The shape of birds’ eggs
Does selection during the incubation period influence egg shape?
Tim R. Birkhead
Department of Animal and Plant Sciences, University of Sheffield, U.K.
The evolution of egg shape in birds: selection during the incubation period. Birkhead, T.R., Thompson. J.E., Biggins, J.D. & Montgomerie, R. 2018. IBIS. DOI: 10.1111/ibi.12658. VIEW
When Thomas Wentworth Higginson’s in 1862 was asked to name the ‘most perfect thing in the universe’ he said: “a birds’ egg”. He must, in part at least, have been referring to the shape of birds’ eggs: smooth, rounded and much more varied than the expression ‘egg-shaped’ ever implies.
In cross section birds’ eggs are perfectly circular, but lengthwise they vary from near-spherical to elongate and often pointed at one or both ends.
Figure 1 Different avian egg shapes (not to scale) of different species © and Courtesy of Trustees of the Natural History Museum, Tring
The natural variation in egg shape across the 10,000 different bird species has triggered two questions among biologists:
- How does a female bird mould the shape of an egg?
- What is the adaptive significance of variation in egg shape?
Our new paper addresses the second question.
A major impediment to answering this question has been the difficulty of quantifying egg shape. The categories such as oval, elliptical and pyriform used extensively in the 19th-century actually grade into each other and are useful only for general description. More pragmatically, for scientific work, biologists, such as D’Arcy Wentworth Thompson in the early 1900s, began to devise mathematical equations that quantified some (but not all) egg shapes. The pyriform egg of the Common Guillemot Uria aalge egg was considered particularly problematical. Subsequently biologists such as F. W. Preston in the 1950s and 1960s, and D. Baker in 2002, built on Thompson’s foundation to create a range of egg shape indices, although from a practical point of view these were not particularly accessible because of the complexity of the maths.
Figure 2 A Common Guillemot egg © T.R. Birkhead
In June 2017, M. C. Stoddard and colleagues published a study in Science on the topic of the evolution egg shape in birds based on an analysis of a staggering 49,000 eggs from 1400 bird species. They concluded that ‘adaptations for flight may have been critical drivers for egg shape variation in birds’. The media was ecstatic that a long-standing puzzle had finally been solved. National Geographic for example, reported that this analysis showed that ‘birds with the higher hand-wing index — the most efficient and thus best fliers — were the ones with the most asymmetric or elliptical eggs’ and that, ‘as a bird’s body evolved for better flight, the shape of its egg might have adapted with it’. The headline in New Scientist claimed: ‘Bird eggs shaped by the way their mother flies’. The implication — certainly for the media, the public and most scientists — was that factors during egg formation drove the evolution of different egg shapes.
This seemed like a major discovery and we were very excited by the paper as we had been studying variation in the shapes and colours of eggs for some time. A closer look at the Stoddard et al. (2017) paper gave us three causes to question the generality of their conclusions. First, when we teased apart the complexities of their statistical analyses we discovered that ‘flight efficiency’ accounted for just 4% of the total interspecific variation in egg shape and that a lot of interspecific variation in egg shape remained to be accounted for.
Second, the huge sample size was compiled using published (online) photographs of eggs in the Museum of Vertebrate Zoology at Berkeley. Those are useful photographs documenting the collection, but they were not taken for the purpose of measuring egg shape accurately. To do that, one needs to ensure that the eggs are perfectly horizontal, otherwise their shape is often distorted in the analysed image, as you can see in the images below.
Figure 3 Photos of Spotted Sandpiper, Actitis macularius, eggs from Berkeley used by Stoddard et al. (2017)
Figure 4 Different shaped eggs (empty egg shells) resting on a horizontal surface, showing how some of them will be foreshortened when photographed from above © T.R. Birkhead
Third, Stoddard et al. used Baker’s (2002) method of quantifying egg shape. But, as the authors themselves noted, this method cannot deal adequately with pyriform egg shapes, many of which had to be excluded from their analyses.
Our own studies of egg shape started with finding a practical method — using Preston’s indices (see Biggins et al. 2018) — that accurately quantifies the shape of all birds’ eggs.
We also asked whether factors after egg formation might drive egg shape evolution. Our studies of guillemot eggs (Birkhead et al. 2018) strongly suggested that events associated with incubation are important and other authors have also suggested that selection during the incubation period influences egg shape, as in the case of wader eggs (e.g. Andersson 1978).
Video The evolution of egg shape in birds
The present study, comparing egg shape across the auks (Alcidae) and penguins (Spheniscidae) —two distantly related families with similar ecologies and life histories — confirmed that the type of site in which the egg is incubated has a major effect, with incubation site explaining more than 65% of the variation in shape.
Andersson, M. 1978. Optimal egg shape in waders. Ornis Fennica 55: 105-9. VIEW
Baker, D.E. 2002. A geometric method for determining the shape of bird eggs. Auk 119: 1179-1186. VIEW
Biggins, J.D., Thompson, J.E. & Birkhead, T.R. 2018. Accurately quantifying the shape of birds’ eggs. Ecology and Evolution, DOI: 10.1002/ece3.4412. VIEW
Birkhead, T.R., Thompson, J.E. & Montgomerie, R. 2018. The pyriform egg of the common murre Uria aalge is more stable on sloping surfaces. Auk Ornithological Advances 135: 1019-31. VIEW
Stoddard, M.C., Young, E.H., Akkaynak, D., Sheard, C., Tobias, J.A. & Mahadevan, L. 2017. Avian egg shape: form, function and evolution. Science 356: 1249-1254. VIEW
About the IBIS paper’s authors
Tim Birkhead‘s main research has been in avian mating system and sperm competition. He has also maintained a long-term study of guillemots on Skomer Island, Wales since 1972,that since the funding was cut by the Welsh Government has relied on public donations (see here). He has published a number of academic and popular science books including The Most Perfect Thing: the Inside (and Outside) of a Bird’s Egg (Bloomsbury),that won the Zoological Society of London’s Silver Medal (2016) for communicating zoology to a general audience. He has also won awards for his undergraduate teaching.
View Tim’s full profile
Jamie Thompson is a research technician working at The University of Sheffield on Tim’s Leverhulme Trust funded research project on avian egg shape. Whilst much of the previous several years has involved him enthusiastically studying guillemots on Skomer Island ,Wales, he has also undertaken research on avian sperm competition, problem solving in zebra finches and the reproductive biology of Galliformes.
John Biggins was until the end of last year the Head of the Department of Mathematics and Statistics. His main research interests are in applied probability with his main work being on the theory of branching processes, and the branching random walk in particular. In a separate research strand, he and Tim Birkhead have collaborated on a number of ornithological projects since the 1970s.
View John’s full profile
Bob Montgomerie is an evolutionary biologist who studies sexual selection and the evolution of gamete structure (sperm and eggs). He first encountered guillemots in 1973 while studying seabirds for the Canadian Wildlife Service in both Newfoundland and the high arctic of Nunavut, Canada. Since then his research has taken him around the world to study more than 50 bird species, largely focusing on reproduction (courtship, ornaments, parental care). With Tim Birkhead and Jo Wimpenny he coathoured Ten Thousand Birds (Princeton) chronicling the history of ornithology since Darwin. To date he has published more than 200 scientific papers and book chapters, and currently writes a blog on the history of ornithology for the American Ornithological Society.
View Bob’s full profile
Follow Bob on Twitter @BobMontgomerie
Top right: A Common Guillemot, Uria aalge, incubating © T.R. Birkhead
Blog posts express the views of the individual author(s) and not those of the BOU.
If you want to write about your research in #theBOUblog, then please see here.