tree of life Archives

18/02/2013

Palaeo-poetry and placental mammals

Recently Science published O’Leary et al.’s – new load of oil to fuel the burning debate on the origins of placental mammals.

Just to be clear: there is an important distinction between mammals in general that includes many fossils from the Jurassic as well as the extant platypus, kangaroo and your grandma; and placental mammals that includes your grandma and the armadillo (but not kangaroos or the platypus) and no fossil before 65Myr. For readers that are not used to the debate concerning the first placental mammal here’s the main question: did the first placental mammals diversify before or after 65Myr (the important KT boundary)?

(1) After Katie

One view suggests that early mammals lived in the shadow of dinosaurs and the demise of these mighty creatures allowed our rat-like ancestors to take over the earth, the seas and the skies “Save yourself mammals”. This idea was proposed by Simpson in the 1950’s and is supported by the fossil record; many dinosaurs (both big and small) were present before the 65 Myr KT boundary, then a catastrophic meteorite impact marked the KT limit and placental mammals radiated after that. This sequence of events seems to be very straightforward but reality appears not to be so simple. Increasing numbers of mammal species from the late Cretaceous are being discovered, (including rather big ones feeding on dinosaurs) and not many species in general are found in the fossil record before 55Myr when all groups of placental mammals seem to suddenly appear (for a full story see Luo’s 2007 Nature review).

(2) Before Katie

In contrast, another group of people, mainly assisted by molecular dating methods, found out that post-KT placental mammalian diversification may just be an artefact of the fossil record (like Meredith et al 2011 in Science again). Their DNA evidence seems to say that placental mammals evolved before the KT limit and that either palaeontologists failed to find them or else the fossil record failed to preserve them. One major criticism that moderate people argue is that there are still problems associated with molecular dating methods. I won’t go into the details (yes I’m trying hard not to) but molecular dating relies on DNA on the one hand (sampling quality and modelling) and on the fossil record on the other hand. So if the people using DNA criticise the fossil record and want to improve the DNA dating estimates, they have to rely on the same fossil record that they are criticising. The snake bites his own tail.

So what about O’Leary and colleague’s paper? They basically support the first theory (placental mammals evolved after KT). Fair enough, it was led by a number of great palaeontologists and based on a massive morphological data set (~4500 characters introduced as phenomics (from the phenotype) as opposed to genomics (from the genome) data) collected on 40 unambiguous fossils and 46 extant placental mammals. Genomic data based on 26 genes of these extant placental mammals was also included. This paper is the result of an impressive and unique collaborative work, but – Ned Stark from Games of Thrones said “nothing someone says before the word “but” really counts” – but this paper is criticisable…

First of all, the data set: although the morphological data is impressive, the taxa sampling effort seems a bit weak, especially for extant placental mammals. Meredith et al used the same genomic data (26 genes) but based on ~164 mammals to answer the same question. Why couldn’t O’Leary use all of this already published mammal DNA? For the second criticism, I’m just going to quote Yoder’s review published in the same issue “Today, sophisticated theoretical and computational methods are used to estimate and calibrate molecular phylogenetic branch lengths (which represent time). Together with improved methods for integrating fossil and molecular data, dates derived from molecular phylogenies have inched closer to those implied by the fossil record. Is the approach used in the O’Leary et al. study directly comparable to these recent molecular phylogenetic studies? Not really, as it turns out.”

No wonder this paper supports the first theory, it is just a precise and massive analysis of the 40 species of the placental mammals fossil record. Personally, I’m really frustrated by how they managed to publish this paper. Since it’s part of my PhD research, I automatically get excited when I see fossils mixing with extant species so I really hoped this paper would link the two approaches instead of supporting the old fashioned view of evolution (the dinosaurs dying and the mammals taking over). I’d like to think that the history of life is a bit more complex and exciting…

A last comment to justify my title and which will be my main critique to this paper is that O’Leary et al. tried to recreate the “hypothetical placental mammal ancestor”.

As I said, this paper could be seen as a summary of the placental mammal fossil record. So why did they break the first rule that keeps palaeontology away from palaeo-poetry (i.e. going too far with palaeontological hypotheses)? Here they reconstructed a whole creature using their morphological data. What they made was essentially a mean (average) placental mammal (a primitive rat-like creature) – a throw-back to the early stages of palaeontological views of mammalian evolution. What did the ancestor of a duck and a beaver look like? Something in between – a platypus for example? As Olaf Bininda-Edmonds said on Ed-Yong’s Nature post “comparing the two estimates is like comparing “apples and oranges”, they haven’t really done anything to resolve this on-going dispute”.

This paper has also caused controversy on twitter. I’ll just cite two opinions.

Gavin Thomas (@Phalaropus)

“The reconstruction is fun – I’d love to see a picture based on 95% CIs for the ancestral states.”

and Rich Grenyer’s answer (@rich_)

“yes indeed. Something like this” (see our title image).

Many parts of the online science community got excited about this paper, you can see further discussions on Jerry A Coyne’s blog (here and here), on Ed Yong’s one (here and here or there) or else on the twitter feed #placental.

Author

Thomas Guillerme: guillert@tcd.ie

Photo credit

http://aerox21.deviantart.com/

11/01/201311/01/2013

Hey Tree of Life! How’s it growing?

Following the influence of science writers such as S.J. Gould, I always try to look back at the historical perspectives of what I’m studying. These days I’m playing with 3Gb trees so I was delighted by Mindell’s 2013 Systematic Biology publication about the Tree of Life.

The idea of placing species into the so called Tree of Life emerged before the Origin of Species with works such as Augier’s Arbre Botanique (1801) (Fig. 1) and Eichwald’s tree (1829 – possibly inspired by Pallas’s 1766 work) (Fig. 1). But the spreading of such trees began only after publications of Lamarck’s scheme (1809,Fig. 2), Darwin’s famous sketched drawing (1859 – Fig. 3) and Haeckel’s beautiful tree (1866 – Fig. 2). It is only within an evolutionary framework that these representations of the relationships among organisms make sense: the idea of descent with modification.

Figure 1: Augier’s Arbre Botanique (1801) & Eichwald’s tree (1829) – from Mindell 2013 (Fig.1)

Figure 2: Lamarck’s scheme (1809) & Haeckel’s tree (1866) – from Wikimedia Commons

Figure 3: Darwin’s Origin of Species unique figure (1859) – from Wikimedia Commons

From that point, we all know how the story continued; from Darwin’s sketch (Fig. 3) to modern phylogenomics (fig. 4). Our understanding of the Tree of Life progressed from Simpson’s (this one, not this one) cladistic methods for looking at morphological relations among vertebrates, through to the discovery of DNA, the first molecular clock and, eventually, the use of complicated Bayesian stuff. Depictions of the Tree of Life evolved from something like a cypress (a nice, straight tree with Neil Armstrong at the top surrounded by monkeys and mosses and jelly fish near the roots) to a three-rooted shrub full of immense dead branches near the centre. If you look at the figures included in this blog, the changes in our understanding of the tree are clear: a gradual reduction of anthropocentrism and inclusion of microscopic organisms.

Figure 4: Tree of Life from David and Alm (2011) – from David and Alm 2011 (Sup. Fig. 15)

So what should we do next? Should we just expand the dataset until we have all the species and all their genome plotted in the Tree of Life? Hopefully there are still lots of less boring things left to do for researchers working in this area today… Two questions are (in my mind) really important to look at: is the Tree of Life only the result of descent with modification and what should we put in the tree?

For the first question, it appears more and more clear nowadays that the Tree of Life is not really a tree but rather something along the lines of a tree-shaped web. Regarding Archaea and Bacteria alone (the majority of organisms in the tree), it is estimated that at least 81% of their genes have been laterally transferred among lineages at some time in the past. This phenomenon is also becoming increasingly evident among Eukaryotes (even among vertebrates!) and recognition of these events should lead to a more web-shaped “Tree” of Life. Incidentally, it is interesting to note that Batsch recognised this web structure in plants as early as 1802 (Fig. 5).

Figure 5: Batsch’s web of plants (1802) – from Mindell 2013 (Fig.1)

Regarding the second question, asking what can be included in the tree of life comes down to how we determine what is living. I remember one question that a classmates had in my phylogenetic lectures; “What is the out group of the tree of life?” The lecturer had just said that a tree without an out group is not valid. The resulting discussion turned into a really long (and interesting) debate about viruses – the question being whether we should put the viruses in the tree of life? We might define living organisms by entities that can replicate their own DNA. So you could argue that if viruses cannot achieve this independent replication then we should prune them out of our tree. Haha ! But wait, it’s not so easy: although most viruses require host cells for reproduction, so do many other “living” organisms like Richettsia or Chlamydia. In addition, some viruses have many genes involved in DNA replication so how should their self-replication abilities be classed ?

Mindell conclude by quoting Brooks and van Veller (2008): “There are two choices. Do we classify a tree with [lateral transfers], or do we try to classify a [lateral transfer] network? If we wish our classifications to reflect what we think we know about evolution, it seems that we will have to opt for the first alternative.” Does this mean that we should go for a tree shaped web including viruses? Let see how the debate will go on…

Author

Thomas Guillerme : guillert[at]tcd.ie

Photo credits

wikimedia commons

Mindell 2013

David & Alm 2011

18/10/201216/11/2012

The tree on the web

Visualising the tree of life is a challenge for even the most artistically attuned in the scientific community. The problem is the sheer number of species that we need to represent, literally millions. But I think the latest attempt meets the challenge. The developers of OneZoom, the name of the new approach, argue that we need to escape the “paper paradigm”. We should instead make full use of the benefits that digital interactive displays grant us. Worrying that our efforts won’t translate to the printed page is an exercise in Luddism.

So, why is OneZoom so successful? The display takes its cues from Google Earth, the virtual globe, but whereas in Google Earth you move from continents to countries in OneZoom you zoom in from class to species. It’s even better than that because the whole tree has a fractal geometry, the self similar patterns you can see with real trees, which allows for an intuitive zoom function. One critique is that the fractal geometry ends up displaying the groups that diverged earlier (e.g. monotremes) as larger than the more recent groups (e.g. placentals), suggesting some significance when there is none. So far only the mammals are on show but it’s early days yet.

There are a number of other really useful features like the ability to play the evolution of the whole tree backwards and forwards in time so you can see exactly when a given species, genus or what have you, diverged. The potential for science communication as well as research is great. But enough of me talking about what is best seen and check it out for yourself. The details of OneZoom are available in PLOS Biology.

Author

Adam Kane: kanead[at]tcd.ie

Photo Credit

Rosindell J, Harmon LJ (2012) OneZoom: A Fractal Explorer for the Tree of Life. PLoS Biol 10(10): e1001406. doi:10.1371/journal.pbio.1001406