen, T., McCairns, R.J.S., Herczeg, G., Merilä, J. (2012). Multiple evolutionary pathways to decreased lateral plate coverage in freshwater threespine sticklebacks. Evolution 66:3866-3875.
Laine, V.N., Herczeg, G., Shikano, T., Primmer, C.R. (2012). Heterozygosity-behaviour correlations in nine-spined stickleback (Pungitius pungitius) populations: contrasting effects at random and functional loci. Molecular Ecology 21:4872-4884.
Hjernquist, M.B., Söderman, F., Jönsson, K.I., Herczeg, G., Laurila, A., Merilä, J. (2012) Seasonality determines patterns of growth and age structure over a geographic gradient in an ectothermic vertebrate. Oecologia 170:641-649.
Bajer, K., Molnár, O., Török, J., Herczeg, G. (2012). Temperature, but not available energy, affects the expression of a sexually selected ultraviolet (UV) colour trait in male European green lizards, Lacerta viridis. PLoS ONE 7(3): e34359. (DOI:10.1371/journal.pone.0034359)
Loehr, J., Leinonen, T., Herczeg, G., O’Hara, R.B., Merilä, J. (2012). Heritability of asymmetry and lateral plate number int he threespine stickleback. PLoS ONE 7:e39843.
Bajer, K., Molnár, O., Török, J., Herczeg, G. (2011). Ultraviolet nuptial colour determines fight success in male European green lizards. Biology Letters 7:866-868.
Gonda, A., Herczeg, G., Merilä, J. (2011). Population variation in brain size of nine-spined sticklebacks (Pungitius pungitius) – local adaptation or environmentally induced variation? BMC Evolutionary Biology 11:75.
Shikano, T., Herczeg, G., Merilä, J. (2011). Molecular sexing and population genetic inference using a sex-linked microsatellite marker in the nine-spined stickleback (Pungitius pungitius). BMC Research Notes 4:119.
Leinonen, T., Herczeg, G., Cano J.M., Merilä, J. (2011). Predation-imposed selection on threespines stickleback (Gasterosteus aculeatus) morphology: a test of the refuge use hypothesis. Evolution 65: 2916-2926.
Trokovic, N., Gonda, A., Herczeg, G., Laurila, A., Merilä, J. (2011). Brain plasticity over the metamorphic boundary: carry-over effect of larval environment on froglet brain development. Journal of Evolutionary Biology 24:1380-1385.
Trokovic, N., Herczeg, G., McCairns, R.J.S., Ab Ghani, N.I., Merilä, J. (2011). Intraspecific divergence in the lateral line system in the nine-spined stickleback (Pungitius pungitius). Journal of Evolutionary Biology 24:1546-1558.
Herczeg, G., Välimäki, K. (2011). Intraspecific variation in behaviour: effects of evolutionary history, ontogenetic experience and sex. Journal of Evolutionary Biology 24:2434-2444.
Alho, J.S., Herczeg, G., Laugen, A.T., Räsänen, K., Laurila, A., Merilä, J. (2011). Allen’s rule revisited: quantitative genetics of extremity length in the common frog along a latitudinal gradient. Journal of Evolutionary Biology 24:59-70.
Kuparinen, A., Cano, J.M., Loehr, J., Herczeg, G., Gonda, A., Merilä, J. (2011). Fish age at maturation is influenced by temperature independently from growth. Oecologia 167:435-443.
Kopena, R., Martín, J., López, P., Herczeg, G. (2011). Vitamin E supplementation increases the attractiveness of males’ scent for female European green lizards. PLoS ONE 6(4):e19410.
Herczeg, G., Matsuba, C., Merilä, J. (2010). Sequence variation in the melanocortin-1 receptor gene (Mc1r) does not explain variation in the degree of melanism in a widespread amphibian. Annales Zoologici Fennici 47:37-45.
Waser, W., Sahoo, T.P., Herczeg, G., Merilä, J., Nikkinmaa, M. (2010). Physiological differentiation among nine-spined stickleback populations: effects of copper exposure. Aquatic Toxicology 98:188-195.
Bajer, K., Molnár, O.R., Török, J., Herczeg, G. (2010). Female European green lizards (Lacerta viridis) prefer males with high ultraviolet throat reflectance. Behavioural Ecology and Sociobiology 64:2007-2014. (DOI 10.1007/s00265-010-1012-2)
Herczeg, G., Turtiainen, M., Merilä, J. (2010). Morphological divergence of North-European nine-spined sticklebacks (Pungitius pungitius): signatures of parallel evolution. Biological Journal of the Linnean Society 101:403-416.
Alho, J.S., Herczeg, G., Söderman, F., Laurila, A., Jönsson K.I., Merilä, J. (2010). Increasing melanism along a latitudinal gradient: local adaptation, ontogenic or environmental plasticity? BMC Evolutionary Biology 10:317.
Herczeg, G., Gonda, A., Merilä, J. (2010). Rensch’s rule inverted – female-driven gigantism in nine-spined stickleback (Pungitius pungitius). Journal of Animal Ecology 79:581-588.
Gonda, A., Trokovic, N., Herczeg, G., Laurila, A., Merilä, J. (2010). Predation- and competition-mediated brain plasticity in Rana temporaria tadpoles. Journal of Evolutionary Biology 23:2300-2308.
Shikano, T., Shimada, Y., Herczeg, G., Merilä, J. (2010). History vs. habitat type: explaining the genetic structure of European nine-spined stickleback (Pungitius pungitius) populations. Molecular Ecology 19:1147-1161.
Bajer, K., Molnár, O., Hegyi, G., Herczeg, G., Laczi, M., Török, J. (2009). Zöld gyíkok színezete és morfológiája: jelzések és funkciók. Állattani Közlemények 94(2):167-176.
Herczeg, G., Gonda, A., Merilä, J. (2009). The social cost of shoaling covaries with predation risk in nine-spined stickleback (Pungitius pungitius) populations. Animal Behaviour 77:575-580.
Hettyey, A., Herczeg, G., Laurila, A., Crochet, P-A., Merilä, J. (2009). Body temperature, size, nuptial colouration and mating success in male Moor frogs (Rana arvalis). Amphibia-Reptilia 30:37-43.
Hettyey A, Herczeg G, Hoi H (2009). Testing the phenotype-linked fertility hypothesis in male Moor Frogs (Rana arvalis) exhibiting a conspicuous nuptial colouration. Amphibia-Reptilia 30:581-586.
Jönsson, K.I., Herczeg, G., O’Hara, R.B., Söderman, F., ter Shure, A.F.H., Larsson, P., Merilä, J. (2009). Sexual patterns of prebreeding energy reserves in the common frog Rana temporaria along a latitudinal gradient. Ecography 32:831-839.
Herczeg, G., Gonda, A., Merilä, J. (2009). Evolution of gigantism in nine-spined stickleback. Evolution 63:3190-3200.
Gonda, A., Herczeg, G., Merilä, J. (2009). Adaptive brain size divergence in nine-spined sticklebacks (Pungitius pungitius)? Journal of Evolutionary Biology 22:1721-1726.
Herczeg, G., Gonda, A., Merilä, J. (2009). Predation mediated population divergence in complex behaviour of nine-spined stickleback (Pungitius pungitius). Journal of Evolutionary Biology 22:544-552.
Gonda, A., Herczeg, G., Merilä, J. (2009). Habitat-dependent and -independent plastic responses to social environment in the nine-spined stickleback (Pungitius pungitius) brain. Proceedings of the Royal Society B 276:2085-2092.
Sos, T., Herczeg, G. (2009). Sexual size dimorphism in Eastern slow-worm (Anguis fragilis cholchica, Reptilia: Anguidae). Russian Journal of Herpetology 16:304-310.
Alho, J.S., Herczeg, G., Merilä, J. (2008). Female-biased sex ratios in subarctic common frogs. Journal of Zoology 275:57-63.
Herczeg, G., Herrero, A., Saarikivi, J., Gonda, A., Jäntti, M., Merilä, J. (2008). Experimental support for the cost-benefit model of lizard thermoregulation: the effects of predation risk and food supply. Oecologia 155:1-10.
Herczeg, G., Török, J., Korsós, Z. (2007). Size-dependent heating rates determine the spatial and temporal distribution of small-bodied lizards. Amphibia-Reptilia 28:347-356.
Herczeg, G., Kovács, T., Korsós, Z., Török, J. (2007). Microhabitat use, seasonal activity and diet of the snake-eyed skink (Ablepharus kitaibelii fitzingeri) with comparisons to the sympatric lacertids in Hungary. Biologia, Bratislava 62:482-487.
Herczeg, G., Saarikivi, J., Gonda, A., Perälä, J., Tuomola, A., Merilä, J. (2007). Suboptimal thermoregulation in male adders (Vipera berus) after hibernation imposed by spermiogenesis. Biological Journal of the Linnean Society 92:19-27.
Herczeg, G., Gonda, A., Perälä, J., Saarikivi, J., Tuomola, A., Merilä, J. (2007). Ontogenetic differences in the preferred body temperature of the European Adder Vipera berus (Serpentes: Viperidae). Herpetological Journal 17:58-61.
Ihász, N., Bajer, K., Kopena, R., Molnár, O., Herczeg, G., Török, J. (2006). Szemben a ragadozóval – a zöld gyík (Lacerta viridis) búvóhelyközpontú menekülési stratégiája. Állattani Közlemények 91:127-138.
Herczeg, G., Gonda, A., Saarikivi, J., Merilä, J. (2006). Experimental support for the cost-benefit model of lizard thermoregulation. Behavioral Ecology and Sociobiology 60:405-414.
Herczeg, G., Szabó, K., Korsós, Z. (2005). Asymmetry and population characteristics in dice snakes (Natrix tessellata): an interpopulation comparison. Amphibia-Reptilia 26:422-426.
Toth, T., Kovács, T., Herczeg, G. (2005). Morphological anomalies in a Finnish Subarctic population of the adder (Vipera berus berus). Litteratura Serpentinum 25:206-215.
Hettyey, A., Laurila, A., Herczeg, G., Jönsson, K.I., Kovács, T., Merilä, J. (2005). Does testis size declines towards the subarctic? – A case study on the common frog, Rana temporaria. Naturwissenschaften 92:188-192.
Harmos, K., Herczeg, G. (2004). A pannongyík elterjedése és természetvédelmi helyzete a Központi-Cserhátban és környékén. Folia Historico-naturalia Musei Matraensis 27:349-357.
Herczeg, G., Kovács, T., Tóth, T., Török, J., Korsós, Z., Merilä, J. (2004). Tail loss and thermoregulation in the common lizard Zootoca vivipara. Naturwissenschaften 91:485-488.
Herczeg, G., Tóth, T., Kovács, T., Korsós, Z., Török, J. (2004). Distribution of Ablepharus kitaibelii fitzingeri Mertens, 1952 (Squamata: Scincidae) in Hungary. Russian Journal of Herpetology 11:99-105.
Herczeg, G., Korsós, Z. (2003). Az interspecifikus kompetíció hatása a pannongyíkra (Ablepharus kitaibelii fitzingeri) egy antropogén hatásoknak kitett élőhelyen. Állattani Közlemények 88:73-84.
Herczeg, G., Kovács, T., Hettyey, A., Merilä, J. (2003). To thermoconform or thermoregulate? An assessment of thermoregulation opportunities for the lizard Zootoca vivipara in the subarctic. Polar Biology 26:486-490.
Herczeg, G., Krecsák, Sz.L., Marsi, Z. (2002). Új bizonyító adat a haragos sikló előfordulásáról Budapest belterületén a Sas-hegyről. Folia Historico-naturalia Musei Matraensis 26:341-344.

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