My PhD viva experience

As I mentioned in my last post I successfully defended my PhD viva (thesis defence) just over three weeks ago. As I was preparing to defend my thesis I found it helpful and often reassuring to read the experiences of other PhD students to get some idea of what I could expect. Of course, every viva will be different but below are some of my thoughts post-viva which I hope my be useful to others.


In the weeks leading up to my viva I tried to prepare as best I could by re-reading my thesis and refreshing my memory of the major papers which I had cited often or based my own work upon. Apart from this I didn’t feel like I could really do much to prepare as I had no idea what questions I would be asked. At the time I found this quite stressful because I felt like I should have been working harder and doing more to prepare. Now with the benefit of hindsight I can see that I did the right things.

What my reviewers most wanted to know was that I understood and could explain my own work. Since I had spent the last 3+ years working on it this was not a problem! What I now know is that the preparation for my viva began on the very first day of my PhD. While it was helpful (and definitely recommended) to refresh my memory of the key points of my work in the weeks leading up to my viva,  the real hard work had already been done over the past years.

The warm-up

On the day of my viva I got up early and read a few notes one last time over breakfast. I then headed into my university for a pre-viva chat and coffee with my supervisor. This really helped calm any nerves I had as he seemed quite confident that I would have no problems.

In the hour before my viva began I had to go and register with the secretary before being shown to the room where I would meet my reviewers. Here I had to wait outside for quite a few minutes and it was at this point that I first started to get really nervous. I think because the time waiting gave me the time to think of all the possible worst case scenarios!

Eventually, I was shown into the room where my reviewers introduced themselves and explained what the viva was and what would happen over the next few hours. I was offered drinks which had been prepared for us (tea/coffee/water) before the viva really began. These first few minutes felt like a warm-up before the real questioning began.


The first thing I was asked to do was to explain in broad terms what my thesis was about, why I did it and what I had discovered. This was possibly the easiest question of the whole day as I had spent so long working on my PhD that I could explain it in my sleep. The only difficulty I had was condensing more than three years of results into fifteen minutes of explanation. This meant leaving out everything but the biggest and most important elements of my work.

On a couple of occasions I found myself spending too much time explaining minor details and had to force myself to move on to more important aspects of my work. Nevertheless, this question really helped me to get into the flow of explaining my research and after a few minutes I really felt like I was firing on all cylinders.

After this warm-up question things got a lot more detailed. My reviewers went through each chapter one by one and stopped wherever they had a question about something I had written or wanted some more detail about a particular point I had raised. In almost all cases these questions were not too difficult, and in most cases they were relatively easy to answer.

However, I don’t want to suggest that there were no difficult questions. On a couple of occasions a question was put to me that I couldn’t answer very well or a criticism of my work was made which I could only agree with. During the viva this made me quite uncomfortable, but now, with the benefit of hindsight, I can see that my reviewers pushed me to look at my work from a different perspective and consider new possibilities. This is probably one of the best things to come out of my viva and will really help me as I write-up some of my chapters for publication.

Towards the end

My viva was just under three hours long. It sounds like a really long time but it really didn’t feel like it. After the first few questions it felt more like a friendly discussion that an exam and I relaxed a little bit and started to feel slightly more confident. This, along with the stress and adrenaline really helped the time to fly by.

Once both my reviewers were satisfied that they had no more questions to ask me I was asked to leave the room for a few minutes while they discussed how I did and what the outcome should be. This was probably the most nerve-wracking moment for me as I really had no idea how well I’d done or what outcome to expect. After maybe five minutes I was asked back into the room where my reviewers both seemed happy and congratulated me on passing with minor corrections. This means I  have up to six months to make a few changes to my thesis which in reality should take no more than a week or two.

And relax!

The feeling of relief on being given the news that I had passed was overwhelming and I didn’t really know how to react. The corrections I need to make are only very small and my reviewers really seemed to like my work.

Once the viva was over I had a celebratory pint in the local pub with my supervisor, a reviewer and some friends. At this point I was still dazed, relieved, tired and hadn’t really full absorbed had what just happened. I think it took a full 24 hours before it really sank  in that all of my work over the past years had been worth it. Often it was fun, at times it was hard, stressful and I felt like I couldn’t do it. But now it has paid off and I am happy!

If you are a PhD student about to defend your thesis and have any questions about the process please feel free to email me or leave a comment. And most of all, good luck!


The end of the beginning

The very first post on this blog was written while I was still an undergraduate at the University of Derby. Today I am happy to say that I have just successfully defended my PhD thesis on the effects of urbanisation on the behaviour of song birds. This means not only can I order a new bank card with “Dr.” on it, but also that my time as a student has finally come to an end.

The first draft of my PhD thesis just before I submitted it.

The question I have to ask myself now is what next?

Right now I am working at the Max Planck Institute for Ornithology in Germany and am loving it. I work with a really fantastic research group and I hope I will be able to stay here a while yet to pursue some of my research ideas. After that, I have no idea! I am considering several possibilities including continuing in academia long-term, working for conservation/environmental charities or applying for jobs in government environmental agencies. If you have completed a PhD I would love to hear in the comments what you did next.

As for this blog, I hope to revive it after its long and unplanned hiatus. As all PhD students know, the final year is the most intense and personally challenging. In the last year I have moved to Germany, written my thesis and published a paper. amongst all this I didn’t find the time for my poor blog! Now, however, I have a little more time and I plan to use it to write a series of blog posts on my PhD experience. Both the good and the bad. If there is anything you would like to know about the PhD experience please let me know below!

I also plan to write about whatever I do next. Although it is still quite unclear at the moment I am optimistic for the future and looking forward to the next challenge. The end of being a student me is really just the beginning of the next stage in my life and I can’t wait!


Urbanisation is changing the way birds sing

ResearchBlogging.orgIn 1800 only 3% of the world’s population lived in urban areas, yet as the industrial revolution picked up pace in the early 18th and 19th centuries the number of people moving from the countryside to work in the newly industrialised cities soared. By 1950 29% of the world’s population were living in cities and by 1985 this had grown to 42% while in 2025 it is estimated to that it will be over 60%1. That is just 10 years from now. In highly developed nations such as those in Western Europe and North America the 50% threshold has already been surpassed which means that if you live in a western nation and do not live in a city you are in a minority.

Across the globe cities are expanding.

It’s not just where people choose to live that is rapidly changing either, but how many of us there are. The human population is currently expanding more rapidly than ever before and has grown from just a few hundred million people less than a thousand years ago to over 7 billion today, and this number is still rising2. As the human population grows cities are expanding quickly to meet the need for additional housing while the surrounding countryside is farmed and developed ever more intensely to provide us with the food, water and other resources we need to live.

Human population growth from around 5 million people in 8000 BCE to over 7 billion today. (figure from Keinen and Clark 2012).
Human population growth from around 5 million people in 8000 BCE to over 7 billion today. (figure from Keinen and Clark 2012).

Naturally, many people are concerned about how we are going to continue to feed and clothe ourselves as populations continue to expand, but there is also a growing concern among many about how such huge numbers of people are affecting the environment and the animals that inhabit it.

For animals which depend on natural habitats such as forests, meadows, or wetlands to survive, the growth of urban areas often spells bad news as these habitats are removed and paved over to make way for new suburbs, factories and roads. In the UK numerous species have declined for just this reason. For example, the bittern, a close relative of herons, was once widespread in reedbeds and wetlands across the UK but is now confined to a tiny area of the south-east after its habitats were drained to make way for agriculture and urban developments.

While many species cannot survive in urbanised areas, others are able to tolerate moderate levels of urbanisation and may continue living within cities despite drastic changes to their habitats. Life in the city is not without its challenges however, even for the most adaptable and resilient of species. Cities typically contain different threats to rural areas such as an abundance of cats which are responsible for killing huge numbers of birds and small mammals4, and high levels of chemical5, light6 and noise pollution7 which all have negative impacts.

This all sounds bad, and it really is, but while many species suffer badly from the effects of increasing urbanisation and habitat loss, there are a few species that have been able to adjust remarkably well to life in urban areas. One group that has been particularly well-studied in this regard are the songbirds and over the past 15 years or so, biologists have discovered some fascinating behavioural adaptations which have allowed some species to become successful city dwellers.

Great tits are one species which are adapting to city life,
Great tits are one species which are adapting to city life.

One of the most notable features of cities across the world is that they are incredibly noisy places. With heavy traffic, building sites, aircraft flying overhead and all manner of other sounds and distractions it’s a miracle anyone can hear anything at all. For songbirds however, all this noise is more than just a distraction, it can seriously affect their chances of finding mates and successfully reproducing and for males it is likely to affect how well they can defend their territories against rivals.

While we may find bird song pleasant to listen to (or annoying depending on how early in the morning it is), for songbirds it has a serious purpose. Males sing during the breeding season to attract females8 and to signal to other males that their territory is occupied and should not be entered9.

City noise can overlap and interfere with these signals making communication among birds difficult and unreliable. The background noise of a city is typically continuous low rumble concentrated at around 2kHz in frequency. Unfortunately for many birds this overlaps neatly with the frequency of their songs and this can make it difficult for other birds to hear them as they do not stand out from the irrelevant background noise.

Clearly this is a problem for birds which rely on song to communicate, yet research has revealed that birds have ways of overcoming this problem and one of them is to increase the frequency at which they sing so that their songs literally rise above the background noise and can be clearly heard.

Great tits sing at a higher frequency in noisy cities than in quieter rural areas (from Mockford and Marshall, 2009).
Sonograms of rural (a) and urban (c) great tit song compared to the background noise of a city (b). You can clearly see that the urban song is a higher frequency than rural song and above the frequency of the city noise. (from Mockford and Marshall, 2009).

Evidence that birds sing at higher frequencies in noisy cities than they do in quieter rural sites has now been found in numerous species including great tits (Parus major)7, blackbirds (Turdus merula)10, European robins (Erithacus rubecula)11 and song sparrows (Melospiza melodia)12.

In great tits the difference in song frequency between urban and rural sites has been measured at 478Hz13 and tests have shown that this is enough to substantially improve the distance over which song can travel in urban environments before it degrades and becomes inaudible14.

Birds may also face challenging noisy conditions in natural environments too such as where running water or wind creates high levels of low-frequency noise and these naturally noisy sites have allowed scientists to confirm that it really is the noise in cities and not some other factor which is causing city birds to sing at high frequencies. Biologists Henrik Brumm and Hans Slabbekoorn recorded the songs of white-throated dippers (Cinclus cinclus) living around noisy fast flowing streams in Scotland and found that they call at frequencies well above that of the background noise and higher than usual for this species suggesting that dippers in this area have adapted their calls to suit their noisy habitat15.

Dippers close to noisy streams sing at a higher frequency than the background noise so their songs can be heard (from Brumm and Slabbekoorn, 2005).

The effect of natural background noise on song frequency has also been shown in African little greenbuls (Andropadus virens) which sing at a higher frequency in areas where the rainforest is merging with open grasslands (known as ecotone forests) than they do deep within the rainforest itself16. Analysis of these two habitats revealed that the background noise in the rainforest is largely concentrated at higher frequencies while in ecotone forests there is more low-frequency noise. By singing at a lower frequency little greenbuls within the rainforest can ensure that their song does not overlap with the higher frequency background noise found in rainforests, while by singing at a lower frequency little greenbuls in ecotone forests avoid the lower frequency background noise in their habitat.

The evidence that birds change the frequency of their songs as an adaptation to noisy conditions may seem quite conclusive but not everyone agrees. An alternative explanation for the observed frequency shifts is that higher frequency song is actually just an unavoidable and possibly unimportant side-effect of singing more loudly, and it is higher volume, not frequency, which allows birds in noisy environments to overcome the background noise17.

In support of this argument Erwin Nemeth and Henrik Brumm of the Max Planck Institute for Ornithology in Germany found that the typical increases in song frequency found in great tits and blackbirds may be too low to substantially improve signal transmission whereas small increases in song amplitude were found to increase the distance over which a bird’s song could be detected much more effectively18.

But why should song frequency increase when birds sing more loudly? Nemeth and Brumm suggest two possibilities. Firstly, the increase in frequency observed in songs in noisy environments could be a side-effect of what is known as the Lombard effect (named after the French scientist Étienne Lombard) in which animals unconsciously increase the volume and frequency of their calls when the level of the background noise rises.

Nightingales in Berlin sing loudly to ensure that they are heard.

The Lombard effect is known to occur in humans (this is why it might feel like you have to shout to be heard at loud parties) and has also been shown in both lab and field studies of songbirds. Lab experiments on elegant crested tinamous (Eudromia elegans)19, and budgerigars (Melopsittacus undulates)20, have shown that these species both sing more loudly and at a higher frequency when background noise increases and the same result has been shown in the field in a study of Nightingales (Luscinia megarhynchos) in Berlin21.

A second reason why song frequency may increase when birds sing more loudly is that both the volume and frequency of bird songs depend on the same song producing organ which could limit how well birds can independently control frequency and volume. In birds this organ is the syrinx which is the bird equivalent of the mammalian larynx or voice box and is located at the base of the windpipe connected to the lungs. Birds produce song by forcing air at high pressure from the lungs through the syrinx causing membranes to vibrate creating sound. This sound can then be modified using numerous tiny muscles which alter the shape and tension of the sound producing membranes.

However, past studies of the avian vocal system have shown that without these tiny muscles altering the structure of the sound, both the frequency and amplitude of bird song unavoidably increase together. In other words, when birds sing louder they cannot help but also sing at a higher frequency22.

Of course, this may be totally irrelevant if birds are able to use muscles to independently control the frequency and volume of their songs but there is evidence to suggest that the frequency and volume of bird songs really are closely intertwined. One of the clearest examples of this comes from a study by Nemeth and his colleagues at the Max Planck institute who recorded blackbirds singing in sound-proof chambers and showed that volume and frequency really were strongly correlated17. When blackbirds sing more loudly they also sing at a higher frequency and this may be totally involuntary. Similar results have been found in other species including zebra finches (Taeniopygia guttata)23 and song doves (Streptopelia risoria)24 suggesting that this pattern may be widespread in birds as a whole.

When blackbirds increase the volume of the song they also sing at a higher frequency. (image credit Juan Emilio).

It has become very clear over the past few years that urban noise is causing bird song to change however, opinion is still divided on whether it is the frequency or amplitude changes that are most important to improving song transmission in noisy environments. It is possible that both have important roles to play in helping birds to adapt to noisy urban areas and hopefully future research will provide an answer to this question.

The study of how urban noise affects bird song is a very active area of research and there are many unresolved questions which are likely to be answered in the next few years. Most importantly we need to find out what the long-term impacts of urban noise are on bird populations. Although many species of birds do seem to be able to adapt to noise we do not know how the dramatic changes we are causing to their environments will affect them in the long-term. Furthermore, many species are not able to adapt to urban areas for numerous possible reasons. They may not possess the behavioural flexibility to cope with new environments or not they might not be physiologically capable of changes their songs or behaviour. That is why studies those discussed here matter, we are changing the planet in ways which have never been seen before and we know that many species are suffering as a result. The first step to protecting animals from these changes is to understand how they are affected and that is just what these studies aim to do.


1. Kegel, B (2014).Tiere in der Stadt: Eine Naturgeschichte. Köln: DuMont Buchverlag. (In German).

2. Keinan, A., & Clark, A. (2012). Recent Explosive Human Population Growth Has Resulted in an Excess of Rare Genetic Variants. Science, 336 (6082), 740-743 DOI: 10.1126/science.1217283

3. Barnosky AD, Matzke N, Tomiya S, Wogan GO, Swartz B, Quental TB, Marshall C, McGuire JL, Lindsey EL, Maguire KC, Mersey B, & Ferrer EA (2011). Has the Earth’s sixth mass extinction already arrived?. Nature, 471 (7336), 51-7 PMID: 21368823

4. van Heezik, Y., Smyth, A., Adams, A., & Gordon, J. (2010). Do domestic cats impose an unsustainable harvest on urban bird populations?. Biological Conservation, 143 (1), 121-130 DOI: 10.1016/j.biocon.2009.09.013

5. Liker A, Papp Z, Bókony V, & Lendvai AZ (2008). Lean birds in the city: body size and condition of house sparrows along the urbanization gradient. The Journal of animal ecology, 77 (4), 789-95 PMID: 18479344

6. Miller, M. (2006). Apparent Effects of Light Pollution on Singing Behavior of American Robins. The Condor, 108 (1) DOI: 10.1650/0010-5422(2006)108[0130:AEOLPO]2.0.CO;2

7. Slabbekoorn, H., & Peet, M. (2003). Ecology: Birds sing at a higher pitch in urban noise. Nature, 424 (6946), 267-267 DOI: 10.1038/424267a

8. Baker, M., Bjerke, T., Lampe, H., & Espmark, Y. (1986). Sexual Response of Female Great Tits to Variation in Size of Males’ Song Repertoires. The American Naturalist, 128 (4) DOI: 10.1086/284582

9. Krebs, J., Ashcroft, R., & Webber, M. (1978). Song repertoires and territory defence in the great tit. Nature, 271 (5645), 539-542 DOI: 10.1038/271539a0

10. Nemeth, E., & Brumm, H. (2009). Blackbirds sing higher-pitched songs in cities: adaptation to habitat acoustics or side-effect of urbanization? Animal Behaviour, 78 (3), 637-641 DOI: 10.1016/j.anbehav.2009.06.016

11. McLaughlin, K., & Kunc, H. (2012). Experimentally increased noise levels change spatial and singing behaviour. Biology Letters DOI: 10.1098/rsbl.2012.0771

12. Wood, W., & Yezerinac, S. (2006). Song sparrow (Melospiza melodia) song varies with urban noise. The Auk, 123 (3) DOI: 10.1642/0004-8038(2006)123[650:SSMMSV]2.0.CO;2

13. Mockford, E., & Marshall, R. (2009). Effects of urban noise on song and response behaviour in great tits. Proceedings of the Royal Society B: Biological Sciences, 276 (1669), 2979-2985 DOI: 10.1098/rspb.2009.0586

14. Mockford, E., Marshall, R., & Dabelsteen, T. (2011). Degradation of Rural and Urban Great Tit Song: Testing Transmission Efficiency. PLoS ONE, 6 (12) DOI: 10.1371/journal.pone.0028242

15. Brumm, H., & Slabbekoorn, H. (2005). Acoustic communication in noise. Advances in the Study of Behavior, 35, 151-209 DOI: 10.1016/S0065-3454(05)35004-2

16. Slabbekoorn H, & Smith TB (2002). Habitat-dependent song divergence in the little greenbul: an analysis of environmental selection pressures on acoustic signals. Evolution; international journal of organic evolution, 56 (9), 1849-58 PMID: 12389730

17. Nemeth, E., Pieretti, N., Zollinger, S., Geberzahn, N., Partecke, J., Miranda, A., & Brumm, H. (2013). Bird song and anthropogenic noise: vocal constraints may explain why birds sing higher-frequency songs in cities. Proceedings of the Royal Society B: Biological Sciences, 280 (1754), 20122798-20122798 DOI: 10.1098/rspb.2012.2798

18. Nemeth, E., & Brumm, H. (2010). Birds and Anthropogenic Noise: Are Urban Songs Adaptive?. The American Naturalist, 176 (4), 465-475 DOI: 10.1086/656275

19. Schuster, S., Zollinger, S., Lesku, J., & Brumm, H. (2012). On the evolution of noise-dependent vocal plasticity in birds. Biology Letters, 8 (6), 913-916 DOI: 10.1098/rsbl.2012.0676

20. Osmanski, M., & Dooling, R. (2009). The effect of altered auditory feedback on control of vocal production in budgerigars (Melopsittacus undulatus). The Journal of the Acoustical Society of America, 126 (2) DOI: 10.1121/1.3158928

21. Brumm, H. (2004). The impact of environmental noise on song amplitude in a territorial bird. Journal of Animal Ecology, 73 (3), 434-440 DOI: 10.1111/j.0021-8790.2004.00814.x

22. Titze, I. R. (1994). Principles of voice production (pp. 279-306). Englewood Cliffs: Prentice Hall.

23. Cynx J, Lewis R, Tavel B, & Tse H (1998). Amplitude regulation of vocalizations in noise by a songbird, Taeniopygia guttata. Animal behaviour, 56 (1), 107-13 PMID: 9710467

24. Elemans, C., Zaccarelli, R., & Herzel, H. (2008). Biomechanics and control of vocalization in a non-songbird Journal of The Royal Society Interface, 5 (24), 691-703 DOI: 10.1098/rsif.2007.1237

Different personalities. Great tits are not all the same. Image credit: Per Tillman.

My study species in four facts
The last few months have been a blur of conferences, workshops. I attended a fantastic two week course on sensory ecology in Sweden (read about that here and here), I went to London and presented a poster of my work at the annual meeting of the Association for the Study of Animal Behaviour and have recently returned from the Joint meeting of the British Ecological Society and the Société Française d’Écologie in Lille. Time has been flying by and now Christmas is upon us but I wanted to write at least one more post for 2014 so here it is. These are four facts about my wonderful study species the great tit (Parus major).

Great tits have personalities.
My study species.

Every great tit has a unique personality

It’s true, you might think that a bird is just a bird and they are all the same but research has shown that this is not the case. In fact, many species of birds including great tits are now known to have distinct personalities and behave differently to one another, just like humans do. Researchers at Groningen University in the Netherlands found that when they exposed adult great tits to a completely new environment which they had never experienced before some birds were confident and bold and explored their new environments quite happily, while others were much more cautious and did not adapt quickly to the unfamiliar surroundings. Further tests have shown that birds which are bold or shy the first time they are tested tend to stay that way in repeated experiments. These results suggest that personality types are probably fixed characteristics which do not easily change. So, for anyone who’s ever said that animals have personalities, you were right. And it’s not just great tits either; research over the past few years has identified distinct personalities in a huge number of species including mammals, birds, insects and even anemones!

Different personalities. Great tits are not all the same. Image credit: Per Tillman.
Different personalities. Great tits are not all the same. Image credit: Per Tillmann.

City great tits just aren’t as colourful as their countryside cousins

With their bright yellow breast feathers and marked and glossy black plumage great tits are hard to miss at the best of times and are particularly conspicuous during the breeding season when the males are singing their hearts out trying to attract mates. The purpose of the great tits striking colouration is not fully understood but it is likely that females are most attracted to those males which have the brightest and boldest feathers. It is unfortunate then for city birds that their feathers just aren’t as bright as their rural counterparts. A likely explanation for this is that the same carotenoid compound which is used to create the bright yellow colour in great tits feathers is also an anti-oxidant which reduces the physiological stress caused by oxidising pollutants in the environment. As urban birds are known to be exposed to much higher levels of pollutants than rural birds it may be that urban birds are forced to divert carotenoids away from their feathers and use them instead to reduce the levels of stress and cell damage that oxidising compounds can cause. Evidence of this comes from a study at Göteborg University in Sweden which found that the increased oxidative stress faced by urban birds correlated well with reduced levels of carotenoid pigments in their feathers.

Great tits can be incredibly aggressive

They might look cute and friendly but in reality great tits can be incredibly violent, and particularly so during the spring and summer when males are competing for the best breeding territories. During this period male great tits mark their territories by singing at regular intervals and this signal tells other males to move on and find an empty territory of their own. This works most of the time but not always. Sometimes males deliberately invade territories and may attempt to mate with the resident female or even try to force the current territory holder out and take over completely. If this happens the first thing the resident male will do is to sing frequently and loudly in an attempt to show their strength and to drive the intruder away. Should this not work things typically escalate quickly as the resident male flies back and forth across the intruder before coming into full-on physical confrontation. It is not common to see fights in the wild as invaders will usually leave before it comes to actual violence. It does happen sometimes though and you can see the result in these incredible videos.

Great tits sing at higher frequencies in cities than in rural areas

Cities are notoriously noisy places full of cars, people and factories, with aircraft flying overhead and noise from building sites, road repair crews or similar urban development projects. This level of noise is something which wild animals have only had to face in the last century or so, and for species which communicate using sound it could make life in cities very difficult as their vocal signals are easily lost amongst the clatter and din of urban life. Great tits are a good example of a species which might be expected to fare badly in urban habitats as the males rely on communicating by song to defend their territories, show aggression towards rivals and attract mates. To make matters worse, the typical song of a great tit lies in exactly the same frequency range as the background noise of most cities and this means that when a great tit sings its song is quickly swamped by urban noise and cannot be heard by other birds. Yet despite this, great tits are one of the most common song birds seen in city parks and at garden bird feeders so how do they manage to cope with the noise problem? The answer is simple but ingenious. Urban great tits have adjusted their typical song frequency so that they now sing around 350Hz higher. That might not sound like much but experiments have shown that this small adjustments means that the song of urban birds travels above the low-frequency rumble of urban noise and can still be heard by other birds, even at long distances. Genius!

When there are high background noise levels great tits adapt by singing at a higher pitch. Figure from Slabbekoorn and Peet (2003).
When there are high background noise levels great tits adapt by singing at a higher pitch. Figure from Slabbekoorn and Peet (2003).


Every great tit has a unique personality

Dingemanse, N. (2002). Repeatability and heritability of exploratory behaviour in great tits from the wild Animal Behaviour, 64 (6), 929-938 DOI: 10.1006/anbe.2002.2006

City great tits just aren’t as colourful as their countryside cousins

Isaksson, C., Örnborg, J., Stephensen, E., & Andersson, S. (2005). Plasma Glutathione and Carotenoid Coloration as Potential Biomarkers of Environmental Stress in Great Tits EcoHealth, 2 (2), 138-146 DOI: 10.1007/s10393-005-3869-5

Great tits sing at higher frequencies in cities than in rural areas

Slabbekoorn, H., & Peet, M. (2003). Ecology: Birds sing at a higher pitch in urban noise Nature, 424 (6946), 267-267 DOI: 10.1038/424267a

Carnival of Evolution #75: A journey

It’s the start of a new month and that means it’s time again for the Carnival of Evolution and this time it’s a special one, the 75th edition! So all aboard as we journey once more through some of the best evolution based blog posts from around the web.

Carnival of Evolution

Our first stop is the History of Evolution so step out and enjoy the view, there are many good things to see here.

Did Darwin and Wallace plagiarise their ideas on Natural selection from an obscure book on Naval timber? George Beccaloni responds with a resounding “NO”!

Why were Darwin’s ideas accepted so quickly in Victorian England? Piers J. Hale thinks one reason is that they endorsed the liberal Whig politics espoused by great thinkers like ” Darwin’s Bulldog” T. H. Huxley.

Christian Wenande tells us about a treasure trove of Darwin’s barnacle specimens recently discovered in Natural History Museum of Denmark.

You may know that Darwin was good friends with Kew botanist Joseph Hooker but did you know that Hooker was the first person Darwin told about his evolutionary ideas which he described in a letter as ” like confessing a murder”.

What would the world be like if Darwin had never existed? Michal Meyer reviews a new book, Darwin Deleted, that asks just that question.

A treasure trove of Darwin's barnacle specimens have recently been discovered in Denmark (photo credit: Natural History Museum of Denmark)
A treasure trove of Darwin’s barnacle specimens have recently been discovered in Denmark (photo credit: Natural History Museum of Denmark)

All aboard! It’s time to move on to our next stop, Evolutionary Ecology.

Greg Laden talks about a new paper published in Science which shows that dinosaurs have been shrinking for at least 50 million years giving rise to those animals we call birds.

Jonathan Weiner discusses the lifes work of legendary ecologists Peter and Rosemary Grant whose 40 years of research on Galapagos finches showed evolution in action in wild animals.

Why are there so many more species in some regions than in others? GrrrlScientist tells us about a fascinating new study which shows that as available niches fill up the rate of speciation slows down, or even stops.

New reasearch shows that two species of Brazilian ants may have evolved without geographic isolation, although not everyone agrees.

Over at The Loom Carl Zimmer talks about an ancient lineage of air-breathing fish called Bichirs which learn to walk better on land if they are raised out of water, even changing the structure of their bones to improve their walking abilities. This may provide clues as to how fish evolved to walk on land.

Travis Park talks about the evolution of penguins and why it is important to calibrate molecular clocks against the fossil record.

Did these two species evolve in sympatry? Mycocepurus goeldii (left) and Mycocepurus castrator (right) are sister species which occupy the same geographic area (image credit: Christian Rabeling, University of Rochester).
Did these two species evolve in sympatry? Mycocepurus goeldii (left) and Mycocepurus castrator (right) are sister species which occupy the same geographic area (image credit: Christian Rabeling, University of Rochester).

And that’s it for evolutionary ecology this time, now our journey must continue as we head into the wonderful world of Evolutionary Theory.

Evolutionary fitness is often misunderstood the mean “healthiest, strongest, biggest, fiercest, and/or fastest”. Fortunately Stephanie Keep is here to tell us what fitness really means.

Charles Goodnight discusses some of his own work showing that within populations similar individuals tend to cluster together to form groups and this may lead to speciation.

Ben Haller talks about the interactions between empirical and theoretical researchers and asks Should theoretical ideas drive new empirical work to look for the patterns and outcomes predicted by theoretical models?  Or should pure “natural history” observations of the real world drive new theoretical work to explain the patterns and outcomes observed?

Our journey is almost at an end but not quite! Hold tight, we have three more posts to go.

Creationist Michael Behe claims the vertebrate glucocorticoid receptor could not have evolved by darwinian means. Thankfully Larry Moran is on hand to dismantle Behe’s arguments which are neither new nor interesting.

Elena Giorgi provides us with a very topical post about her work using ideas based on the evolution of HIV to develop a vaccine for Ebola.

Bradly Alicea shows us some really neat models of breeding networks that could help us better understand how behaviour and physiology lead to structured genetic variation in populations.

And finally, microbes can evolve the ability to form antibiotic resistant biofilms. Luckily for us people like Elyse Hope are working on ways to thwart them so that we still have effective antibiotics.


And now we come to the end of the line. We hope you had a pleasant journey and will travel with us again soon. The next Carnival will be held in October at Eco-Evolutionary Dynamics, if you would like to submit anything you can submit it on the Facebook page. You can keep up with the Carnival of Evolution via Facebook and Twitter.


Can we use anthropomorphic language in animal behaviour research?

A few months ago during the coffee break at an animal behaviour conference I was talking to a colleague about her research when she told me that to suggest that animals could feel fear or be afraid was anthropomorphism, the mistake of assigning of uniquely human characteristics to other animals. This view is not at all uncommon among practicing scientists and the term anthropomorphism is often extended to include a whole range of behavioural traits and emotions such as impatience, joy, expectation, boredom, anger, happiness or sadness, and yet there is good evidence that these emotions are not unique to humans. For example, dogs have been shown to exhibit jealousy, elephants have empathy, and Capuchin monkeys get visibly angry when treated unfairly as this video shows.

The possibility than animals can think for themselves is also often questioned and yet we know that some animals such as chimps and dolphins have a sense of self. There are also examples of animal behaviours that surely require some degree intelligence and forward planning such as innovative tool use by chimps and deception by ravens.

Deep in thought. Do chimps think like we do?
Deep in thought. Do chimps think like we do?

One of the main arguments against using anthropomorphic language to describe animal behaviours is that there is no way to know how an animal is really feeling, we can only describe what it looks like it’s feeling but not what’s actually happening inside it’s head. But the same is also true of humans, yet no one would question the use of anthropomorphic language to describe human behaviours.

If a person says they are excited we don’t actually know that what they feel as excitement is the same as what you or I feel as excitement, to them it may be a very different thing. All we can do is observe how that person acts and behaves and decide for ourselves if that matches up with our interpretation of what excitement is. The same is true of any emotion. If I say I am or happy or bored how could you tell that what I feel as happiness or boredom is the same as what you or anyone else feels as those things? At some point we have to use our subjective judgement to decide how a person is feeling. Do their actions match our expectations for a happy person? Then we can say they are happy. Do they behave as if they are sad? Then we can say they are sad. If this applies to humans then surely it can also apply to animals, at least in some cases.

I am not suggesting that we should abandon all caution and start using anthropomorphic terms carelessly. What I am suggesting is that so long as we clearly define our terms we should be able to use words like ‘afraid’ or ‘excited’ to describe animal behaviours when those terms well match what we see. If I want to describe anger in animals I should be able to use the word anger so long as I clearly state what I mean by the term and the behaviour that I am describing well matches what most of us would recognise as anger.

Of course, there are cases where using anthropomorphic terms really isn’t appropriate. If I read a paper that described ‘angry’ aphids or ‘jealous’ earthworms I would be dubious that those animals really could feel those emotions. In other cases I would have much less of a problem. Can chimps get angry? I think so. Are rats afraid of predators? It seems likely.

For many scientists, including people who I work with, the fear of using anthropomorphic language seems deeply ingrained and I think this affects how we view the behaviour of animals. Not all animals are mindless automatons that blindly follow their pre-programmed instincts. Many animals, especially among the vertebrates, have complex behaviours and emotions which are best described using the same terms we use for those things in human animals. I think it is time we started describing animals behaviours exactly as we see them. We must define our terms and we must be clear but so long as we are there should be little problem to this approach.

For those that still doubt that animals have can have thoughts and emotions like ours I recommend this TED talk.

For another view I also strongly recommend this post by Jilly at her blog fluffysciences

What do you think? If you have an opinion please leave a comment below.



Male peacock (Parvus cristatus) aim to attract females by out-perform their rivals displays.

Cooperative sperm, killer sperm and the competition for reproductive success

In the closing paragraph of on the origin of species Darwin famously said that nature was a war in which individuals struggle against each other and the environment for survival. However, while survival may be important from an individuals point of view, from an evolutionary perspective mere survival is not enough. Reproduction is what matters and success or failure at producing offspring is what determines an individual’s evolutionary success. Of course, survival is important too, but only when it leads to reproduction.

In most species the reproductive success of females is limited by the rate at which they can produce offspring. When a female is pregnant or carrying eggs she has no choice but to wait until she has given birth or laid her eggs before she can reproduce again, and this can take a long time. Males have no such constraints to their reproductive success and can potentially mate with hundreds of females over their lifetime and raise an enormous number of offspring. The only thing stopping them is that there just aren’t enough females to go around. This shortage of females coupled with the need to reproduce leads to intense, and often aggressive, competition among males for limited mating opportunities.

Male red deer (Cervus elephus) compete for females by fighting
Male red deer (Cervus elephus) compete for mating opportunities by fighting

Male red deer (Cervus elephus) fight for their chance to mate by using their huge antlers to batter their rivals into submission, while male northern elephant seals (Mirounga angustirostris) grow to enormous sizes allowing them to dominate harems of many females and guard them against the advances of smaller, weaker males. Not all species are so aggressive in their tactics. Males of many bird species such as peacocks (Pavo cristatus) and birds of paradise produce fantastic and colourful displays with which they attempt to attract females, as do a large number of insects and fish. In these species, rather than fighting with each other, males try to out-perform and out-class each other in the hope that females will choose them while their rivals are left unwanted on the sidelines. This may seem a more peaceful strategy but make no mistake, although these males don’t actively fight each other the competition between them is every bit as intense as among more aggressive species.

Male peacock (Parvus cristatus) aim to attract females by out-perform their rivals displays.
Male peacock (Pavo cristatus) aim to attract females by out-performing the displays of rival males.

So fighting or displaying are two ways in which males can improve their reproductive chances, but what happens in species in which each female mates with lots of different males in quick succession? How is a male to improve his odds of being the true genetic father of the offspring? Well, as is often the case evolution has found a way and that way is called sperm competition (yes, really).

In species in which females mate promiscuously males compete not just for mating opportunities but also for direct access to eggs. In these cases competition between males happens after mating has occurred as the sperm of multiple males compete with each other within the females reproductive tract as they race towards the eggs. In species in which sperm competition is known to exist an incredible variety of different sperm adaptations have been found, all of which serve to improve the sperms chances of reaching the eggs first.

For individuals of many species adaptation to sperm competition simply means producing more sperm so as to swamp the sperm of their rivals and increase the odds that some of their sperm will make it to the eggs before anyone elses. For other species adaptation to sperm competition is more complex. For example, the wood mouse, Apodemus sylvaticus, has evolved sperm that have a hook-like structure on the head which allows them to intertwine with one another to form long sperm ‘trains’ which are much faster at swimming than individual sperm.

The sperm of the wood mouse (Apodemus sylvaticus). (a) Image of the sperm head with the hook clearly visible. (b) 50 sperm hooked together. (c) A clip from video footage of the sperm train. (d) Another view of the sperm train with an arrow and asterisk marking the position of hooks. (e) One sperm latching onto another. (f) Another view of a sperm hook.
The sperm of the wood mouse (Apodemus sylvaticus). (a) Image of the sperm head with the hook clearly visible. (b) 50 sperm hooked together. (c) A clip from video footage of the sperm train. (d) Another view of the sperm train with an arrow and asterisk marking the position of hooks. (e) One sperm latching onto another. (f) Another view of a sperm hook. Image from Moore et al. (2002).

In a similar and recently discovered case, a team led by Morgan Pearcy of the Université libre de Bruxelles looked for evidence of sperm competition in the desert ant, Cataglyphis savignyiThe queen ants of this species mate with up to 14 males in rapid succession and store their sperm jointly in a special storage organ called the spermatheca. Only those sperm which make it to this storage organ have any chance of fertilising an egg and so competition for access to the spermatheca is intense. In response to this pressure C. savignyi males have evolved highly cooperative sperm that team up into bundles of 50-100 cells which can swim much faster than they could alone and so are better able to outcompete their rivals.

Sperm from the desert ant Cataglyphis savignyi work together to increase their swimming speed. Image from Pearcy et al, (2014).
Sperm from the desert ant Cataglyphis savignyi work together to increase their swimming speed. Image from Pearcy et al, (2014).

It is not just the way sperm behave that can change due to sperm competition, the shape and function of sperm can change too. For example, Philip Byrne and his colleagues from the University of Western Australia found that in a group of Australian frogs those species under the most intense sperm competition produced sperm with the longest tails, possibly to improve their swimming speed. Other species known to have oddly shaped sperm include the water beetle Dytiscus marginalis which has sperm that fuse at the head into pairs with two tails, and the tiny fruit fly Drosophila bifurca which at 6cm long produces the longest sperm on earth.

Some species have taken a more sinister approach to sperm competition and have evolved infertile “parasperm” which contain enzymes capable of breaking down the sperm of rivals. A similar and fantastically named kamikaze sperm hypothesis has even been proposed for humans in which some sperm are adapted to kill the sperm of rivals rather than fertilise eggs. The evidence for this hypothesis is equivocal at best but given the adaptations that have been discovered in other species it is not entirely unbelievable. In fact, given the adaptations that have been discovered so far, almost nothing is completely unbelievable.


Sperm competition by producing large quantities of sperm
Moller, A. (1989). Ejaculate Quality, Testes Size and Sperm Production in Mammals Functional Ecology, 3 (1), 91-96 DOI: 10.2307/2389679

Sperm trains in the wood mouse
Moore H, Dvoráková K, Jenkins N, & Breed W (2002). Exceptional sperm cooperation in the wood mouse. Nature, 418 (6894), 174-7 PMID: 12110888

Cooperative sperm in the desert ant
Pearcy M, Delescaille N, Lybaert P, & Aron S (2014). Team swimming in ant spermatozoa. Biology letters, 10 (6) PMID: 24919705

Sperm competition in Australian frogs
Byrne PG, Simmons LW, & Roberts JD (2003). Sperm competition and the evolution of gamete morphology in frogs. Proceedings of the Royal Society B: Biological Sciences, 270 (1528), 2079-86 PMID: 14561298

The two tailed sperm of the water beetle
Mackie JB, & Walker MH (1974). A study of the conjugate sperm of the dytiscid water beetles Dytiscus marginalis and Colymbetes fuscus. Cell and tissue research, 148 (4), 505-19 PMID: 4836644

The world’s largest sperm in drosophila
Bjork A, Dallai R, & Pitnick S (2007). Adaptive modulation of sperm production rate in Drosophila bifurca, a species with giant sperm. Biology letters, 3 (5), 517-9 PMID: 17594959

Killer ‘parasperm’
Buckland-Nicks, J. (1998). Prosobranch parasperm: Sterile germ cells that promote paternity? Micron, 29 (4), 267-280 DOI: 10.1016/S0968-4328(97)00064-4

Kamikaze sperm
Baker, R., & Bellis, M. (1989). Elaboration of the Kamikaze Sperm Hypothesis: a reply to Harcourt Animal Behaviour, 37, 865-867 DOI: 10.1016/0003-3472(89)90074-2

Criticism of the kamikaze sperm hypothesis
Moore, H., Martin, M., & Birkhead, T. (1999). No evidence for killer sperm or other selective interactions between human spermatozoa in ejaculates of different males in vitro. Proceedings of the Royal Society B: Biological Sciences, 266 (1436), 2343-2350 DOI: 10.1098/rspb.1999.0929