What defines a ‘trait’ in ecology? Part 1 of ‘Ecologists Please Try Harder’

If anyone reading this personally knows me, they will be able to tell you that one of my pet peeves at the moment revolves around the definitions of commonly used words in ecology such as ‘trait’ and ‘niche’. This article is going to focus on what is specifically meant by a ‘trait’, in an ecological sense. I’m going to save ecological niche for another day – in my first article for this blog, I briefly expressed the variable contexts in which it is commonly used, or misused, using a postgraduate conference as an example.

The background behind the desire to do this stems from, in my experience, an apparent inability for anyone to quantify or empirically validate what a trait really is. During my recent MSc, I posed the simple question ‘what is a trait?’ to most of the class, and also to a couple of the professors that taught us (not naming any names, but they both loved insects). Surprisingly, not a single one of them could answer, despite the diversity of associated backgrounds (geologists, palaeontologists, zoologists, entomologists, botanists, etc.)! Considering the rampant use of this term in the literature as, I feel some clarity needs to be given to the precise definition, or concept, of the trait.

To begin with a few simple definitions, typing “define: trait” into Google gave the initial [relevant] definition of “A genetically determined characteristic”; so essentially, any aspect of a phenotype. A definition further down is “Qualities that make one organism different from another”. Further down still, another is “A particular aspect of the phenotype that can be measured or observed directly”. As an sub-ironic side note, Word listed ‘trait’ as a synonym of aspect when I checked during writing this.. Anyway, the second of these is pretty subjective, and also incredibly broad, and is fairly similar to the inverse of homology. The third is an extension of the first, but still provides no resolution in terms of scale – it implies that any aspect of the phenotype, from hair length to number of skin cells on middle finger, can be used as a trait. Similarly, any process, be it chemical, biological or mechanical (or any combination thereof) can also be regarded as a trait. What about in geometric morphometrics? A program I wanted to use recently regarded every single x and y of each co-ordinate a single trait (that was 200 ‘traits’ representing a 2D snout profile!!)

So, if a trait is just any aspect of phenotype, then what is its heuristic value? Morphology is typically broken down into characters and character states, focussed around the concept of biological homology and primarily for use in cladistics. It can also be broken down into functional domains, such as the regions of the axial skeleton (e.g., cervical, dorsal, sacral and caudal vertebrae, and the skull). This has an inherent value, in that each domain has individual parameters that define it, be it from something simple like vertebral count, to the variable strains induced on the skeletal elements during locomotion.

Recently, I discussed the concept of traits briefly with a PhD student at the University of Leicester. He mentioned that the value of the term to ecologists is that it is all-encompassing. I disagree, and, like with homology, a formally well-defined concept, believe that the value of the term lies in the precision of the definition and what it describes. To have a scientific value, the term must imply something that is transparent, and comparable on the necessary levels. I believe that the term ‘trait’ offers neither of these.

To look at occurrences of its academic use, I simply whacked ‘trait’ into Google Scholar, refining it to articles from 2011. It came up with quite a lot of irrelevant garbage, so the search was refined to ‘trait ecology’. Now you’re talking! Out of the host that were found, I selected just a few randomly to see how they used traits in their studies. Below is a list of the papers selected, and an attempt to disseminate what they mean by the term ‘trait’ in each particular and independent study. The references for each are below, and I can provide pdfs if necessary.

Cornelissen et al.

Leaf pH is used as a biochemical plant functional trait. Variation here is attributed to either a species’ genetic makeup or phenotypic plasticity, and additional contributing factors mentioned are seasonal variation and environmentally controlled physiological variability. So there are a host of potential parameters that control trait variation – how do you infer how much of a governing influence one or numerous ones are? We see no empirical or theoretical background again for deciding what a trait is and what is not, and what controls putative trait variation.

Coulson et al.

Utilises a model that classes populations as “fluctuating distributions of phenotypic traits and genotypes”. I think they just use body mass here, combined with genotype in influencing reproductive success. Body mass results from the complex interaction of biological parameters (e.g., feeding efficiency), and climate, as well as latitude and plausibly Cope’s Rule, amongst others. Why not just use ‘body mass’? Why use ‘trait’..? It’s taking something well-defined and quantitative, and dumping it in a bucket with every other aspect of phenotype.

Diniz-Filho et al.

In point 3 of their Abstract (or Summary?!), the first two mentions of traits are immediately followed by “(i.e., body mass)”. They then mention that they were going to deconstruct this trait into phylogenetic and specific components and that each one of these would be a trait too. You can probably see where this is going.. (edit: they actually used ‘niche’ and ‘trait’ in the same sentence at one point. More atrocious than climate change denial).

Hérault et al.

Utilises 17 functional traits (i.e., related to leaf economics, stem economics and life history) in plants, including maximum height, specific leaf area (leaf area/leaf mass), seed mass, wood density. Now, I don’t know about you guys, but when you measure an aspect of morphology, isn’t that called morphometrics? Each one of these putative quantitative traits  is based on a different metric that may or not be independent of each other, and may or may not be related to intrinsic biology as opposed to extrinsic factors.

Laughlin et al.

Functional traits synonymised with “phenotypic properties”, in that traits are environmentally reactive and predict performance. How do you quantify this? Is a trait such as body/leaf/seed mass not the influence of many other smaller-scale parameters? It is a gross over-simplification of phenotypic complexity that closer inspection through character analysis can reveal.

Pavoine et al.

These guys/gals actually mention traits in a phylogenetic context. If you read the following paragraph, it’s pretty obvious that all they’ve done is replaced the word ‘morphology’ with ‘trait’.

However, many approaches have conflated phylogenetic information with trait values (particularly where trait information is unavailable), relying on the underlying hypothesis that closely related species are more likely to have similar traits than distantly related species. Studies that have combined the analyses of traits with phylogenies, in a context of community assembly, have revealed that convergence in trait states can occur among unrelated species.

At this point, they haven’t mentioned what a trait is, or what aspects they are actually studying. Within their analyses, they treat all ‘traits’ as independent, and despite calibrating for phylogeny, means that the study (imo) is methodologically flawed.

Piqueray et al.

Identification of traits that increase the likelihood that species will support an extinction debt. The authors actually make a note of what they define to be a trait, the first time I have ever came across this: “a measurable characteristic of an organism”. This only serves to convey the subjective basis of trait acquisition and analysis; I mean, c’mon, measurable characteristic? Not vague at all.. The authors describe another trait following this, “only present in grazed grasslands” (page 1622). This is not a characteristic, it’s a characteristic of a characteristic. The point again is, if a trait can physically be anything, then what is it’s value as a concept?

Raes et al.

Molecular trait variation in the context of ecosystem processes in microbial communities using environmental shotgun sequencing (i.e., metagenomics). Lateral gene transfer and species dispersal are described as functional traits in a spatial context, or so it seems.

“[investigate] the factors influencing functional dispersal (defined here as the functional effects of species dispersal as well as horizontal gene transfer and phage-mediated gene flow), i.e., the movement of functional traits through geographical space” (page 2).


Note that I’m not saying any of these studies are wrong or flawed due to their use of ‘traits’, I’m simply highlighting the unconstrained variability in which the term is used. There are more references, but they really all follow the same routine. Given the amount of work and debate that goes into understanding and defining the concept of homology, why hasn’t the same attention and scrutiny been given to an almost equally used and valuable concept?

Following this and considering that the term is used so widely, if it is removed or modified, how would people cope? I don’t think it needs strict removal, more just stringently defining as a concept. It needs to be transparent, so that people know when to apply it, and when not to, and also what it can be specifically applied to. It needs to be refined to exclude and include certain factors, and each factor should be purely independent (as in homology). The problem possibly stems from the fact that ‘trait’ has a purely denotative definition; the “any phenotypic aspect” bit. But the connotative definition is unconstrained and poorly defined. I’d like to think that the above examples illustrate this, as if I say ‘trait’ to any of the authors, it is likely they will consider the subjective definition as applies to their personal research.

Also, what of missing data? Phylogeneticists shiver at the very thought of it when constructing a character matrix. So what of missing trait information? If you are regressing twenty ‘traits’ against environmental variables to find a correlation, are you sure that this is enough to capture the entire suite of functionally coupled parameters? Analogous to genomic sequencing, you have to use every piece of available data to reconstruct your trees – the same reasoning should apply to trait data. Except we’ve already seen that a trait is any aspect of phenotype, so it might take a while..

Penultimate point: using body mass as a functional trait is something I’m extremely sceptical about. The opening clause of the abstract from Cooper and Purvis (2010) is probably the most important quote pertaining to this: “Body size correlates with virtually every aspect of species biology” . This is with respect to mammals, and the most significant result of their study is that body mass evolution “has been influenced by a complex interplay among geography, climate, and history”. This should be a strong warning to studies using body mass as a functional trait.

Finally, note that all of the above references are in biological journals. Palaeontologists seem to be doing fine without using ‘traits’! (not that I’m at all biased) Oh, and only a couple of them did any form of phylogenetic regression (i.e., to account for ‘trait’ covariation resulting from common ancestry), so..yeah. But then, I probably don’t get it, not exactly being an expert in the field..

Clarity, constructive comments, and sarcasm would be appreciated.


Edit: http://www.onto-med.de/obml/ws2011/obml2011report.pdf#page=15 Just found this. For additional reading, looks useful. Enjoy! I’m off to play Skyrim..



Cornelissen, J. H. C., Sibma, F., Van Logtestijn, R. S. P., Broekman, R. A. and Thompson, K. (2011) Leaf pH as a plant trait: species-driven rather than soil-driven variation, Functional Ecology, 25, 449-455

Cooper, N. and Purvis, A. (2010) Body size evolution in mammals: complexity in tempo and mode, The American Naturalist, 175(6)

Coulson, T., MacNulty, D. R., Stahler, D. R., vonHoldt, B., Wayne, R. K. and Smith, D. W. (2011) Modelling effects of environmental change on wolf population dynamics, trait evolution, and life history, Science, 334, 1275-1278

Diniz-Filho, J. A. F., Cianciaruso, M. V., Rangel, T. F. and Bini, L. M. (2011) Eigenvector estimation of phylogenetic and functional diversity, Functional Ecology, 25, 735-744

Hérault, B., Bachelot, B., Poorter, L., Rossi, V., Bongers, F., Chave, J., Paine C. E. T., Wagner, F. and Baralato, C. (2011) Functional traits shape ontogenetic growth trajectories of rain forest tree species, Journal of Ecology, 99, 1431-1440

Laughlin, D. C., Fulé, P. Z., Huffman, D. W., Crouse, J. and Laliberté, E. (2011) Climatic constraints on trait-based forest assembly, Journal of Ecology, 99, 1489-1499

Pavoine, S., Vela, E., Gachet, S., Bélair, G. and Bonsall, M. B. (2011) Linking patterns in phylogeny, traits, abiotic variables and space: a novel approach to linking environmental filtering and plant community assembly,  Journal of Ecology, 99, 165-175

Piqueray, J., Bisteau, E., Cristifoli, S., Palm, R., Poschlod, P. and Mahy, G. (2011) Plant species extinction debt in a temperate biodiversity hotspot: community, species and functional trait approaches, Biological Conservation, 144, 1619-1629

Raes, J., Letunic, I., Yamada, T., Jensen, L. J. and Bork, P. (2011) Toward molecular trait-based ecology through integration of biogeochemical geographical and metagenomic data, Molecular Systems Biology, 7, doi:10.1038/msb.2011.6

The Young Systematists Forum, 2011

Having just attended a rather neat conference down in London at the Natural History Museum (NHM), I figured a little summary would make a decent first blog entry. This is my first ever attempt at writing to the general public as well as a scientific audience (or whoever wants to read it really), so if it’s crap, please do let me know.

This year the Young Systematists Forum (YSF), hosted by the Systematics Association, had the greatest turn-out since the conference began aeons ago in the year 2000. The attendees looked eager and ready to devour the broad range of information, despite the starting time of 9.30am (a terrifying prospect for any student). I’d estimate there were at least 100 people in attendance, which is perhaps a little scary for any MSc/PhD student giving a first oral presentation. The multi-disciplinary nature of the field of systematics was apparent straight away – the abstract booklet contained such an immense range of topics, from fossil coelacanths and angiosperm phylogeny, to cladistic methodology and mammoth ecology.

http://www.systass.org/ysf/ <– Webby. Abstract booklet not uploaded yet, but I can send upon request.

Now, as a devout vertebrate palaeontologist, I must say I was overall a little disappointed. Eighteen talks, and only two involved fossils, and just the one poster out of twenty seven, which was mine! Now, given the recently solidified view that fossils and the nature of the fossil record provide critical information in, for example, molecular clock calibration (e.g., Warnock et al. 2011, and refs. within) or macroevolution (Quental and Marshall, 2009), this seemed like a slightly biased series of presentations. That’s all I’m going to say about that, for now.

On to the talks. The first session was hosted by Ellinor Michel, who works with the ICZN in London, and the Zoology Dept. at the NHM. She also lectures on taxonomic principles for the MSc course run at the museum, so was a familiar face to many.

The first talk was on pollen morphology, a stimulating topic to arouse the audience for the next 9 hours of talks. I’m going to digress here. When doing an MSc project, I think that the overall aim is not to do something mind-blowing, but simply to teach yourself new techniques with a dataset that could prove useful for the associated research group. I also think it’s critical that you develop a project personally, instead of simply analysing a small sub-set of your supervisor’s data and falling into the trap of becoming a junior lab assistant. When doing a PhD, you want to be researching a problem or area that is going to have an impact on the scientific community or domain of study. It has to have purpose, a meaning. During very few of these talks, did I see any semblance of the authors knowing why they were doing a particular project. This is my personal feeling, but it may simply be that many chose not to convey this. Setting out with clear-cut aims, objectives and implications is presentation-101 imo though. This leads back on to the first talk.

Descriptive anatomy of pollen with the aim of lineage discrimination. Although well-presented, the speaker failed to reveal what was significant about his group, the Violaceae. Concluding that his studies were, well, inconclusive, he mentioned that pollen needs further systematic evaluation in this group. Despite the multi-genomic analysis that precedes his work on the same group. OK. Unfortunately, he then lost my respect completely. He had constructed a phylogeny to map his few putative synapomorphies on. Now, forgive me for being blunt, but when you are doing a project based on systematics and using cladistic methods, presenting at a systematics conference, you better know what you’re talking about. Especially when you have an entire generation of graduates/masters/doctorate candidates watching you. So when someone asks what optimisation criteria you used to construct your phylogeny, you think back to your basics of cladistics, and answer accordingly. You don’t ask what it means. For those who are not familiar with this terminology, it is nicely summarised in Agnarsson and Miller (2008). To have undertaken research involving cladistics and not know the fundamentals is unforgivable. Rant over.

A quick second digression, but somewhat relevant. About half the speakers (including the above chap) got mixed up between the term ‘cladogram’, and ‘phylogeny’. This is one of my pet peeves, along with confusion between crown and stem groups (http://bit.ly/sptDTk, see this recent article by Mike Keesey). A cladogram is explicitly something created using cladistics, and represents nothing more than a hierarchical branching patterns between taxa. A phylogeny is different; it shows evolutionary trends representing ancestor-descendant relationships between species. The differences are subtle, and I may be completely wrong, (hopefully not), but distinguishing between them and recognising the different implications is important.

Next up was on giant fossil coelacanths! Not much to say here. It was almost a non-stop stream of comparative fish anatomy, in a French accent. Conclusions were that it was possible that gigantism in coelacanths had independently occurred in at least two lineages. Cool.

Following on, bacteria. I’ve got to admit, I don’t know a whole lot about these guys. Some cool images, and horrendous taxonomic nomenclature (Chroococcidiopsis, say it 100 times with a German accent), and then it was time for something completely different. For me, anyway.

Bio-ontologies, an unambiguous way to represent subjective knowledge. It sounds appealing, right? As far as I am aware, the speaker was presenting a new method of describing concepts of words in systematics, to remove subjectivity and refine their semantic power, combined with novel software developed at the Paris NHM. She used the concept of homology as an argument, effectively adding an alternative approach to an already hotly-debated topic. This, of course, was raucously discussed during and after the talk. Break time! And the first poster session. 27 posters to discuss is a bit much, and to be perfectly honest, the idea of reading about the plethora of invertebrate and botanical molecular-based studies was not as appealing as the idea of having a nicotine fix. I’ll just skip to the next session..

Morphometrics! That’s what we like! Kyle, a course-mate from the MSc at the NHM was up, talking about its application in assessing ontogenetic trajectories in mammoths. I believe the software he used was MorphoJ  (available here: http://bit.ly/uoPuzc), which I’m actually not yet acquainted with (we hope to have lunch next week..), having only formally used the tps series (see Links page) and custom software at the NHM designed by Norman MacLeod. It was all good, apart from using positions of particular specimens in Procrustes-transformed data within principal components ordinations to assess changes in shape, which really should have been observed using either a thin-plate spline model (somewhat unreliable) or a strobe-plot. Although, as he used 3-D landmarks, this might have been a little difficult. Regardless, there are alternative methods of visualising shape deformation than speculation (e.g., mapping the highest PCA scores back on to the landmarks, and qualitatively observing deformation trajectories). He got really buggered by a couple of the questions, unfortunately, which was understandable as both questions were nonsense. Firstly, it was asked why ‘discrete character coding’ wasn’t used instead of landmarks. The whole point of the project was to quantitatively assess shape variations, which can’t be done using cladistics. In the future, the results here could be refined to assess the validity of using discrete character states on cranial landmarks, as has been done quite effectively in an MSc project conducted last year on the course, but using Felidae. The second question should have been a piece of piss to answer for someone who’d just done a morphometrics project and having been lectured on it a few months beforehand. Sorry Kyle, but it’s true. “What was the proportion of type 1, type 2, and type 3 landmarks used?” My response would have been “Who gives a toss? (Or why?)”. Type 1 landmarks are those which are topographically homologous, not necessarily biologically homologous. They represent structures which can be explicitly defined on all specimens, such as suture intersections. Type 2 landmarks are those that represent points of geometric significance, such as minima and maxima of curvatures, distal terminations etc. Type 3 are those which are defined or interpolated in terms of other landmarks. The point is that they are all geometrically comparable. So, asking the proportion of these is insignificant. For example, semi-landmarks are a form of secondary landmark, and can contain substantially more information than primary ones when used in abundance to profile outlines. I failed to understand, and glowered at an empty coffee mug for a while.

As a side-note, I love geometric morphometrics. Its statistical power, relative ease in grasping, and broad range of applications make it an invaluable tool in many areas of palaeontology. Imagine, every single character and character state in that matrix it took you so long to create being quantifiable, and the character states formally delimited in a statistically rigorous manner, in such a way that true homologies can not only be irrefutably defined, but also viewed in shape space. I might write an article on this later. Digression x over.

Next up were two vaguely similar talks, both on gastropod morphometrics. The first was excellent! Not surprisingly, it took the award for best talk. A combined biogeographical analysis of thecosomate gastropods using GIS, molecular sequencing and geometric morphometrics allowed the presenter to successfully discriminate geographical gastropod domains. And it came with a meaning! Apparently, these particular gastropods are particularly susceptible to acid dissolution (aragonite shells), so can be used as a proxy for ocean acidification variations. The only thing I’d like to have seen again, is the valve shape changes apparent over these phenotypic and molecular-defined boundaries, and if they could be related to function or environment. The second talk, using a method known as co-ordinate point extended-eigenshape analysis (with Norm’s fingerprints all over it) assessed disparity in Lake Tanganyika’s endemic gastropod fauna. Results seemed to imply that gastropods from analogous but spatio-temporally divergent adaptive radiations occupied near-identical regions of morphospace, which is pretty cool! More was required on casual factors though; for example, were the unoccupied morphospaces less hydrologically stable, or make them look more edible..?

The fourth talk was entertaining. The idea that continental lakes could potentially be used as analogues for island biogeographical patterns seemed appealing, and the sort of crazy idea one would expect from a student from Amsterdam (something in the water, perhaps?). The results, however, seemed to be controversial, and possibly based on incomplete sampling and/or knowledge of the species’ boundaries in the analysed specimens (using CO1, a slowly evolving gene, to assess rapid divergence may have been problematic).

Finally for this session, was an ecological analysis of fiddle crabs in Indonesia, looking at the modes of sympatry between species. Methods were largely observational, using the infamous quadrat methodology, and only preliminary observations were conveyed. Ready for another digression?

The ‘niche’. So far, this word has been used to describe a lake, intra-lacustrine habitats, a quadrat, a mudflat and coeval beach deposits. I posed the question to several conference attendees, and also to the interweb, asking what a ‘niche’ was, and got largely vague responses. It appears that ‘niche’, is understood (or not) by many as a concept to represent any aspect of any environment. Then, clarity. Maybe? John Nudds, palaeontologist, ex-supervisor, Lagerstatten and beer expert, replied “It’s a type of music related to Garage, a British thoroughbred racehorse, or possibly the relational position of an organism’s species“. OK, nice. I trust him. Still, if that’s the definition, does it mean spatially, temporally, sympatrically, allopatrically, ecologically, parasitically? You see where I’m going – it could mean anything, or any combination of things. Therefore, what is it’s use? Considering, unless someone can tell me explicitly what it means, I think it means whatever people want it to. Much like the word ‘trait’. God I hate that word.. Perhaps these vague definitions require some kind of ontological analysis..

Break two. Lunch. Back to the staff canteen after a few months away. Reminiscing, and being miffed at ‘niches’ took up most of that time. Especially as someone had pronounced the word with the ‘ch’ as in ‘leaches’. Practically vomited.

Round three. By now, the lack of sleep and series of misfortunes in getting to London on time was kicking in. Nonetheless, what followed was a series of diverse and interesting talks, despite all being about molecular systematics. Firstly, preliminary results were given regarding the phylogenetic status of economically significant beetles (Elateridae) in Canada. As crop pests, this seemed like quite an important study. But then, what is the point of spending all this time and effort assessing something that you actually just want to kill? And how is a Canadian farmer going to assess the haplotypic affinity of the beetles in his wheat crop from his tractor?

Next up, the most critical talk of the day, as it was directly applicable to almost every other given. Using nymphalid butterflies, it was demonstrated that using identical alignments, you get conflicting trees resulting from using either coalescent or concatenated methods of input. Excellent! Considering this was using a huge data set (12.5kBp, 87 species) using multiple genomic loci, it not only demonstrated the effects of taxonomic and genomic sampling, but also on the impacts of tree construction methodology, something that most other analyses had largely overlooked. This was followed by a largely similar study using Araneae, with largely preliminary work mentioned, so I’ll give it a miss here.

The next one was pretty cool, and took second prize. Phylogeography of a model weed (yep) using haplotype configurations. I think that the main thing to take away, is that there is a geographically-defined system of haplotype differentiation in this particular plant , and secondarily, that if you spend too much time studying at Oxford, accidental quotes such as “explosive seed dispersal that once got in my eye” will be over-looked. I envy the innocent.

That was it. Time for jasmine tea and to tell people about geometric morphometrics and dinosaurs. Got a bit of interest, but most people had resorted to trying to make their own fermentation tanks by this point, with at least two hours to wait until the pub.

On to the final session. It might just have been a trick of the light, but there seemed to be significantly fewer people in the theatre.. The first talk was possibly the most odd of the entire day. I don’t think the chap had actually done anything to present, it was more this concept and series of protocols he wanted to establish to assess population genetics within a couple of Antarctic brittle stars. He had a background in marketing, and it seemed like he was propositioning the audience (comprising one hundred skint students) almost to fund his work. I was confused. It was at this time that my phone battery died, so I had to stop live-tweeting the event (#YSF13), which was also going to be the primary basis for this entry. Ah well.

The next one was reconstructing the ‘phylogeny’ of a particularly speciose clade of angiosperms, the magnoliids. Again, presenting just preliminary work, there wasn’t much to conclude, except that, with the current taxon and genetic coverage, resolution was looking pretty promising, at least with very deep relationships.

Nearly there now. Going on to cladistic methods, the next talk was one of the best-presented of the conference, and with pretty important impacts. They must be feeding those guys at Cambridge something.. The talk revolved around the assessment of ‘hidden’ support in simultaneous and combined analyses. I won’t go into too much depth, but with the creation of intricate new metrics to assess the behaviour of hidden support, I’m looking forward to the publication(s).

Penultimately, an assessment of various competing topologies that reflect the diversification tempo and trajectories of diversification. Now, over the last year, there has been a significant amount of work published within this field (e.g., Stadler (2011), Purvis et al. (2011) and Venditti et al. (2011)). I don’t think they added anything particularly challenging or novel to these three citations, although this may change as additional genomes are acquired and sequenced, and methods of fossil calibration become more robust.

The last talk was on early metazoan evolution using nuclear genomes in extant sponges. In the first bit of the talk, not a single mention of fossils was given, so I had a nap and missed the rest.

And that was it! Rewards were distributed ceremoniously, for some reason in the cramped Bird Gallery (the most easily congested hall in the entire NHM), and free wine was unceremoniously consumed. All in all, there was an eclectic mix of topics, people, and quality throughout the day. Given the pressure, I have to congratulate every single presenter for their courage, and for the most part, unfaltering delivery. Martin Hughes deserves a special mention at the end for asking a question to every single speaker that day. The organisers are again duly noted for, well, organising the event. Although I don’t thank them too much, as they made a typo in my abstract title.

End transmission.



Agnarsson, I. and Miler, J. A. (2008) Is ACCTRAN better than DELTRAN? Cladistics, 24, 1032-1038

Purvis,  A., Fritz, S. A., Rodriguez, J., Harvey, P. H. and Grenyer, R. (2011) The shape of mammalian phylogeny: patterns, process and scales, Proceedings of the Royal Society B, 366, 2462-2477

Quental, T. Q. and Marshall, C. R. (2009) Extinction during evolutionary radiations: reconciling the fossil record with molecular phylogenies, Evolution, 63(12), 3158-3167

Stadler, T. (2011) Mammalian phylogeny reveals recent diversification rate shifts, PNAS, 108(15), 6187-6192

Venditti, C., Meade, A. and Pagel, M. (2011) Multiple routes to mammalian diversity, Nature, 479, 393-396

Warnock, R. C. M., Yang, Z. and Donoghue, P. C. J. (2011) Exploring uncertainty in the calibration of the molecular clock, Biology Letters8, (doi:10.1098/rsbl.2011.