In review / under revision:

  • Ramesh, A,Gismann, J. , Groothuis, A.G.G., Weissing, F.J. & Nicolaus, M. Mesocosm experiments reveal the loss of migratory tendencies in a recently isolated population of three-spined sticklebacks Biology Letters
  • Lamers K.; Nicolaus M.; Burgess M.; Pearce-Higgins J.; Nilsson J.-Å. & Both C. Northward dispersal in early springs: the effects of spring phenology on long-distance dispersal in pied flycatchers.  Journal of Animal Ecology 


  • Nicolaus, M., Ubels R. & Both C. Eco-evolutionary consequences of dispersal syndromes during colonization in a passerine bird. The American Naturalist. In press
  • Nicolaus, M., Wang, X., Lamers K.P., Ubels R. & Both C. (2022) Unravelling the causes and consequences of dispersal syndromes in a wild passerine. Proceedings of the Royal Society of London B. 289 (1974): 20220068
  • Verkuil Y., Nicolaus M., Ubels R., Dietz M., Samplonious J., Galema A., Kiekebos K., Knijff Pd & Both C. (2022) DNA metabarcoding quantifies relative abundances of arthropod taxa in songbird diets: validation with camera-recorded diets. Ecology & Evolution. In press
  • Ramesh A., Groothuis A.G.G, Weissing F.J. & Nicolaus M. (2022) Habitat fragmentation induces rapid phenotypic divergence of migratory and isolated sticklebacks. Behavioral Ecology. 33(1): 167-177.


  • Ramesh A., Matos Domingues M., Stamphuis E, Groothuis A.G.G, Weissing F.J. & Nicolaus M. (2021) Does genetic differentiation underlie behavioral divergence in response to migration barriers in sticklebacks? A common garden experiment Behavioral Ecology and Sociobiology. 75(12): 161-172.
  • Mouchet A., Cole E.F., Matthysen E., Nicolaus M., et al. (2021) Heterogeneous selection on exploration behavior within and among West European populations of a passerine bird. Proceedings of the National Academy of Sciences of the United states of America. 118: e2024994118
  • De Jong M.E., Nicolaus M., Fokkema R.W. & Loonen M.J.J.E (2021) State-dependence explains individual variation in nest defence behaviour in a long-lived bird. Journal of Animal Ecology 90: 809-819


  • Lamers P., Nicolaus M., Nilsson J-Å & Both C. (2020) Descriptive and experimental evidence for timing-mediated polygyny risk in a pied flycatcher (Ficedula hypoleuca) population. Journal of Avian Biology 51(2): e02190


  • Mutzel  A., Olsen A-L., Mathot K.J., Araya-Ajoy Y.G., Nicolaus M., Wijmenga J., Wright J.; Kempenaers B. & Dingemanse N.J. (2019) Effects of manipulated levels of predation threat on parental provisioning and nestling begging. Behavioral Ecology, 30(4): 1123–1135
  • Nicolaus M., Barrault S. & Both C. (2019) Diet and provisioning rate differ predictably between dispersing and philopatric pied flycatchers. Behavioral Ecology, 30(1): 114-124


  • Nicolaus M. & Edelaar P. (2018) Comparing the consequences of natural selection, adaptive phenotypic plasticity, and matching habitat choice for phenotype-environment matching, population genetic structure, and reproductive isolation in meta-populations. Ecology and Evolution, 8(8): 3815-3827


  • Mathot K.J., Olsen A.-L., Mutzel A., Araya-Ajoy Y.G., Nicolaus M., Westneat D.F., Wright J., Kempenaers B. & Dingemanse N.J. (2017) Provisioning tactics of great tits (Parus major) in response to long-term brood size manipulations differ across years. Behavioral Ecology, 28 (6): 1402–1413


  • Araya-Ajoy Y.G., Kuhn S., Mathot K.J., Mouchet A., Mutzel A., Nicolaus M., Wijmenga J.J., Kempenaers B. & Dingemanse N.J. (2016). Sources of (co)variation in alternative siring routes available to male great tits (Parus major). Evolution 70(10): 2308-2321
  • Nicolaus M., Piault R.,  Ubels R., Tinbergen J.M. & Dingemanse N.J. (2016). The correlation between coloration and exploration behaviour varies across hierarchical levels in a wild passerine bird. Journal of Evolutionary Biology 29 (9): 1780-1792
  • Nicolaus M., Tinbergen J.M., Ubels R., Both C. & Dingemanse N.J. (2016). Density fluctuations represent a key process maintaining personality variation in a wild passerine birds. Ecology Letters, 19: 478-486 (for press article about it click here)


  • Nicolaus M., Mathot K.J., Araya-Ajoy Y.G., Mutzel A., Wijmenga J., Kempenaers B. & Dingemanse N.J. (2015). Does coping styles predict optimization? An experimental test in passerine bird. Proceedings of the Royal Society of London B 282: 20142405
  • Mathot K.J., Nicolaus M., Araya-Ajoy Y.G., Dingemanse N.J. & Kempenaers B. (2015). Does metabolic rate predict risk-taking behaviour? A field experiment in a wild passerine bird. Functional Ecology. 29 (2): 239-249


  • Nicolaus M., Brommer J.E., Ubels R., Tinbergen J.M. & Dingemanse N.J. (2013). Exploring patterns of variation in clutch size–density reaction norms in a wild passerine bird. Journal of Evolutionary Biology 26: 2031-2043.
  • Stuber E.F., Araya-Ajoy Y.G., Mathot K.J., Mutzel A., Nicolaus M., Wijmenga J.J., Mueller J.C. & Dingemanse N.J. (2013). Slow explorers take less risk: a problem of sampling bias in ecological studies. Behavioral Ecology 24: 1092-1098.
  • Michler S.P.M., Nicolaus M., van der Velde M., Radersma R., Ubels R., Both C., Komdeur J. & Tinbergen J.M. (2013). Local offspring density and sex ratio affect sex allocation in the great tit. Behavioral Ecology 24: 169-181.


  • Nicolaus M., Tinbergen J.M., Bouwman K.M., Michler S.P.M., Ubels R., Both C., Kempenaers B. & Dingemanse N.J. (2012). Experimental evidence for adaptive personalities in a wild passerine bird. Proceedings of the Royal Society of London B 279: 4885-4892.
  • Nicolaus M., Michler S.P.M., Ubels R., van der Velde M., Bouwman K.M., Both C. & Tinbergen J.M. (2012). Local sex ratio affects the cost of reproduction. Journal of Animal Ecology 81: 564-572.
  • Nicolaus M., Michler S.P.M., Jalvingh K.M., Ubels R., van der Velde M., Komdeur J., Both C. & Tinbergen J.M. (2012). Social environment affects juvenile dispersal in great tits (Parus major). Journal of Animal Ecology 81: 827-837.


  • Michler S.P.M., Nicolaus M., Ubels R., van der Velde M., Both C., Tinbergen J.M. & Komdeur J. (2011). Do sex-specific densities affect local survival of free ranging great tits? Behavioral Ecology 22: 869-879.
  • Michler S.P.M., Nicolaus M., Ubels R., van der Velde M., Komdeur J., Both C. & Tinbergen J.M. (2011). Sex-specific effects of the local social environment on juvenile post-fledging dispersal in great tits. Behavioral Ecology and Sociobiology 65: 1975-1986.
  • Nicolaus, M. (2011) Review of: The maintenance of variation in avian personality: the role of the social environment par Eva Fucikova, Ardea 99: 243-244.


  • Nolan P.M., Dobson F.S., Nicolaus M., Karels T.J., Mcgraw K.J. & Jouventin P. (2010). Mutual mate choice for colorful traits in King Penguins. Ethology 116: 635-644.


  • Nicolaus M., Both C., Ubels R., Edelaar P. & Tinbergen J.M. (2009). No experimental evidence for local competition in the nestling phase as a driving force for density-dependent avian clutch size. Journal of Animal Ecology 78: 828-838.
  • Nicolaus M., Michler S.P.M., Ubels R., van der Velde M., Komdeur J., Both C. & Tinbergen J.M. (2009). Sex-specific effects of altered competition on nestling growth and survival: an experimental manipulation of brood size and sex ratio. Journal of Animal Ecology 78: 414-426.
  • Nicolaus, M. (2009) Reproductive rates under competition. PhD Thesis, University of Groningen, (reviewed by M. van de Pol (2010) Ardea 98: 120-121).


  • Nicolaus M., Bouwman K.M. & Dingemanse N.J. (2008). Effect of PIT tags on the survival and recruitment of Great Tits Parus major. Ardea 96: 286-292.
  • Dobson F.S., Nolan P.M., Nicolaus M., Bajzak C., Coquel A. & Jouventin P. (2008). Comparison of color and body condition between early and late breeding King Penguins. Ethology 114: 925-933.
  • Jouventin P., Nolan P.M., Dobson F.S. & Nicolaus M. (2008). Coloured patches influence pairing rate in King Penguins. Ibis 150: 193-196.


  • Bried J., Nicolaus M., Jarne P., Dubois M.P. & Jouventin P. (2007). Population biology of the wandering albatross (Diomedea exulans) in the Crozet and Kerguelen archipelagos, southern Indian Ocean, approached through genetic and demographic methods. Journal of Zoology 272: 20-29.
  • Nicolaus M., Le Bohec C., Nolan P.M., Gauthier-Clerc M., Le Maho Y., Komdeur J. & Jouventin P. (2007). Ornamental colors reveal age in the king penguin. Polar Biology 31: 53-61.