Avian incubation is a critical part of reproduction and was extensively covered in the OUP book Avian Incubation published in 2002. This new book updates the older one by introducing new concepts that have arisen over the intervening years as well as updating key areas of research. The new book does not replace Avian Incubation but, rather, reinforces and supplements it. Therefore, this book covers four key areas: nest, eggs, incubation, and the study of avian reproduction. After an initial introduction about incubation research and a chapter reviewing the fossil record of avian reproduction chapters 3-6 focus on nest biology, including construction behaviour, functional properties, nest ecology, and the potential effects of climate change. Chapters 7-11 deal with egg biology with chapters addressing microbiology, allometry, energetics, and eggs as signals. Behaviour, energetics of incubation and the effects of incubation on post-hatching phenotype are reviewed in chapters 12-14. The study of avian reproduction is a new topic that was not covered in the previous book 13 years earlier. Chapters 15-17 deal with techniques for studying nests and the processes of incubation, the application of incubation science to conservation, and the contribution of citizen science to the study of reproduction of wild birds. The final chapter looks forward to new areas of study. Overall, this book collates and describes a range of recent developments in avian reproduction and complements the previous book in bringing the field right up to date.

Nests, Eggs, and Incubation: New ideas about avian reproduction

This chapter discusses several different aspects of the energy balance and physiological homeostasis of incubating birds, ranging from systematic, geographical and life history related variation in energy costs of incubation, to thermal considerations for birds on the nest, links between energy expenditure and fitness, and non-energetic costs of incubation. Our review of the literature shows that, across all bird species, this energy costs amounts to 3.4 times the basal metabolic rate (BMR). This is 15% lower than the cost of chick rearing (2.9 × BMR) for all birds, but is roughly equal to chick-rearing costs in those species in which only the female incubates. Energy costs are typically higher in challenging conditions, such as during incubation in harsh climates. This can impair fitness of parents and offspring, but little is understood about the physiological basis for such costs. We highlight and discuss possible mechanisms by which increased energy expenditure in incubating birds might hamper adult survival and, independently, carry over to also affect nestling phenotype and performance. We end by drawing attention to situations where the primary currency for incubation is not energy-based, which we exemplify by a discussion on the water economy of incubation in desert birds.

The energetic costs of incubation

Interspecific brood parasitism represents a prime example of the coevolutionary arms race where each party has evolved strategies in response to the other. Here, we investigated whether common cuckoos (Cuculus canorus) actively select nests within a host population to match the egg appearance of a particular host clutch. To achieve this goal, we quantified the degree of egg matching using the avian vision modelling approach. Randomization tests revealed that cuckoo eggs in naturally parasitized nests showed lower chromatic contrast to host eggs than those assigned randomly to other nests with egg-laying date similar to naturally parasitized clutches. Moreover, egg matching in terms of chromaticity was better in naturally parasitized nests than it would be in the nests of the nearest active non-parasitized neighbour. However, there was no indication of matching in achromatic spectral characteristics whatsoever. Thus, our results clearly indicate that cuckoos select certain host nests to increase matching of their own eggs with host clutches, but only in chromatic characteristics. Our results suggest that the ability of cuckoos to actively choose host nests based on the eggshell appearance imposes a strong selection pressure on host egg recognition.

https://royalsocietypublishing.org/doi/pdf/10.1098/rspb.2013.2665

Brood parasites lay eggs matching the appearance of host clutches

In oviparous taxa such as birds, eggs can be considered ‘sealed capsules’ that contain all of the nutrients and energy used during embryonic development (Carey 1996) and which only require external heat and oxygen to allow for complete development to hatching. Nevertheless eggs have complex macroand micronutrient compositions that serve not only as nutrients (resources) but potentially as ‘signals’, from the mother, that affect embryonic and post-hatching development and which, in turn, potentially affect offspring and adult phenotype and fitness in myriad ways (Williams 2012). Since these egg components are all maternally derived, i.e. transferred from mother to egg during egg formation, they have been considered important pathways for maternal effects, including non-genetic contributions of the mother to her offspring, and have become a major focus of life-history studies (Bernardo 1996; Mousseau and Fox 1998a; Badyaev and Uller 2009). Although maternal effects were originally considered to generate troublesome ‘noise’ in effective breeding programmes, the last 15 years have seen a paradigm shift in viewing such effects as potentially important and flexible ‘tools’ for the mother to adjust her offspring’s phenotype to current environmental conditions, with potentially important consequences for evolution of phenotype. Thus, much of the work on this topic has been conducted at the interface between ecology, evolutionary biology, and physiology within the context of how mothers might ‘adaptively’ adjust the development and the phenotype of their offspring to match prevailing environmental conditions in order to increase fitness (Mousseau and Fox 1998a; Groothuis and Schwabl 2008; Love and Williams 2008a; Uller 2008). There have been many reviews of different maternally derived egg components, but these have generally considered variation in single components, or groups of related components (e.g. macronutrients, Hill 1995; hormones, Groothuis et al. 2005; antioxidants, Blount et al. 2000; antibodies, Grindstaff et al. 2003). Nevertheless these reviews have all highlighted the same unresolved, or poorly understood, issues: a) the extent to which, and mechanism(s) by which, females might control or regulate maternal transfer to eggs; b) the extent to which embryos are passive recipients of maternal transfer, or whether they modulate the effects of maternally derived egg components; c) the mechanism(s) by which maternal substances might affect offspring development and ultimately adult phenotype to mediate long-term fitness; but also, d) the relative lack of studies supporting long-term fitness consequences of maternally derived egg components (cf. short-term, single trait studies), especially in free-living birds. Here, we will focus on these same issues but rather than consider egg macronutrients (e.g. lipid, protein), hormones, antioxidants, or immunoglobulins separately, we attempt to take an integrated approach to consider the effects of overall egg quality on embryonic development and the post-hatching phenotype. This is based on the idea that if females use maternally derived egg components to fine-tune offspring phenotype to prevailing environmental conditions, increasing offspring fitness independently of egg size per se, then there should be some ‘optimum’ combination of macroand micronutrients with which females attempt to furnish their eggs (assuming that the different components do not act independently of each other, or affect similar ‘targets’ in the offspring). Any optimum

Egg quality, embryonic development, and post-hatching phenotype: an integrated perspective

Bright colouration in animals has long attracted the attention of physicists, chemists and biologists. As such, studies on the functions of colours are interdisciplinary, focusing on the mechanisms of colour production and maintenance, the physical and chemical properties of the colour-producing elements, and visual systems and behaviour of potential receivers. Blue colouration has received a large share of research attention and is fascinating for several reasons: blue has been attributed to a very broad range of functions, blue is achieved by a great variety of mechanisms (although their production and maintenance costs are currently unclear), and the blue part of the spectrum (450–490 nm) can be perceived by most taxa. This review explores the breadth of studies that propose a function for blue colouration. In so doing, it discusses the diversity of ways in which blue colours are produced both as pigments and structural colours, and that blue visual pigments are common across a broad range of taxa. This analysis of the current literature emphasizes the importance of multidisciplinary hypothesis testing when attempting to elucidate the function of colours, the need for manipulative over correlative evidence for the function of colours, and, as colour research becomes evermore interdisciplinary, the need for well-defined consistent terminology.

https://zslpublications.onlinelibrary.wiley.com/doi/pdf/10.1111/jzo.12001

On the perception, production and function of blue colouration in animals

We analysed the eggs of the European starling Sturnus vulgaris for variability of UV and blue-green colouration. We noted the calcium deposition in eggshells, egg measurements, light conditions in nests and starling eye properties. Egg colouration shows high within-clutch similarity. Intensity of the blue-green egg colour neither depends on Ca deposition in the eggshell nor correlates with egg measurements, clutch size and laying date. This suggests that the intensity of the blue-green colour of starling eggs does not reflect female condition, which is reported to affect egg and clutch parameters. Low light intensity in cavities enables only mesopic vision, which is characterised by degraded visual acuity and colour sense, as well as Purkinje shift. As a consequence, the probability of precise assessment of the differences in eggs colour by starling males, crucial for the signalling hypothesis (Moreno & Osorno 2003), is not high. On the other hand, irrespective of its intensity, blue-green is the last col...

Is UV and blue-green egg colouration a signal in cavity-nesting birds?

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Is UV and blue-green egg colouration a signal in cavity-nesting birds?