Journal of Ornithology

, Volume 157, Issue 1, pp 303–310 | Cite as

Maternal influence on eggshell maculation: implications for cryptic camouflaged eggs

  • Camille DuvalEmail author
  • Phillip Cassey
  • P. George Lovell
  • Ivan Mikšík
  • S. James Reynolds
  • Karen A. Spencer
Original Article


Egg camouflage may explain the adaptive significance of avian eggshell pigmentation in ground-nesting species. Eggshell maculation (spots) is predominantly due to protoporphyrin, but both biliverdin (antioxidant) and protoporphyrin (pro-oxidant) may be present in spotted eggshells. Because of their role in oxidative stress, the deposition of eggshell pigments might be condition-dependent. However, because of the fitness benefits of eggshell coloration, cryptic eggshell appearance should be strongly conserved in ground-nesting species regardless of female condition and eggshell pigment concentrations. We investigated whether Japanese quails (Coturnix coturnix japonica) maintained eggshell maculation under food restriction. We quantified eggshell maculation (i.e., percentage of spot coverage) using digital photography, and both protoporphyrin and biliverdin concentrations of eggs laid by females either on a food-restricted or an ad libitum diet. Females on a high quality diet, which are known to decrease the deposition of eggshell protoporphyrin, decreased eggshell maculation compared with food-restricted females that maintained it. For the first time, we propose an experimental study which suggests that eggshell maculation depends on female body condition and that manipulating eggshell maculation may be the strategy used by females to potentially optimize egg camouflage.


Body condition Camouflage Coturnix coturnix japonica Eggshell maculation Protoporphyrin 


Maternaler Einfluss auf die Befleckung der Eierschale: Folgen für kryptische getarnte Eier

Die Tarnung von Eiern könnte die adaptive Bedeutung der Pigmentierung der Eierschale bodenbrütender Vögel erklären. Die Befleckung der Eierschale ist hauptsächlich auf Protoporphyrin zurückzuführen, aber sowohl Biliverdin (ein Antioxidant) als auch Protoporphyrin (ein Prooxidant) können in gefleckten Eierschalen enthalten sein. Aufgrund der Rolle von Eierschalenpigmenten bei oxidativem Stress könnte ihre Einlagerung in die Eierschale konditionsabhängig sein. Aufgrund der Fitnessvorteile einer Färbung der Eierschale sollte ein kryptisches Aussehen der Eier bei Bodenbrütern jedoch hochkonserviert sein, unabhängig von der Kondition der Weibchen und der Konzentration der Eierschalenpigmente. Wir haben untersucht, ob Japanwachteln (Coturnix coturnix japonica) die Befleckung ihrer Eierschale bei Futterknappheit beibehielten. Wir haben die Eierschalenbefleckung (d.h. den Anteil der Abdeckung mit Flecken) mit Hilfe digitaler Fotografie quantifiziert sowie die Protoporphyrin- und Biliverdin-Konzentrationen der Eier von Weibchen ermittelt, die entweder eingeschränkt oder ad libitum Futter erhielten. Weibchen mit hochwertiger Kost, die bekannterweise die Einlagerung von Protoporphyrin in die Eierschale reduzieren, reduzierten die Befleckung der Eierschale im Vergleich zu Weibchen mit eingeschränkter Kost, welche sie beibehielten. Zum ersten Mal liefern wir eine experimentelle Studie ab, die darauf hindeutet, dass die Befleckung der Eierschale von der Kondition des Weibchens abhängt und dass Weibchen die Befleckung der Eierschale beeinflussen könnten, als Strategie, um die Tarnung der Eier potenziell zu optimieren.



We thank Carole Chestnut and Malcolm McColl from Cochno Farm (Glasgow) for their help with animal husbandry. Funding was provided by the Human Frontier Science Program (RGY0069 to PC), a Birmingham University Teaching Assistantship (to CD) and a Biotechnology and Biological Sciences Research Council (BBSRC) David Phillips Fellowship (BBE024459, to KAS). PC is an Australian Research Council (ARC) Future Fellow (FT0991420).

Compliance with ethical standards

All of the procedures were agreed by the local ethics committee at the University of Glasgow and the experiment was conducted under the Animals (Scientific Procedures) Act 1986 (under PIL 30/8939 held by CD and PPL 60/4068 held by KAS).

Conflict of interest

The authors declare that they have no conflicts of interest.

Supplementary material

10336_2015_1278_MOESM1_ESM.xls (38 kb)
Supplementary material 1 (XLS 38 kb)


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Copyright information

© Dt. Ornithologen-Gesellschaft e.V. 2015

Authors and Affiliations

  • Camille Duval
    • 1
    • 6
    Email author
  • Phillip Cassey
    • 2
  • P. George Lovell
    • 3
  • Ivan Mikšík
    • 4
  • S. James Reynolds
    • 5
  • Karen A. Spencer
    • 1
  1. 1.School of Psychology and NeuroscienceUniversity of St AndrewsSt AndrewsUK
  2. 2.School of Earth and Environmental SciencesUniversity of AdelaideAdelaideAustralia
  3. 3.Division of Psychology, Social and Health SciencesThe University of AbertayDundeeUK
  4. 4.Institute of PhysiologyAcademy of Sciences of the Czech RepublicPragueCzech Republic
  5. 5.Centre for Ornithology, School of Biosciences, College of Life and Environmental SciencesUniversity of BirminghamBirminghamUK
  6. 6.Tournon Sur RhôneFrance

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