Reading Evolutionary Biology Group

Selected Publications


Pagel, M. In Press. Language: Why Us and Only Us? Trends in Ecology and Evolution. [Article]


Grollemund, R., Branford, S., Meade, A., Venditti, C. and Pagel, M. 2015. Bantu expansion shows that habitat alters the route and pace of human dispersals. PNAS, 112, 13296-13301. [Abstract & PDF]

Baker, J., Meade, A., Pagel, M. and Venditti, C. 2015. Adaptive evolution toward larger size in mammals. PNAS, 112, 5093-5098. [Abstract & PDF]

Pagel, M. 2015. They'll do more good than harm. Pp 145-148 in (Brockman, J, ed) What to think about machines that think. New York: Harper Perennial.

Hruschka, D, Branford, S, Smith, E, Wilkins, J, Meade, A, Pagel, M and Bhattacharya, T. 2015. Detecting regular sound changes in linguistics as events of concerted evolution. Current Biology, 25, 1-9. [Abstract & PDF]


Calude, A., & Pagel, M. (2014). Frequency of Use and Basic Vocabulary. In L. Filipovic & M. Putz (Eds.), Multilingual Cognition and Language Use: Processing and Typological Perspectives (pp. 45-72). Amsterdam/Philadelphia: John Benjamins Publishing Company.

Pagel, M. 2014. A speaker's map of the mind. In Mapping it Out (H.U.Obrist, ed), London: Thames and Hudson.

Venditti, C., & Pagel, M. (2014). Plenty of room for punctuational change. Trends in Ecology and Evolution, 29, 71-72. [PDF]

Pagel, M. 2014. Foreword to Modern phylogenetic comparative methods and their application in evolutionary biology: concepts and practice (L. Z. Garamszegi, ed). Heidelberg: Springer.


Pagel, M. 2013. Linguistics and the Evolution of Human Languages. Chapter VIII.9, pp 786-794 in the Princeton Guide to Evolution (J. Losos, Editor). Princeton: Princeton University Press.

Pagel, M. 2013. Adapted, yes, but to whom or what? Cliodynamics: The Journal of Theoretical and Mathematical History, 4, 117-119.

Pagel, M. 2013. Why make art? CIsland Magazine: Ideas, Writing and Culture, p. 19, Spring 2013.

Mark Pagel, Quentin D. Atkinson, Andreea S. Calude, and Andrew Meade, May 2013. Ultraconserved words point to deep language ancestry across Eurasia. PNAS, 110, 8471-8476. [Paper]

Eric Lewin Altschuler, Andreea S. Calude, Andrew Meade, Mark Pagel, May 2013. Linguistic evidence supports date for Homeric epics. BioEssays, 35, 417-410. [Abstract] [PDF]


Pagel, M. 2012. War of Words. New Scientist, 8 December 2012, 38-41. Reprinted in New Scientist, The Collection: The Human Story, 2014. [New Scientist Article] [News Article]

Pagel, M. 2012, Wired for Culture: Origins of the Human Social Mind. W. W. Norton & Company. [Publisher] [Facebook]

Pagel, M. 2012. First steps for birds. Nature, 491, 337. [PDF]

Pagel, M. 2012. Adapted to Culture. Nature, 482, 297-299. [PDF]

Pagel, M. 2012. Forked Tongues: the evolution of human languages. Planet Earth (NERC publication), Spring 2012. PDF


TED Global 2011, Edinburgh. Pagel, M. July 2011. How language transformed humanity. [TED Global]

RTVE. Pagel, M. April 2011. Los orĂ­genes de las especies. [RTVE]

Calude, A and Pagel, M. 2011, How do we use language? Shared patterns in the frequency of word-use across seventeen World languages Philosophical Transactions of the Royal Society B 366: 1101-1107. [PDF] This article is also printed in the book "Culture Evolves", Oxford University Press.

Venditti, C., Meade, A. and Pagel, M. 2011. Multiple Routes to Mammalian Diversity. Nature, 479, 393-396. [PDF]

Wilson Sayres, M.A., Venditti, C., Pagel, M. and Makova, K.D. 2011. Do variations in substitution rates and male mutation bias correlate with life history traits? A study of 32 mammalian genomes . Evolution, 65, 2800-2815. [PDF]

Gomes, N.M.V, et al., Pagel, M. and Wright, W. 2011. Comparative biology of mammalian telomeres: hypotheses on ancestral states and the roles of telomeres in longevity determination. Aging Cell, 10, 1-10. [PDF]


Venditti, C. and Pagel, M. 2010. Speciation as an active force in evolution. Trends in Ecology and Evolution, 25, 14-20. [Abstract] [pdf]

Venditti, C., Meade, A. and Pagel, M. 2010. Phylogenies reveal new interpretation of speciation and the Red Queen, Nature 463, 349-352 (21 January 2010) doi:10.1038/nature08630 [Abstract] [pdf]


Organ, C., Janes, D., Meade, A. And Pagel, M. 2009, Genotypic sex determination enabled adaptive radiations of extinct marine reptiles, Nature 461, 389-392 (17 September 2009) doi:10.1038/nature08350; [Abstract] [pdf]

Pagel, M. 2009 Human language as a culturally transmitted replicator. Nature Reviews Genetics, Nature Reviews Genetics 10, 405-415 (June 2009) | doi:10.1038/nrg2560. [pdf]

Pagel, M. 2009. Natural selection one hundred fifty years on. Nature 457, 808-811 (12 February 2009) [Abstract] [pdf]


Pagel M. 2008. Rise of the digital machine. Nature (Essay), 452, 699. [Essay]

Pagel, M. and Meade, A. 2008. Modelling heterotachy in phylogenetic inference by reversible jump Markov chain Monte Carlo. Phil. Trans Roy Soc. B, 363,3955-3964. [Abstract and pdf]

Venditti, C and Pagel, M. 2008. Model misspecification not the node-density artifact. Evolution, 62, 2125-2126. [Abstract] [pdf]

Meade, A. and Pagel, M. 2008. A phylogenetic mixture model for heterotachy. Pages 29-41 in Evolutionary Biology from Concept to Application (P. Pontarotti, ed). Springer Verlag: Heidelberg. [Web Site]

Venditti, C., Meade, A. and Pagel, M. 2008. Phylogenetic mixture models can reduce the node-density artifact. Systematic Biology, 57, 286-293. [Abstract] [pdf]

Pagel, M. and Pomiankowski, A. 2008. Evolutionary Genomics and Proteomics. Sinauer: Sunderland MA. [Web Site]

Atkinson, Q., Meade, A., Venditti, C., Greenhill, S., Pagel, M., 2008, Languages Evolve in Punctuational Bursts, Science, Vol. 319. no. 5863, p. 588, (1 February 2008) [Abstract] [pdf]


Pagel, M., Atkinson, Q., Meade, A., 2007. Frequency of word-use predicts rates of lexical evolution throughout Indo-European history. Nature 449, 717 - 720 (11 October 2007) [Abstract] [pdf]

Organ, C., Shedlock, A., Meade, A., Pagel, M., Edwards, E., 2007. Origin of avian genome size and structure in non-avian dinosaurs. Nature 446, 180 - 184 (08 Mar 2007) [Abstract] [pdf]

Pagel, M., Meade, A., and Scott, D. 2006. Assembly rules for protein interaction networks. BMC Evolutionary Biology, BMC Evolutionary Biology 2007, 7(Suppl 1):S16 doi:10.1186/1471-2148-7-S1-S16. [Abstract] [pdf]

Barker, D., Meade, A., and Pagel, M. 2007. Constrained models of evolution lead to improved prediction of functional linkage from correlated gain and loss of genes. Bioinformatics, doi:10.1093/bioinformatics/btl558 [Abstract] [PDF]


Pagel, M. Venditti, C., Meade, A. 2006 Large Punctuational Contribution of Speciation to Evolutionary Divergence at the Molecular Level. Science, 314, 119-121. [Abstract] [Paper]

Pagel, M. 2006. Darwinian cultural evolution rivals genetical evolution. Brain and Behavioural Sciences, 29, 360- 360.

Venditti, C., Meade, A. and Pagel, M. 2006. Detecting the Node-Density Artefact in Phylogeny Reconstruction. Systematic Biology 55: 637-643

Pagel, M. and Meade, A. 2006. Bayesian analysis of correlated evolution of discrete characters by reversible-jump Markov chain Monte Carlo. American Naturalist, 167, 808-825.

Lester, R.L., Meade, A., and Pagel, M. 2006. The slow road to the eukaryotic genome. BioEssays, 28, 57-64.

Pagel, M and Meade, A. 2006. Estimating Rates of Lexical Replacement on Phylogenetic Trees of Languages. Phylogenetic methods and the prehistory of languages (Peter Forster and Colin Renfrew eds.). McDonald institute Monographs, 173-182.


Sibly, R.M., Barker, D., Denham, M.C., Hone, J. and Pagel, M. 2005. On the regulation of populations of mammals, birds, fish and nsects. Science, 309 607-610 (Perspectives on pp 567-8).

Barker, D. and Pagel, M. 2005. Predicting functional gene links using phylogenetic-statistical analysis of whole genomes. PLoS Computational Biology, 1, 24-31. DOI: 10.1371/journal.pcbi.0010003 [Paper]

Pagel, M. and Meade, A. 2005. Mixture models in phylogenetic inference. Pp 121-142 in O. Gascuel (ed) Mathematics of Evolution and Phylogeny. Oxford: Oxford University Press.

Pagel M, Meade A. 2005. Bayesian estimation of correlated evolution across cultures: A case study of marriage systems and wealth transfer at marriage. Pp 235-256 in Mace R, Holden CJ, Shennan S, editors. The Evolution of Cultural Diversity: a phylogenetic approach. London: University College London Press.


Pagel, M. and Mace, R. 2004 The cultural wealth of nations. Nature, 428, 275-278.

Pagel, M. 2004. Limpets break Dollo´s Law. Trends in Ecology and Evolution, 19, 278-280.

Pagel, M., Meade, A. and Barker, D. 2004 Bayesian estimation of ancestral character states on phylogenies. Systematic Biology, 53, 673-684.

Pagel, M. and Meade, A. 2004. A Phylogenetic Mixture Model for Detecting Pattern-Heterogeneity in Gene Sequence or Character-State Data. Systematic Biology, 53, 571-581.

Pagel, M. and Bodmer, W. 2004. The evolution of human hairlessness: cultural adaptations and the ectoparasite hypothesis. Pp 329-335 in S.P. Wasser (ed) Evolutionary Theory and Processes: Modern Horizons (papers in honour of Eviatar Nevo). London: Kluwer Academic Publishers.


Pagel, M. 2003. Evolutionary biology: Polygamy and parenting. Nature 424, 23-24.

Webster, A.J., Payne, R.J.H. and Pagel, M. 2003. Molecular phylogenies link rates of evolution and speciation. Science, 301, 478.

Pagel, M. and Bodmer, W. 2003. A naked ape would have fewer parasites. Biology Letters (Proc. Roy. Soc. Lond. B. Suppl, 270, S117-S119) DOI 10.1098/rsbl.2003.0041.

Freckleton, R.P., Harvey, P.H., and Pagel, M. 2003. Bergmann´s rule and body size in mammals. American Naturalist, 161, 821-825.


Domb, LG., and Pagel, M. 2002. Sexual selection and costly signaling: reply. Nature, 420, 143.

Pagel, M. and Lutzoni, F. 2002. Accounting for phylogenetic uncertainty in comparative studies of evolution and adaptation. Pp 148-161 in Biological Evolution and Statistical Physics (M. Lässig, and A. Valleriani, eds). Berlin: Springer-Verlag.

Pagel, M. 2002. Modelling the evolution of continuously varying characters on phylogenetic trees: the case of hominid cranial capacity. Pp 269-286 in Morphology, Shape and Phylogenetics (ed, N. MacLeod and P. Forey): Taylor and Francis.

Freckleton, R., Harvey, P.H., and Pagel, M. 2002. Phylogenetic analysis and ecological data: a review of the evidence. American Naturalist,160, 712-726.

Cook, J.M., Rokas. A., Pagel, M. and Stone, G.N. 2002. Evolutionary shifts between host Oak sections and host plant organs in Andricus gall wasps. Evolution, 56, 1821-1830.

Pèrez-Barbería , F.J., Gordon, I.J., and Pagel, M. 2002. The origins of sexual dimorphism in body size in ungulates. Evolution, 56, 1276-1285.

Pagel, M. 2002. Discussion comments on Bayesian phylogenetic inference from animal mitochondrial genome arrangements (B. Larget, D. Simon, and J.B. Kadane) paper read to the Royal Statistical Society meeting of 22 May, 2002. Journal of the Royal Statistical Society, Series B, vol. 64 (2002), p. 757.


Domb, L.G. and Pagel, M. 2001 Sexual swellings advertise female quality in wild baboons. Nature, 410, 204-206 (plus News and Views, p158).

Lutzoni, F. ,Pagel, M., and Reeb, V. 2001. Major fungal lineages derived from lichen-symbiotic ancestors. Nature, 411, 937-940.

Payne, R.J.H. and Pagel, M. 2001. Inferring the origins of state-dependent courtship traits. American Naturalist, 157, 42-50.

Marjon A.M. van Dijk, Ole Madsen, François Catzeflis, Michael J. Stanhope, Wilfried W. de Jong and Pagel, M. 2001. Protein sequence signatures support the ´African clade´ of mammals. Proceedings of the National Academy of Sciences, 98, 188-193.

Jenkins, G.M., Pagel, M., Gould, EA. de A. Zanotto, P.M. and Homes, E.C. 2001. Evolution of base composition and codon usage bias in the genus Flavivirus. Journal of Molecular Evolution, 52 (4): 383-390.


Pagel, M. 2000 Phylogenetic-evolutionary approaches to bioinformatics. Briefings in Bioinformatics, 1, 117-130.

Cézilly, F., Dubois, F., and Pagel, M. 2000. Is mate fidelity related to site fidelity? A comparative analysis in Ciconiiforms. Animal Behavior, 59, 1143-1152.

Pagel, M. 2000. The History, rate, and pattern of World linguistic evolution. In The Evolutionary Emergence of Language (C Knight, M. Studdert-Kennedy, and J. Hurford, eds), Cambridge: Cambridge University Press, pp 391-416.

Pagel, M. 2000. Maximum likelihood models for glottochronology and for reconstructing linguistic phylogenies. In Time-Depth in Historcial Linguistics (C. Renfrew, A. MacMahon, L. Trask eds). Cambridge: The McDonald Institute of Archaeology, pp 189-207.

Pagel, M. 2000 Statistical analysis of comparative data. Trends in Ecology and Evolution, 15, 418.


Pagel, M. 1999 Inferring the historical patterns of biological evolution. Nature (Research Article), 401, 877-884.

Pagel, M. 1999. The maximum likelihood approach to reconstructing ancestral character states of discrete characters on phylogenies. Systematic Biology, 48, 612-622.

Pagel, M. 1999. Mother and father in surprise genetic agreement. Nature, 397, 19-20. (News and Views article)

Olsson, M., Pagel, M., Shine, R., Madsen, T., Doums, C., Gullberg, A., Tegelström, H. 1999. Sperm choice and sperm competition: suggestions for field and laboratory studies. Oikos, 84, 172-175.

Robert, M., Sorci, G., Møller, A.P., Hochberg, M.E., Pomiankowski, A., and Pagel, M. 1999. Retaliatory cuckoos and the evolution of host resistance to brood parasites. Animal Behavior, 58, 817-824.


Pagel, M., Møller, A.P., and Pomiankowski, A. 1998. Reduced parasitism by retaliatory cuckoos selects for hosts that rear cuckoo nestlings. Behavioral Ecology, 9, 566-572.

Dubois, F., Cézilly, F. and Pagel, M. 1998. Mate fidelity and coloniality in waterbirds: a comparative study. Oecologia, 116, 433-440.

Møller, A.P. and Pagel, M. 1998. Developmental stability and signalling among cells. Journal of Theoretical Biology, 193,497-506.

Krakauer, D.C., Zanotto, P.M. de A, and Pagel, M. 1998. Prions' progress: patterns and rates of molecular evolution in relation to spongiform disease. Journal of Molecular Evolution, 47, 133-145.


Pagel, M. 1997. Inferring evolutionary processes from phylogenies. Zoologica Scripta (Journal of the Royal Swedish Academy) 25th Anniversary Special Issue on Phylogenetics and Systematics, 26(4), 331-348.

Lutzoni, F. and Pagel, M. 1997. Accelerated molecular evolution as a consequence of transitions to mutualism. Proc. Natl. Acad. Sci. USA, 94, 11422-11427.

Mace R. & Pagel, M.. 1997. Phylogenies and cultural evolution. Evolution and Human Behaviour 18, 349-351

Pagel, M. 1997. Desperately concealing father: a theory of parent-infant resemblance. Animal Behaviour, 53, 973-981.

Pagel, M. and Krakauer, D.C. 1997. Is the propagation of prion molecules in different hosts an example of Lamarckian inheritance? Trends in Ecology and Evolution, 12,194.

Pagel, M. and Dawkins, M.S. 1997. Peck orders in laying hens: 'futures contracts' for non-aggression. Behavioural Processes, 40, 13-25.

Payne, R.J.H. and Pagel, M. 1997. Why do animals repeat costly signals? Animal Behaviour, 54, 109-119.

Poiani, A. and Pagel, M. 1997. Evolution of avian cooperative breeding: comparative tests of the nest predation hypothesis. Evolution, 51, 226-240.

Cotgreave, P. and Pagel, M. 1997. Predicting and understanding rarity: the comparative approach. To appear in (W. Kunin and K. Gaston, eds) The Biology of Rarity: the causes and consequences of rare-common differences. Chapman and Hall, 237-261.


Krakauer, D.C., Pagel, M., Southwood, T.R.E. and Zanotto, P.M. de A. 1996. Phylogenesis of prion protein. Nature, 380, 675.

Krakauer, D.C. and Pagel, M. 1996. Prions: the genealogy of a toxic protein. Social Biology and Human Affairs, 61, 45-54.

Pagel, M. and Krakauer, D.C. 1996. Prions and the new molecular phenetics. Trends in Ecology and Evolution, 11, 487-488.

Pagel, M. and Payne, R.J.H. 1996. How migration affects estimation of the extinction threshold. Oikos, 76, 323-329.

Krakauer, D. and Pagel, M. 1996. Selection by somatic signals: The advertisement of phenotypic state through costly intercellular signals. Phil. Trans. Roy. Soc. Lond. (B), 351, 647-658.

Payne, R.J.H. and Pagel, M. 1996. When is false modesty a false economy? An optimality model of escalating displays. Proc. Roy. Soc. Lond (B), 263, 1545-1550.

Payne, R.J.H. and Pagel, M. 1996. Escalation and time-costs in displays of endurance. Journal of Theoretical Biology,183,185-193.

Mace, R. and Pagel, M. 1996. Tips, branches, and nodes: seeking adaptations through comparative studies. In L. Betzig (ed) Human Nature: A Critical Reader. Oxford: Oxford University Press, pp 297-310.


Pagel, M. 1995. Sexual selection and oestrous advertisement. Animal Behaviour, 49, 1401-1402.

Krakauer, D.C. and Pagel, M. 1995. Spatial structure and the evolution of honest cost-free signalling. Proceedings of the Royal Society(B), 260,365-372.

Mace, R. and Pagel, M. 1995. A latitudinal gradient in the density of human languages in North America. Proc. Roy. Soc. Lond. (B), 261, 117-121.


Pagel, M. 1994. Evolutionary biology: Parents prefer pretty plumage (News and Views). Nature, 371, 200-201.

Pagel, M. 1994. The adaptationist wager. In Phylogenetics and Ecology (P. Eggleton & R.I. Vane-Wright, eds). London: Academic Press, 29-51.

Pagel, M. 1994. The evolution of conspicuous oestrous advertisement in Old World monkeys. Animal Behaviour, 47, 1333-1341.

Pagel, M. 1994. Detecting correlated evolution on phylogenies: a general method for the comparative analysis of discrete characters. Proceedings of the Royal Society (B) 255, 37-45.

Mace, R. and Pagel, M. 1994. The comparative method in anthropology. Current Anthropology, 35, 549-564.


Pagel, M. 1993. Honest signalling among gametes. Nature, 363, 539-541.

Pagel, M. 1993. Seeking the evolutionary regression coefficient: an analysis of what comparative methods measure. Journal of Theoretical Biology, 164, 191-205.

Pagel, M. 1993. Evolutionary biology: The design of animal signals (News and Views). Nature, 361, 18-20.

Pagel, M. and Harvey, P.H. 1993. Evolution of the juvenile period in mammals. In Juvenile Primates: life history, development, and behavior (M. Pereira & L. Fairbanks, eds). Oxford: Oxford University Press, 28-37.


Pagel, M. 1992. A method for the analysis of comparative data. Journal of Theoretical Biology, 156, 431-442.

Pagel, M. and Harvey, P.H. 1992. On solving the correct problem: Wishing does not make it so. Journal of Theoretical Biology, 156, 425-430.

Pagel, M. and Johnstone, R. 1992. Variation across species in the size of the nuclear genome supports the junk-DNA hypothesis for the C-value paradox. Proceedings of the Royal Society, 249, 119-124.

Pomiankowski, A. and Pagel, M. 1992. Sexual selection and MHC genes. Nature, 356, 293-294.


Pagel, M. and Mace, R. 1991. Keeping the ivory trade banned (Commentary). Nature (Lond.), 351, 265-266.
Reprinted in B. Seyler (1994) Understanding Argument. McGraw Hill, pp 488-493.

Pagel, M.D., Harvey, P.H., and Godfray, H.C.J. 1991. Species-abundance, biomass, and resource-use distributions. American Naturalist, 138, 836-850.

Harvey, P.H., Pagel, M.D. and Rees, J. 1991. Mammalian metabolism and life histories. American Naturalist, 137, 556-566

Pagel, M.D., May, R.M., and Collie, A. 1991. Ecological aspects of the geographic distribution and diversity of mammal species. American Naturalist, 137,791-815.


Elgar, M.A., Pagel, M.D., and Harvey, P.H. 1990. Sources of variation in mammalian sleep. Animal Behaviour, 40, 991-994.

Keymer, A.E. and Pagel, M.D. 1990. Predisposition to helminth infection. In Hookworm Disease: Current Status and New Directions (G.A. Schad and K.S. Warren, eds). Taylor and Francis: London, 177-209.

Pagel, M.D. and Harvey, P.H. 1990. Diversity in the brain sizes of mammal neonates: allometry, energetics, or life history tactic? Bioscience, 40, 116-122.

Blackburn, T.M., Harvey, P.H., and Pagel, M.D. 1990. Species number, population density and body size relationships in natural communities. Journal of Animal Ecology, 59, 335-346.

Trevelyan, R., Harvey, P.H., and Pagel, M.D. 1990. Metabolic rates and life histories in birds. Functional Ecology, 4, 135-141.


Pagel, M.D. and Harvey, P.H. 1989. Taxonomic differences in the scaling of brain weight on body weight among mammals. Science, 244, 1589-1593.

Pagel, M.D. and Harvey, P.H. 1989. Comparative methods for examining adaptation depend on evolutionary models. Folia Primatologia (Special Issue), 53, 203-220.

Harvey, P.H. and Pagel, M.D. 1989. Comparative studies in evolutionary ecology: using the database. In Towards a More Exact Ecology : 75th Anniversary Symposium Volume of the British Ecological Society (J. Whittaker and P.J. Grubb, eds), Blackwell Scientific Publications, 209-230.


Pagel, M.D., Trevelyan, R. and Harvey, P.H. 1988. The evolution of bowerbuilding. Trends in Ecology and Evolution, 3, 288-290.

Pagel, M.D. and Harvey, P.H. 1988. How mammals produce large-brained offspring. Evolution, 42, 948-957.

Pagel, M.D. and Harvey, P.H. 1988. Recent developments in the analysis of comparative data. Quarterly Review of Biology, 63, 413-440.
Reprinted in Japanese as: Pagel, M.D. and Harvey, P.H. 1992. Deta no hikaku bunseki ni okeru saikin no shipno. In Misc. Publ. Nat. Inst. Agro-Environ. Sci. (Japan)

Pagel, M.D. and Harvey, P.H. 1988. The taxon-level problem in mammalian brain size evolution: facts and artefacts. American Naturalist, 132, 344-359.

Harvey, P.H. and Pagel, M.D. 1988. The allometric approach to species differences in brain size. Journal of Human Evolution, 3, 461-472.

Elgar, M.A., Pagel, M.D., and Harvey, P.H. 1988. Sleep in mammals. Animal Behaviour, 36, 1407-1419.


Pagel, M.D. and Greenough, J.A. 1987. Explaining species size-ratios in nature. Trends in Ecology and Evolution, 2, 114-115.