Un-reclaiming the name – I am not a zoologist

zoologist

[Disclaimer – this is just my opinion. I do not speak for everyone at EcoEvo@TCD]

Recently on Twitter there has been a call to “reclaim the name” of Botany accompanied with the hashtag: #iamabotanist. The response has been really cool – lots of different scientists working on different questions have posted pictures of themselves on Twitter, often with their plants. It’s amazing the diversity of researchers out there who identify as botanists.

But why try to reclaim the name Botany? The issue is that Botany as a discipline is seen as rather old-school and irrelevant to current scientific challenges. For these reasons it tends to be unpopular with undergraduates and also with university governing boards. More and more Botany departments are being closed or merged with other departments, and Botany courses are being revamped and renamed to attract more students. Zoology departments are suffering similar fates. Like Botany, Zoology is considered an outdated discipline. It tends to fare better with undergraduate students because there are always people who want to work in a zoo or think they might get to cuddle a panda!

I appreciate what the #iamabotanist campaign was trying to do, but I’m not sure I agree. I work in a Zoology department, but I am not a Zoologist. This isn’t because I think Zoology is irrelevant as a discipline, it’s because I’m far more interested in the questions I’m asking, than in the taxa I use to test my hypotheses. Yes, the mammals I work on are adorable and fascinating, but what drives me as a scientist is trying to understand their evolution and ecology, and how the two things are connected. I’ve mostly worked on mammals so technically I’m a mammalogist. I’m happy with this label, but it’s not what I’d call myself if anyone asked. I’d identify as an (macro)evolutionary biologist, or an evolutionary ecologist. I test my ideas on mammals because these are the group I have most data for, but I’m equally fascinated by insects, bacteria, epiphytic plants, parasitic helminths etc. I think we do a disservice to the science if we focus too much on one taxonomic group.

Zoology and Botany at Trinity are particularly diverse disciplines. We have a couple of “classical” taxonomists/systematists, but also phylogeneticists, landscape ecologists, behavioural ecologists, demographers, evolutionary biologists, conservation biologists, developmental biologists and parasitologists. We teach courses across discipline boundaries, and often the person doing research closest to our own is in the other department. But sadly the Botany-Zoology divide still exists, mostly for reasons of history and geography (we are in separate buildings). This is holding back science, rather than pushing it forward.

Maybe we need to identify as botanists or zoologists (or any other taxon specific -ologists) less often, rather than more often. Forcing general questions and principles down taxon-specific lines seems rather backwards. It also isn’t helpful to our students if they only learn about animals and not the plants they eat, or only learn about plants and not the animals they are being eaten by. This interconnectedness is particularly important in light of the challenges of global change and the current extinction crisis.

So in conclusion, I think animals are cool, but I’m not a zoologist.

Author: Natalie Cooper, ncooper[at]tcd.ie, @nhcooper123

Image Source

Levels of Selection

Okasha 2006

Thanks to the magical (and sometimes frustrating!) technological capabilities of Google+, every fortnight we have international phylo/macro journal club meetings which span three continents and even include elements of time travel (the Australian participants are always in the future!). Among the varied topics we cover, one of our recent sessions was a discussion of Samir Okasha’s book, Evolution and Levels of Selection. Evolutionary biology is an empirical science which also receives attention from philosophers. The two approaches are often difficult to reconcile so Okasha’s book is a welcome bridge for the gap between philosophical theory and practical biology. We mainly focused on the chapter which deals with species selection, clade selection and macroevolution and addresses the units of selection debate.

As a brief background, the debate hinges on the unit(s) and level(s) at which evolution by natural selection operates. Biological structures are inherently hierarchical. As researchers we tend to focus on specific organisational levels, so community ecologists will have very different concerns and interests to cell biochemists. Biological structures are shaped by evolution but the question is at which level(s) in the biological hierarchy does natural selection act? The theory of natural selection is an abstract concept; as Lewontin describes, the tripartite conditions for evolution to occur are phenotypic variation, differential fitness and heritability. So which level of biological organisation satisfies these requirements? Well it seems like it depends on who you read and who influenced your evolution teachers!

In the UK and Ireland we are generally taught evolution from a gene-centric point of view – the Dawkins school of thinking in which selection acts at the level of the individual and the unit of selection is the gene, and only the gene. However, across the pond, there seem to be more proponents and acceptors of higher or multi-level selection theory; the idea (following Stanley, Gould and Eldredge among others) that natural selection is not restricted to genes as the sole units of selection. It’s a confusing debate, especially when it comes to teasing apart the concepts of levels and units of selection; Okasha argues that a gene’s-eye view (genes as the units) can still be adopted for selection acting at various hierarchical levels. Furthermore, concepts of species level selection tend to become confused with group selection – a notoriously controversial concept which is guaranteed to set alarm bells ringing for many people.

Returning to Lewontin’s criteria, the basic idea of species level selection is simple. If species vary in some sort of traits and that variation gives rise to differential extinction or speciation rates, then some types of species will become more common than others. This approach is particularly common from palaeontological or macroevolutionary perspectives. If you’re interested in long-term evolutionary trends such as patterns of differential lineage abundances or extinction and speciation trends, it’s intuitive to treat the species as the level at which selection acts. This highlights a fundamental component of this debate: the gene-only-level of selection is usually advocated by microevolutionists; those who are interested in changes at the genetic level. In contrast, multilevel selection theory receives support from macroevolutionists who, due to their fundamentally different approaches, consider individual species to be their smallest units of interest.

When you think about species selection it is often easy to confound it with clade selection yet Okasha draws a clear distinction between the two concepts. Clades are, by definition, monophyletic; comprised of a single ancestral species and all of its descendant species. Unlike species, clades cannot split to create new clades with ancestor-descendant relationships because any new clade will inevitably be nested within the old clade (the diagram in Okasha’s book makes all of this far clearer than my description!)

Figure
Clade A is part of the larger clade B but it is not the offspring of clade B (offspring must have an independent existence from their parents and be able to outlive them).

Speciation and extinction rates are clearly not uniform; some lineages radiate into many different types of species which enjoy happy evolutionary lives (think of our arthropod-dominated world) while other evolutionary lineages produce fewer species. The question is whether these patterns are the result of species-level, macroevolutionary processes or whether emergent, species-level properties can be explained from selection acting at the genetic level. As an “acid test” for genuine species selection, Okasha proposes Elizabeth Vrba’s view that species selection must in principle (though not necessarily in practice) “be able to oppose selection at lower hierarchical levels”. Otherwise species level selection merely describes processes which can also be explained from a genetic-selection stance. For example, species selection may have been involved in the evolution or maintenance of sexual reproduction; the advantages of sexuality at the species level may have outweighed the two-fold cost of sex at the individual level and therefore favour the evolution of sexual over asexual lineages.

However, there seems to be a general paucity of clear examples which conform to Vrba’s acid test. One intriguing suggestion as to why this may be the case is time. The generation times of species producing new lineages are clearly far longer than the generation times of individuals’ reproduction so perhaps comparatively sluggish species selection processes have not had sufficient time to oppose evolutionary patterns which arise from individual selection?

Confused? It’s an interesting debate but certainly not one for the faint hearted and the fact that each philosopher/scientist/punter on the street seems to have jargon and slightly differing definitions of their own only serves to  cloud the murky waters further. It is, however, interesting to contemplate how our own research backgrounds and the inclinations of our teachers influence our approach to the debate. If you’re interested in these kinds of questions then Okasha’s book is well worth the read or else you could join in with our Phylo/Macro journal club meeting; wherever you are in the world we’re on a Google+ hang out near you!

Authors: Thomas Guillerme (guillert[at]tcd.ie, @TGuillerme) and Sive Finlay (sfinlay[at]tcd.ie, @SiveFinlay)

Image Source: Okasha 2006, Evolution and the Levels of Selection

Seminar series: Tom Ezard, University of Southampton

Forams

Part of our series of posts by final-year undergraduate students for their Research Comprehension module. Students write blogs inspired by guest lecturers in our Evolutionary Biology and Ecology seminar series in the School of Natural Sciences.

This week; views from Sarah Byrne and Sean Meehan on Tom Ezard’s seminar, Birth, death and macroevolutionary consequences.

Splitting Hares – easier said than done?

In a recent talk given by Tom Ezard, a research fellow and evolutionary ecologist, the definition of a species was examined and challenged. While defining a species may seem a simple task for just about anybody and in particular a room full of people with a biology background, the actual definition can be harder to understand when thinking about fossil or species’ records and gaps across time. Ezard highlights that a dynamic approach is needed when discussing speciation and the definition of a species. Claiming that you shouldn’t define a species at one particular moment in time, he details that large gaps in the fossil record make it very difficult to have a fully complete picture about speciation events. In other words, making inferences about speciation events from a certain snapshot in time could overlook the dynamic process of change that occurs over time and give us inaccurate theories about the macroevolution of species.

Following on from the definition of a species, Ezard was interested in the fossil record and how it can give us information about the species record and also, more importantly, about diversity. He was interested in finding out where these gaps in the fossil record had occurred and what impacts they could possibly have. In graphs he provided, it was clear that there was a difference between data over time with more species surges found in recent data in comparison with the past, indicating the number of species has increased over time. However, it’s a little misleading because as time develops we learn more about how to indentify species of have better techniques to do so, it is therefore unclear as to whether or not there has been a big increase in species.

To better explain some complicated parts of the speciation theory, Ezard used a baseball analogy which I was thankful for, showing a picture of various baseballs over time. Ezard explained how techniques improve over time and how the original was very different to the new and modern ball. All of the baseballs of various different ages, textures and shapes remained part of one game (or one species) and that there was no split into a new game (or new species). He stressed that this continuation was very important in understanding macroevolution and when identifying species, that it was vital to look at gaps in the lineage. This brings us back to the fact that the fossil record needs to be examined further and the question of what is meant by a species may need to be redefined. Ezards definition of a species as ‘a single line of descent, a sequence of populations evolving separately from others seems closer to the real definition than previously thought.

Speciation was also a key factor of Ezard’s talk and he was interested in identifying budding speciation events while still being able to identify their ancestors. Two main types of speciation and evolution were discussed in the talk, one type; anagenesis refers to a change along a branch of a phylogeny or the evolution of a gradual change within a species over time. This theory was backed by Darwin and eventually leads to a speciation event. In contrast, cladogenesis, where a population stays stable until a big speciation event happens suddenly and then a splitting occurs between species that ensures they can then not reproduce with each other.

The split can be caused by either biotic or abiotic factors with disagreements regularly occurring between geologists and modern evolutionary biologists over whether the biotic factors (such as competition) or the abiotic factors (such as climate) are the main key drivers affecting species ecology and diversification. So, what is the main driver affecting species ecology and in turn speciation and diversification? Ezard was interested in finding this out.

Using observational studies, algorithmic processes and a multivariate complex approach, Ezard was able to account for ecological differences between species. Lotka’s equation gave an estimate of birth and death models that detailed speciation probability and extinction risk. Species respond differently to global drivers of change and these differences have macroevolutionary consequences. The Red Queen Hypothesis mentioned above, a biotic factor that describes how predator and prey are continually adapting to out-do each other affects species much more so than climate does, and in comparison, climate, an abiotic factor has much more of an effect on extinction.

So, it seems that a combination of both factors are important although they affect both speciation and extinction at different rates. Ezard indicated that, in order to understand diversity, it was first necessary to understand the biotic factors that impact the split and to then devise a model to draw these two areas together. Ezard’s enthusiastic and engaging approach clearly showed his passion for the subject and the interesting topic left me with a lot to think about it.

Author: Sarah Byrne

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Lumpers and Splitters: Apparently they’re not varieties of potato

What is a species? This question seems so fundamental to biology that surely the experts have answered it by now, right? Wrong. Defining a species is a difficult thing, and each new definition seems to come up short in certain criteria. For example Ernst Mayr’s widely used definition of a species: “groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups” completely disregards species which reproduce asexually. For this reason I like Simpson’s evolutionary concept for defining a species and this is precisely what Tom Ezard uses for his work on macroevolutionary dynamics. This concept holds that each species represents a single line of descent; it begins with a speciation event and terminates with extinction. Ezard used the evolution of the baseball to demonstrate this concept. Although the modern baseball is considerably different from its original ancestor, it is still a baseball and there have been no ‘speciation’ events or splits in the lineage to form a new type of ball.

It was Darwin who first coined the term ‘lumpers and splitters’. Lumpers are those biologists who tend to ‘lump’ many ‘species’ in together as one. The splitters are those biologists who like to make as many ‘species’ as possible. In his 1945 work ‘The Principles of Classification and a Classification of Mammals’ George G. Simpson notes rather sardonically: “splitters make very small units – their critics say that if they can tell two animals apart, they place them in different genera … and if they cannot tell them apart, they place them in different species. … Lumpers make large units – their critics say that if a carnivore is neither a dog nor a bear, they call it a cat.” So we can see that this problem is an old one, and that Simpson’s evolutionary concept is very useful for defining species in macroevolutionary studies.

In order to study macroevolutionary dynamics one needs a fairly detailed picture of a clade’s development, and not many organisms provide a suitable fossil record for a detailed study. Fortunately Ezard and his team found the perfect organisms for this purpose; the Foraminifera. These creatures are marine dwelling amoeboid protists. When they die they sink to the bottom and leave behind their calcium shells or tests. They are deposited and preserved on the sea floor and in the right conditions over time can form stratified layers of fossils which give a very complete picture of their evolution over time. Also,the stable isotope ratios of oxygen in the shells can be used to reconstruct palaeo-climatic conditions. These attributes make them incredibly useful in the study of macroevolutionary dynamics.

So, what are the driving forces of speciation? Is there one factor which influences this process above all the others? This is what Ezard and his team set out to investigate. The foraminifera had an interesting story to tell. It was found that incipient species diversify the fastest. This was found to be primarily due to biotic factors or ‘Red Queen’ factors. As a clade grows older it was found that diversification slows due to diversity dependence. However, it was found that extinction is primarily influenced by climatic or Court Jester factors. These findings are important in order to grasp a general understanding of macroevolutionary dynamics. It means that impacts of diversity and climatic fluctuations are not felt uniformly across a phylogeny.  More simply put, it means that the extent of the effect of biotic and abiotic factors on a clade depend on how old it is.

In summary, what Ezard and his team found was that there is no dominant macroevolutionary force, but that, a combination of biotic and abiotic variables drive speciation and extinction. They also found that species’ ecologies are important driving forces in these processes.

Author: Sean Meehan

Image Source: Wikicommons