University of Durham

Martin Smith

Laboratory for invertebrate palaeontology and phylogenetics

We are interested in the origins of biodiversity. Our research uses and develops quantitative approaches to reconstruct evolutionary history from the fossil record, with a particular emphasis on the unusual organisms from Burgess Shale-type deposits and their microscopic counterparts, the Small Carbonaceous Fossils.

Timing the pace of the Cambrian Explosion

530 million years ago, the fossil record documents a step change in the complexity and abundance of life on Earth. Our understanding of this ‘Cambrian Explosion’ is fundamentally incomplete. Was it one event or multiple bursts? How did diversification relate to contemporary environmental changes? Can it be explained by presentday evolutionary processes?

The Leverhulme Trust have awarded us funding to develop mathematical techniques that integrate geological observations from across the globe, generating a single record of biological and environmental changes through time. This will establish the rate at which animal biodiversity arose, and the triggers and consequences of this riot of evolution.

To accurately reconstruct the evolutionary events of the Cambrian radiation, we will map the global record of palaeontological, sedimentological and geochemical change onto a common timeline, accounting for preservational biases. The first challenge is to fit independent rock deposits into a single timeline, which currently relies on subjective decision-making. We will construct an objective statistical model for precise stratigraphic correlation, adapting established archaeological techniques to establish the relative ages of rock sequences in the Cambrian and beyond. Secondly, a novel model of morphological evolution will link fossil finds of known ages to points in the tree of life, providing dates and rates of branching points and morphological innovation. Finally, applying our models to a detailed dataset of a major animal lineage will link evolutionary progress to environmental change. Testing the influence of different factors interpreted as contributors to, or results of, elevated evolutionary rates will establish the feedbacks and interactions between the planet and early animal life.

Comparing evolutionary family trees

M.R. Smith

Phylogenetic analysis is the science of reconstructing evolutionary relationships from observations such as morphology and genetic sequences. There are various methods of reconstructing relationships, each with different strengths and weaknesses. As well as working to improve the methods, and the quality of data that is analysed, we have developed new ways to compare trees – a first step to testing how well different methods reconstruct true evolutionary patterns.

Smith, M.R. (2020). "Information theoretic Generalized Robinson-Foulds metrics for comparing phylogenetic trees". Bioinformatics, in the press.

Brazeau, M.D., Guillerme, T. & Smith, M.R. (2019). "An algorithm for morphological phylogenetic analysis with inapplicable data". Systematic Biology, 68 (4): 619–631.

Smith, M.R. (2019). "Bayesian and parsimony approaches reconstruct informative trees from simulated morphological datasets". Biology Letters, 15:20180632.

Solving the riddle of the hyoliths

Danielle Dufault

Hyoliths are one of the most abundant fossils from the Palaeozoic, the era before the dinosaurs. Looking like an ice cream cone with a lid, their identity has long been a mystery: were they aberrant snails, or a long-extinct lineage with no close relatives – a 'failed evolutionary experiment'?

Amazing new fossils from North America and China have finally solved this 175 year old mystery. Preserving the body, not just the shell, these fossils reveal for the first time a crown of tentacles surrounding the mouth – a distinctive feature that links the fossils to modern brachiopods and horseshoe worms. This shows that Hyoliths lived on the sea floor, plucking food particles from passing water. Settling the affinity of hyoliths sheds new light on the origins of animal body plans during the sudden burst of evolution that is the Cambrian Explosion.

Smith, M.R. (2020). "Finding a home for hyoliths". National Science Review, 7 470–471.

Moysiuk, J., Smith, M.R. & Caron, J-B. (2017). "Hyoliths are Palaeozoic lophophorates". Nature 541: 394–397.

Sun, H.-J., Smith, M.R., Zeng, H., Zhao, F.-Z., Li, G.-X. & Zhu, M.-Y. (2018). "Hyoliths with pedicles illuminate the origin of the brachiopod body plan". Proc. Roy. Soc. B, 285: 20181780.

The humble fungus that brought life to land

M. R. Smith

These tiny branching filaments may look unremarkable, but they represent the oldest fossils of terrestrial organisms. New fossils collected in field excursions to Sweden, New York and Scotland allowed me to reconstruct how the hitherto enigmatic organism Tortotubus lived and grew.

These strands represent fragments of extensive subterranean networks, which would have stabilised and nourished the soils in which early plants took root.

Smith, M.R. (2016). "Cord-forming Palaeozoic fungi in terrestrial assemblages". Botanical Journal of the Linnean Society 180 (4): 452–460.

Mini fossils that pack a punch

T.H.P. Harvey / PalAss

Small Carbonaceous Fossils (SCFs) are a diverse suite of non-mineralized microfossils representing bits of early organisms. Widespread in space and time, these tiny elements provide an unrivalled view on the rate and tempo of Cambrian evolution. By establishing the identity of a range of problematic SCFs, my research is telling the evolutionary tales locked up in these beautiful fossils.

Smith, M.R., Hughes, G.M.G., Vargas, M.C. & de la Parra, F. (2016). "Sclerites and possible mouthparts of Wiwaxia from the temperate palaeolatitudes of Colombia, South America". Lethaia 49 (3): 393-397.

Smith, M.R., Harvey, T.H.P. & Butterfield, N.J. (2015). "The macro-and micro-fossil record of the Cambrian priapulid Ottoia". Palaeontology 58 (4): 705–721.

Smith, M.R. & Ortega Hernández, J. (2014). "Hallucigenia’s onychophoran-like claws and the case for Tactopoda". Nature 514 (5722): 363–366.

Caron, J.-B., Smith, M.R. & Harvey, T.H.P. (2013). "Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians". Proceedings of the Royal Society B 280 (1767): 20131613.

The 'weird worm' Hallucigenia

M. R. Smith / Smithsonian

Hallucigenia has always baffled scientists. A 500-million-year-old enigma, this bizarre spiny worm is known from a handful of fossils and just two locations. A study with coauthors Tom Harvey and Jean-Bernard Caron revealed that, in fact, it had relatives all over the world. Whilst examining its defensive spines, we spotted a resemblance with a global family of small spiny fossils: both have a subtle surface ornament and a structure like a stack of ice-cream cones. It seems that quirky Hallucigenia was no recluse: along with its relatives, it formed a cosmopolitan community that spanned the Cambrian seas.

When Hallucigenia was first described in the 1970s, the spines along its back were mistaken for legs, and its head was mistaken for its tail. Now, state-of-the-art electron microscopes have yielded new details from the spines, claws and head of Hallucigenia, exposing its place in life’s evolutionary tree, and how it helps us to reconstruct the common ancestor of all moulting animals.

Smith, M.R. (2017). "Fossil Focus: Hallucigenia and the origin of animal body plans". Palaeontology Online 7 (5): 1–9

Smith, M.R. & Caron, J.-B. (2015). "Hallucigenia’s head and the pharyngeal armature of early ecdysozoans". Nature 523 (7558): 75–78.

Smith, M.R. & Ortega Hernández, J. (2014). "Hallucigenia’s onychophoran-like claws and the case for Tactopoda". Nature 514 (5722): 363–366.

Caron, J.-B., Smith, M.R. & Harvey, T.H.P. (2013). "Beyond the Burgess Shale: Cambrian microfossils track the rise and fall of hallucigeniid lobopodians". Proceedings of the Royal Society B 280 (1767): 20131613.

The 'maybe mollusc' Wiwaxia

M.R. Smith M.R. Smith / Royal Society

The loveable fossil Wiwaxia bears an intimidating coat of scale-mail armour punctuated with lengthy spines. But what is the animal underneath this tough exterior? Could it be a slug, related to the molluscs? Or does it really represent an 'earthworm in disguise'? I've focussed on two clues to assess its affinity. Firstly, its mouthparts have several similarities with the teeth of modern molluscs - and look nothing like worm teeth. Secondly, the animal grows just like some slug-like molluscs that inhabit modern rock pools. However, compelling links with annelid worms suggest that Wiwaxia has a deep evolutionary position and might tell us how both these two body plans became established.

Zhang, Z.-F., Smith, M.R. & Shu, D.-G. (2015). "New reconstruction of the Wiwaxia scleritome, with data from Chengjiang juveniles". Scientific Reports 5: 14810.

Zhao, F.-C., Smith, M.R., Yin, Z.-J., Zeng, H., Hu, S.-X., Li, G.-X. & Zhu, M.-Y. (2015). "First report of Wiwaxia from the Cambrian Chengjiang Lagerstätte". Geological Magazine 152: 378–382.

Yang, J., Smith, M.R., Lan, T, Hou, J.-B., Zhang, X.-G. (2015). "Articulated Wiwaxia from the Cambrian Stage 3 Xiaoshiba Lagerstätte". Scientific Reports 4: 4643.

Smith, M.R. (2014). "Ontogeny, morphology and taxonomy of the soft-bodied Cambrian “mollusc” Wiwaxia", Palaeontology 57 (1): 215-229.

Smith, M.R. (2012). "Mouthparts of the Burgess Shale fossils Odontogriphus and Wiwaxia: implications for the ancestral molluscan radula", Proceedings of the Royal Society B: Biological Sciences 279 (1745): 4287-4295.

The 'squishy squid' Nectocaris

M.R. Smith / Nature

From 1911, only a single specimen of Nectocaris was known. But since the 1980s, Royal Ontario Museum collectors have recovered dozens more. This new material transformed our idea of what Nectocaris looked like, and colleague Jean-Bernard Caron and I estabilshed that it resembled a modern-day squid or cuttlefish. This was astonishing: such cephalopods aren't meant to have evolved until much, much later. This could mean that there’s a huge (200 million year) gap in the fossil record, which would turn the conventional concept of cephalopod evolution on its head. Alternatively, Nectocaris may represent an incredible example of convergent evolution - the independent invention of a modern body plan. This question hinges on how far we trust the fossil record; for now, at least, the jury is out.

Smith, M.R. (2020). "An Ordovician nectocaridid hints at an endocochleate origin of Cephalopoda". Journal of Paleontology 94 (1): 64–69.

Smith, M.R. (2013). "Nectocaridid ecology, diversity and affinity: early origin of a cephalopod-like body plan". Paleobiology 39 (2): 345–357.

Smith, M.R. & Caron, J-B. (2011). "Nectocaris and early cephalopod evolution: Reply to Mazurek & Zatoń". Lethaia 44 (4): 369–372.

Smith, M.R. & Caron, J.-B. (2010). "Primitive soft-bodied cephalopods from the Cambrian", Nature 465 (7297): 469–472.

The 'soggy seaweed' Nematothallus

M.R. Smith / Palaeontology

Tiny fragments of fossilized cuticle, referred to as Nematothallus, have long been implicated in the origin of land plants. Cuticle was an essential adaptation before plants could invade the land – without it, they'd dry up far too quickly. But plants are not the only organisms to have cuticle – so do some seaweeds, lichens, fungi, and animals. How can you tell the difference based on fossil scraps that are less than a millimeter in size?

M.R. Smith / Palaeontology

With co-author Nick Butterfield, I recovered new details of Nematothallus’s reproductive organs, some of which were even preserved with spores attached. Details of these organs allowed our fossils to be reinterpreted as coralline red algae, a type of seaweed that forms parts of coral reefs today. But unlike their modern counterparts, our fossils did not lay down limestone in their skeletons — filling an interesting gap in the evolutionary history of seaweeds.

Smith, M.R. & Butterfield, N.J. (2013). "A new view on Nematothallus: coralline red algae from the Silurian of Gotland". Palaeontology 56 (2): 297-321.

Martin R. Smith

I am interested in the origins of complex animal life, as revealed by the fossil record. Teasing evolutionary information from ancient rocks requires sophisticated mathematical techniques; I develop bioinformatic and phylogenetic methods to better understand the patterns and processes documented by the palaeontological record.

After obtaining an undergraduate degree (BA and MSc) in Natural Sciences from the University of Cambridge, specializing in Earth Sciences and undertaking a Masters’ research project in Palaeobotany with Nick Butterfield, I proceeded to a PhD at the University of Toronto under Jean-Bernard Caron. Based at the Royal Ontario Museum, this was my first encounter with the Burgess Shale fossils that now form a central part of my research portfolio. I subsequently returned to Cambridge to take up a Junior Research Fellowship at Clare College, before moving in 2015 to Durham University, where I am an Assistant Professor (i.e. Lecturer) in Palaeontology.

Martin R. Smith portrait

Dr. Martin R. Smith SFHEA
Assistant Professor (Palaeontology)
Department of Earth Sciences
University of Durham, DH1 3LE
Room ES313

+44 (0) 191 334 2320

Matthias Sinnesael

To reconstruct past interactions between life and the planet, it is necessary to construct a detailed timeline that links evolutionary events with environmental change. This requires the correlation of geochemical and sedimentary observations with an incomplete fossil record.

Matthias is PDRA on a Leverhulme-funded project that will construct Bayesian models of stratigraphy and morphological evolution, which will allow diverse sources of geological data to be integrated into a single timeline, allowing tests of causality and correlation. These models will be applied to the Cambrian Explosion, establishing the character, causes and consequences of this profound evolutionary diversification—thus illuminating the origins of diverse animal-dominated ecosystems and their impact on Earth processes.

Matthias has a PhD on Ordovician cyclostratigraphy, has contributed to cyclostratigraphic methodology, studied K-Pg boundary sections, Devonian cyclostratigraphy, and carbonate geochemistry. Research interests include integrated stratigraphy, paleoclimatology and cyclicity.

Matthias Sinnesael portrait

Dr. Matthias Sinnesael
Postdoctoral research associate
+44 (0) 191 334 2347

Alex Butryn

Alex has an interdisciplinary background, with an undergraduate degree in Natural Sciences and an MPhil in Computational Biology. His Master’s project with Roger Benson used “realistic simulations” to investigate the effect of missing data on phylogenetic inference. These experiences were crucial in developing research skills and shaping his academic interests, which revolve around palaeontology, phylogenetics and applied mathematics. Upon completing his Master’s, Alex worked as a tutor, education consultant and business analyst, before returning to academia.

Alex's Leverhulme-funded PhD research combines evolutionary theory, statistical methods and data from the fossil record to contribute to the understanding and evaluation of the evolutionary interrelationships of extinct taxa.

Alex Butryn portrait

Mr. Alexander Butryn
Postgraduate student

Alavya Dhungana

Alavya has a Leverhulme-funded PhD position that will collect new palaeontological data to generate a new, annotated morphological dataset of representing the Cambrian fossil record of all Ecdysozoan phyla. Analysis by appropriate, stratigraphically constrained models will yield the first detailed, temporally calibrated view of the evolutionary origins of arthropods and related phyla during the Cambrian explosion.

Alavya Dhungana portrait

Mr. Alavya Dhungana
Postgraduate student


Our Leverhulme-funded research project exploring the pace of the Cambrian Explosion is a collaborative effort with Tom Nye (Statistics, Newcastle) and Andrew Millard (Archaeology, Durham).


Previous group members include:

Masters by Research

We welcome applications from undergraduate students looking to undertake MScR training.

Applications are due in early April, with interviews soon thereafter. Applicants will automatically be considered for a Stephen Mills Scholarship, which covers tuition fees at the home/EU rate. We may be able to consider self-funded applications received after this date.

We welcome enquiries or project proposals from potential applicants, who should be on track to receive a strong Bachelor's degree. Overseas applicants may be eligible for Erasmus, Fullbright, Chevening or other scholarships. Durham University also offer means-tested scholarships.

Potential projects

Do clades exhibit an early burst of disparity?

It has often been claimed that major groups are characterized by an early burst of disparity (e.g. Hughes et al. 2013; Oyston et al. 2015). Extrinsic and intrinsic factors have been proposed, but no satisfactory explanation is forthcoming.

In the context of recent work by Budd and Mann, we suggest that two null models may offer a satisfactory account for this claim:

  1. High early disparity is a prerequisite for becoming a ‘major clade’;
  2. High early disparity is an artefact of using phylogenetic datasets, which do not contain autapomorphies.

This project will use the software REvoSim to simulate evolutionary data in order to compare observed patterns of disparity to expected values under unconstrained evolutionary change. This will make it possible to test whether 'early bursts' of evolution represent an authentic feature that requires a macroevolutionary explanation, or whether they represent an necessary prerequisite for survival to the present and thus an emergent feature of natural selection in the presence of extinction.

Why do Cambrian organisms lack mineralization?

The Ediacaran–Ordovician transition sees a profound shift in the role and use of mineralization within organisms. This project aims to test the hypothesis that mineralization intensity corresponds to predation pressure, and, in turn, primary productivity. This provides a possible explanation for the increase in mineralization through the Cambrian explosion, and the selective nature of mineralization in the Ediacaran – with high productivity settings dominated by highly mineralized cloudinid-dominated reefs, in contrast to the non-mineralized nature of deep-water vendobionts.

Using a combination of extant and fossil datasets, it will seek to establish three relationships that, if true, would support the hypothesis:

  1. Is the cost of mineralization (or not mineralizing) related the probability of predation?
  2. Is the abundance of predators a good proxy for the probability of predation?
  3. Does (primary) productivity predict the abundance of predators?

Reading: Wood, R., and Zhuravlev, A. Yu., 2012, Escalation and ecological selectively of mineralogy in the Cambrian Radiation of skeletons: Earth-Science Reviews, v. 115, p. 249–261.

Using δ13Corg to improve the timeline of the Cambrian explosion

This project would involve fieldwork and labwork to obtain a δ13Corganic curve from stratigraphic sections within the UK that span radiometrically dated bentonites, and/or U/Pb dates. This carbon isotope data will be fed into a developing framework of Cambrian stratigraphy established by the Leverhulme project team. This will result in (a) improved resolution and accuracy in the timing of Cambrian deposits in the UK; (b) tighter constraint on correlations throughout the Cambrian period, with implications for the rate and timing of evolutionary change through the Cambrian explosion.

An improved Bayesian framework for global stratigraphic correlation

We will soon have an established Bayesian framework for stratigraphic correlation based on δ13Corg, U/Pb dates and biostratigraphy. We will be looking for MScR and PhD students to develop additional modules that will allow a broader suite of stratigraphic information to be incorporated into this model. Possible focal areas include Strontium isotope data; integration of hiatuses; addition of fossil groups and divergent preservational modes.

Cambrian cyclostratigraphy

The Cambrian Period (~541-485 million years ago) is well known for the Cambrian Explosion. The Cambrian Explosion corresponds with a period in Earth’s history that is characterised by a large diversification of life. An important aspect of reliably reconstructing such changes in biodiversity hinges on our ability to correlate different sections worldwide and construct precise timescales for these events. Establishing such timelines is a challenging task. Early Palaeozoic temporal uncertainties are often in the order of millions of years. One novel way forward is the use of astronomical climate cycles for timescale calibration (a.k.a. cyclostratigraphy or astrochronology). Astrochronology has the potential to construct timescales in the order of tens of thousands of years. This project will investigate the potential astrochronology of the Cambrian Lie De Vin Formation in the Moroccan Anti-Atlas. We will develop an understanding of the sedimentology, integrated stratigraphy and apply numerical time-series analysis techniques to test the hypothesis of the recording of an original astronomical signal in this section.

All the better to see you with: controls on visual acuity in Cambrian communities

The evolution of vision offered significant advantages to the earliest organisms to bear eyes. Simple physical considerations allow visual acuity to be accurately reconstructed from suitably preserved fossils. This project will constrain the optical capabilities of soft-bodied fossils from Cambrian Lagerstätten, using measurements obtained from published specimens. Visual acuity will be linked to environmental factors to establish the primary evolutionary forces controlling vision in the Cambrian, with a view to better constraining the ecology and environmental parameters of early animal ecosystems.

How does morphology evolve?

Understanding the biological processes by which observable phenotypic evolution occurs is a fundamental goal of macroevolutionary study – and modelling morphological evolution is key to reconstructing the relationships and evolutionary history of life.

This project will test a series of hypotheses of how morphological variety originates by identifying patterns of evolution in a collection of hundreds of existing morphological datasets from a wide range of vertebrate, invertebrate and non-metazoan groups. The successful student will establish whether widespread tenets of palaeontological phylogenetics (e.g. equal weights parsimony) can truly account for observed morphological diversity, improving our understanding of how evolutionary processes lead to morphological novelty. This in turn will improve models of phylogenetic inference, and our understanding of evolutionary history.

Reliability of phylogenetic results

Much of modern biology is underpinned by frameworks of relationships arising through phylogenetic analysis. As such, it is more important than ever to be able to distinguish results that are supported by strong evidence from those likely to be overturned as new data accumulate.

Hillis and Huelsenbeck (1992) propose the skewness of tree lengths as a proxy for the reliability of phylogenetic results obtained from a dataset. This project will implement this reliability measure and apply it to palaeontological data matrices in order to resolve disagreements between competing phylogenetic studies and to establish the relative contributions of fossil and extant taxa to phylogenetic reconstructions.


We are currently looking to recruit excellent students to our postgraduate research programmes.

A diverse range of projects are available, based on independent fieldwork, museum collections, and laboratory-based data analysis.

The most popular routes to PhDs in the Palaeoecosystems group are:

  • RCUK-funded projects (home students): these are advertised each autumn through the NERC Iapetus doctoral training program; applications are due by early January. If you'd like to propose your own project, get in touch over the summer.
  • Durham Doctoral Studentships: these highly competitive positions allow a home or overseas student to take up a project advertised under the RCUK umbrella, or one of their own devising. Applicants are advised to contact potential supervisors well in advance of the mid-January deadline.
  • Personal scholarships: many overseas students are eligible for international funding routes, which may be tied to an advertised project or to a project devised jointly with a potential supervisor – feel free to get in touch.

Potential projects

Convergence or cryptic evolution? The origins of the cephalopod body plan

500 million years ago, life on earth was fundamentally transformed by the geologically rapid emergence of complex animal-dominated ecosystems. This ‘Cambrian Explosion’ permanently altered the dynamics of biology and geology on a planetary scale – but what evolutionary processes led to the sudden dominance of macroscopic organisms, and their concomitant interactions with the oceans, sediment and biosphere?

Cephalopods – a charismatic group of molluscs that includes cuttlefish, octopus, and the extinct ammonites and belemnites – offer an illuminating perspective on the dynamics of Cambrian evolution. The conventional view interprets a suite of snail-like Cambrian fossils as representing the gradually evolving roots of the cephalopod lineage. This view has recently been challenged by Nectocaris, an early Cambrian fossil that strikingly resembles a modern squid. If Nectocaris is a cephalopod, then cephalopods evolved from a non-mineralized ancestor in the height of the Cambrian explosion, with an early evolutionary history that largely escaped the fossil record.

This controversial interpretation supports an interpretation that sees anatomical blueprints of the major animal lineages becoming ‘fixed’ in the earliest Cambrian, and overturns the core tenets of cephalopod evolution: reconstructing the ancestral cephalopod as internally shelled, physiologically energetic, and jet propelled contradicts the conventional viewpoint of evolutionary ‘progress’ from sluggish forebears to ‘advanced’ modern taxa.

If Nectocaris is not a cephalopod, however, then a squid-like body plan must have arisen twice, with today’s squid ‘reinventing’ the blueprint established by Nectocaris. This remarkable degree of convergence implies that physical processes constrain biological possibility, with evolution only able to arrive at a finite number of discrete ‘body plans’.

This PhD proposal will conduct the first quantitative test of the relationships of Nectocaris. This objective evaluation will evaluate its implications for cephalopod origins, and for the nature of evolutionary innovation more generally.

See full project description and how to apply.

An improved Bayesian framework for global stratigraphic correlation

We will soon have an established Bayesian framework for stratigraphic correlation based on δ13Corg, U/Pb dates and biostratigraphy. We will be looking for MScR and PhD students to develop additional modules that will allow a broader suite of stratigraphic information to be incorporated into this model. Possible focal areas include Strontium isotope data; integration of hiatuses; addition of fossil groups and divergent preservational modes.

Postdoctoral Fellowships

We invite potential postdoctoral fellows with compatible research interests to join the Palaeoecosystems group, and welcome the opportunity to support any potential applications. Besides national and international schemes (e.g. NERC/Leverhulme/Royal Society), Durham University funds its own Addison Wheeler Fellowships.

We will soon be recruiting a postdoctoral research assistant to work on the stratigraphy of the Cambrian explosion, with an anticipated start date in 2021/22; contact Martin Smith for details.

Undergraduate Research

A range of undergraduate-level research projects are available for Durham and non-Durham students who wish to gain palaeontological experience. Bursaries may be available from the Palaeontological Association (Deadline: late February).

As a Senior Fellow of the Higher Education Academy, I apply evidence-based pedagogical theory to develop an engaging, balanced and outcome-directed teaching approach.

Jiahe Max Luan

I currently lead a second year undergraduate course in Palaeoecosystems, contribute to Palaeobiology teaching at a first-year introductory level, and supervise third- and fourth-year research projects.

We have produced a variety of software packages to facilitate phylogenetic analysis and the interpretation of results.

Ternary: create ternary plots.

Packages for calculating and evaluating distances between phylogenetic trees: TreeDist, TBRDist, Quartet.

TreeSpace: interactive user interface for the creation, visualization and evaluation of tree landscapes.

TreeTools: utilities for the creation, modification and analysis of phylogenetic trees.

TreeSearch: phylogenetic tree search in R, with an appropriate treatment of ‘inapplicable’ characters.