Last week I presented at the University of Wollongong Post-Graduate Conference. I have adapted my presentation below as it was a good overview of my project to date. By way of some background, each year the biology post-graduate students are set a challenge to incorporate an object or personality into their presentations. This year it was Leonardo Dicaprio, so keep an eye out for some celebrity cameos.
So without further ado, let’s start this talk by having a quick think about why animals form groups. We might imagine that in a perfect world, the ideal way to ensure that you maximise your genetic contribution would be to breed as soon as possible and as many times as possible. This would involve leaving the natal territory as soon as possible to pursue individual breeding opportunities.
So why then do we see so many examples in nature of reproductively mature animals which routinely delay or (in extreme cases) completely forgo their own reproductive opportunities in order to join and remain within a group as a subordinate non-breeding member?
The answer is that we don’t really know. We have a few ideas and theories about the costs and benefits of group living, but a general explanation has remained elusive. A major obstacle standing in the way of achieving a general explanation is that there are a lack of studies on marine organisms, which is what my study will focus on.
One of the most promising frameworks with which to study this behaviour is the cooperative breeding framework. This framework contains several hypotheses. I’ll go through just a few which I would like to test and which I will refer to throughout the presentation.
1) Ecological constraints – EC looks at a situation where ecological factors force animals into groups. eg. high predator abundance might cause animals to group in order to obtain a protective benefit through the dilution effect or by making use of discrete habitat patches which provide defence.
2) Life-history – LH hypothesis and EC are closely linked. LH hypothesis looks at inherent life-history traits of animals which might lead to a situation where group formation if more beneficial. For example, animals with long life-spans might cause a breeding habitat to become ‘saturated’. i.e. no vacant breeding ground for new recruits to make use of. In this case it might be more beneficial for the next generation of reproductively mature individuals to join a group and wait for the breeding habitat to become available (avoids conflict).
Most of these studies on birds, insects and mammals to date have either been broad phylogenetic comparisons or fine scale experimental manipulations. Both methods have merit and have in fact been responsible for the huge advances in this field, but there are few studies combining the two approaches. It is important to combine these methods as the broad scale studies can draw correlative conclusions across multiple taxa, but they don’t offer causative explanations. Which is where the experimentation becomes important. However, fine scale experimentation can only focus on a few taxa so the results are often difficult to apply generally across multiple taxa.
So how am I going to approach this question?
I will use a genus of coral reef fishes which show considerable variation in social organisation as a model. Gobiodon species are found in high abundance on tropical reefs. There are over 20 known species.
I will start by conducting a broad phylogenetic comparison of the Lizard island population of Gobiodon. This will involve making a genetic phylogeny of the species at Lizard Island (the above picture shows the phylogeny of the Red Sea population). Phylogenies show how closely related species are to each other. Species radiating from a common node are more closely related to each other, than to species originating from other nodes. The nodes represent some common ancestor. Looking at where sociality occurs on this phylogeny is important as we can see whether the behaviour arose at a single evolutionary point or whether it has evolved multiple times. Looking at the tree above, social behaviour does appear to have evolved multiple times.
Using this phylogeny as a base, I can map ecological and life history traits of each species. This will show which traits the social species share and which the asocial species have in common with each other. I will use this information to identify traits to manipulate experimentally to try to induce social behaviour in an asocial species or vice-versa.
I have chosen these fishes because they show great variation in social structure. for example G. histrio is stubbornly pair forming like Romeo and Juliet. While G. rivulatus forms large social groups more akin to the Great Gatsby, although I suspect that there might be some reproductive shares going to subordinate individuals in the Great Gatsby…
But back to Gobiodon; they are also highly site attached, which makes observing and cataloguing their social systems, ecological traits and life history traits far simpler and experimenting logistically easier with them. Once they have chosen a coral to settle in, that is where they stay. Even when the water level drops below their corals during extreme low tides, they will hunker down and remain within the coral, exposed to air for a couple of hours. They have a high hypoxia tolerance and air breathing ability which enables them to do this.
I have chosen Lizard Island in far north queensland as my study site because:
a) it has an exclusive resort where celebs like leo can be found. Unfortunately, they don’t let the researchers stay here. They tuck us around the corner in this photo.
b) most of the species of gobiodon are known to occur here (and possibly a new species). It’s also worth noting the size of these fishes in the picture below. The fish in the second and fourth pictures on the top row are actually sitting on my gloved hand.
c) There is a well established research station here run by the Australian Museum which makes field work and experiments much easier.
We started this work by finding Gobiodon colonies around lizard island and capturing, counting and identifying the species. To capture the fish we use a mixture of clove oil and ethanol which anesthetises the fish and then we ‘waft’ them out of the coral. Once we capture the fish we hold them in plastic bags until the end of the dive and then take them to a boat to be processed.
To build the genetic phylogeny we need to obtain genetic material. I’ve been taking fin clips from the fish for this. We just snip off about 1/8th of the caudal (tail) fin area while the fish are anesthetised. It’s not uncommon to see these fish with much larger chunks taken out of their fins, usually from conspecific disputes. The fins do grow back quickly so we’re not doing any permanent damage to the fish.
While we have the fish on board and anesthetised, we measure their SL and TL and we give them little tattoos. These are a flourescent elastomer tag inserted under the skin so that we can identify the fish again when we come back in order to determine some life-history traits like dominant turn over rates or growth rates or mortality.
To collect the information about the ecological traits to map onto my phylogeny, I have been seeking out Gobiodon colonies and taking Coral measurements.
To measure habitat saturation we have been using x-transects centred on a colony of Gobiodon. We move along each axis of the transect and catalogue all of the corals known to be inhabited by Gobiodon species. We record whether they were inhabited or not, what they were inhabited by, how many individuals are in each coral, the size and species of each coral.
This gives us an indication of how many and what types of colonies are surrounding the focal colony and how much available habitat there is in the immediate area.
I need to complete the phylogeny now to map these LH and ecological traits and see if there is any correlation between sociality and these traits.
The downside to working in beautiful tropical locations is that they are prone to cyclone activity.
Cyclone Ita came right over Lizard Island in April this year. The photos below are taken from the same sites (left to right) before and after the cyclone. In February, my assistants, and I had tagged about 600 individual fish with a plan to come back in 6 months to re-capture and re-measure these individuals and determine growth rates and dominant turn over rates. I returned in August and found 8 of the original 600 tagged fish.
But moving on, I am still interested in the evolution of social behaviour in these fish, but I will focus more on the evolutionary advantages of sociality or asociality in re-colonising a reef after a disturbance. And I’m hoping that I’ll be able to see that recovery in the data coming out of the x-transects that we’re using to measure habitat saturation.
Anecdotally, there appeared to be more uninhabited corals than there were in February, though I can’t verify this statistically because I used different methods (we were looking at a different question in February). There also appeared to be more juveniles present in August than in February.
Moving on to some preliminary results, these tables show the results from a statistical method called a Generalized Linear Model or GLM for short. Don’t worry about that or all of the technical looking numbers, all you need to know is that a significant result is indicated in red or a highly significant result in yellow. For most of the species above, there is a significant result for average diameter of the coral. That just means that there was a strong relationship between the size of the coral (the predictor) and the number of individuals living within the coral (the response). i.e. the size of the coral could be used to predict the number of fish living within it for those species with a significant result.
I’ve found that the group size of some of the social Gobiodon species is related to the coral size, but not to the size of the largest individual (alpha), which is interesting as Marian Wong and Pete Buston (who presented at UoW a couple of weeks ago) had found that there was a relationship between both coral size and the size of the alpha with the group size in the anemone fish Amphiprion percula. G. oculolineatus does appear to follow this pattern. What I can take away from this is that the determinant of group size is probably species specific and will therefore be more difficult for me to make general conclusions about.
Looking at some of the before-after cyclone data that was comparable, the corals that did survive the cyclone showed positive growth. However when we looked at the site as a whole, the average size of the corals had decreased. This was to be expected since, as you can imagine, a major disturbance like a cyclone would smash up the larger corals into smaller corals. The smaller corals also have less surface area so are more likely to survive a cyclone.
I’ve also found that, as you would expect there was a decline in the coral goby abundance. However, the second graph is more interesting. Some species, like G. erythrospilus, G. rivulatus and G. unicolor appear to be occurring in larger groups post-cyclone. This is possibly an indication that they are in a phase of recruitment.
This will require further exploration so I have another trip planned for January. During this trip I will be re-conducting the x-transects in order to examine this trend across multiple sites. What I will be looking for is whether there is a detectable shift in the goby community. There might be a higher proportion of social species present which could indicate that social species have some kind of advantage in recolonising a reef after a disturbance (or vice-versa).
I will now need to finish the genetic phylogeny and map on the ecological traits that I have collected so far. I have another round of field work booked in for January. I will be conducting more of the cross transects to see if there has been a detectable community shift since my last visit. I also want to set up a pilot experiment looking at the effects of habitat quality and habitat saturation on a subordinate individual’s decision to move or not.
I would also like to set up and run the life history traits work again. i.e. the capturing and tagging component, as this is a really important part of the cooperative breeding framework which I’d like to explore.
To finish up I’d like to say a final thank you to my assistants for their help in the field. It really is a big commitment for them to come and help me out for weeks at a time. Thank you very much! My work could not happen without your help.
If anyone has any questions about my project, please leave a comment below. Thank you!