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Contributions > Papers


Lloyd, G. T., Bapst, D. W., Friedman, M. and Davis, K. E., 2016. Probabilistic divergence time estimation without branch lengths: dating the origins of dinosaurs, avian flight, and crown birds. Biology Letters, 12, 20160609.
     Dryad upload of code and data

Clarke, J., Lloyd, G. T. and Friedman, M., 2016. Little evidence for enhanced phenotypic evolution in early teleosts relative to their 'living fossil' sister group. Proceedings of the National Academy of Sciences USA, 113, 11531-11536.

Bapst, D. W., Wright, A. M., Matzke, N. J. and Lloyd, G. T., 2016. Topology, divergence dates and macroevolutionary inferences vary between different tip-dating approaches applied to fossil theropods (Dinosauria). Biology Letters, 12, 20160237.
     Dryad upload of data and analytical files

Wright, A. M., Lloyd, G. T. and Hillis, D. M., 2016. Modeling character change heterogeneity in phylogenetic analyses of morphology through the use of priors. Systematic Biology, 65, 602-611.

Wang, M. and Lloyd, G. T., 2016. Rates of morphological evolution are heterogeneous in Early Cretaceous birds. Proceedings of the Royal Society B, 283, 20160214.
     Dryad upload of code and data

Lloyd, G. T., 2016. Estimating morphological diversity and tempo with discrete character-taxon matrices: implementation, challenges, progress, and future directions. Biological Journal of the Linnean Society, 118, 131-151.
     Dryad upload of simulation code and tutorial

Benson, R. B. J., Butler, R. J., Alroy, J., Mannion, P. D., Carrano, M. T. and Lloyd, G. T., 2016. Near-stasis in the long-term diversification of Mesozoic tetrapods. PLoS Biology, 14, e1002359.
     Dryad upload


Close, R. A., Friedman, M., Lloyd, G. T. and Benson, R. B. J., 2015. Evidence for a mid-Jurassic adaptive radiation in mammals. Current Biology, 25, 2137-2142.

Wang, M., Zheng, X.-T., O'Connor, J. K., Lloyd, G. T., Wang, X.-L., Wang, Y., Zhang, X.-M. and Zhou, Z.-H., 2015. The oldest record of Ornithuromorpha from the Early Cretaceous of China. Nature Communications, 6, 6987.

Bell, M. A. and Lloyd, G. T., 2015. strap: an R package for plotting phylogenies against stratigraphy and assessing their stratigraphic congruence. Palaeontology, 58, 379-389.
     Dryad upload of R tutorial and example data sets

Brusatte, S. L., Butler, R. J., Barrett, P. M., Carrano, M. T., Evans, D. C., Lloyd, G. T., Mannion, P. D., Norell, M. A., Peppe, D. J., Upchurch, P. and Williamson, T. E., 2015. The extinction of the dinosaurs. Biological Reviews, 90, 628-642.


Brusatte, S. L., Lloyd, G. T., Wang, S. C. and Norell, M. A., 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across dinosaur-bird transition. Current Biology, 24, 2386-2392.
     Dryad upload of R code and data


Lloyd, G. T. and Friedman, M., 2013. A survey of palaeontological sampling biases in fishes based on the Phanerozoic record of Great Britain. Palaeogeography, Palaeoclimatology, Palaeoecology, 372, 5-17.
     Dryad upload of R code and data


Lloyd, G. T., Pearson, P. N., Young, J. R. and Smith, A. B., 2012. Sampling bias and the fossil record of planktonic foraminifera on land and in the deep sea. Paleobiology, 38, 569-584.

Smith, A. B., Lloyd, G. T. and McGowan, A. J., 2012. Phanerozoic marine diversity: rock record modelling provides an independent test of large-scale trends. Proceedings of the Royal Society B, 279, 4489-4495.

Lloyd, G. T., Young, J. R. and Smith, A. B., 2012. Comparative quality and fidelity of deep-sea and land-based nannofossil records. Geology, 40, 155-158.

Lloyd, G. T., Wang, S. C. and Brusatte, S. L., 2012. Identifying heterogeneity in rates of morphological evolution: discrete character change in the evolution of lungfish (Sarcopterygii; Dipnoi). Evolution, 66, 330-348.
     Dryad upload of R code and data

Lloyd, G. T., 2012. A refined modelling approach to assess the influence of sampling on palaeobiodiversity curves: new support for declining Cretaceous dinosaur richness. Biology Letters, 8, 123-126.
     R function and tutorial
     Dryad upload of functions and data
     Piece in Adelaide Advertiser describing me as a "leading palaeontologist"

Lloyd, G. T., Young, J. R. and Smith, A. B., 2012. Taxonomic structure of the fossil record is shaped by sampling bias. Systematic Biology, 61, 80-89.
     Dryad upload of data


Brusatte, S. L., Benton, M. J., Lloyd, G. T., Ruta, M. and Wang, S. C., 2011. Macroevolutionary patterns in the evolutionary radiation of archosaurs (Tetrapoda: Diapsida). Transactions of the Royal Society of Edinburgh: Earth and Environmental Science, 101, 367-382.


Sakamoto, M., Lloyd, G. T. and Benton, M. J., 2010. Phylogenetically structured variance in felid bite force: the role of phylogeny in the evolution of biting performance. Journal of Evolutionary Biology, 4, 733-736.


Brusatte, S. L., Benton, M. J., Ruta, M. and Lloyd, G. T., 2008. The first 50 million years of dinosaur evolution: macroevolutionary pattern and morphological disparity. Biology Letters, 4, 733-736.

Lloyd, G. T., Davis, K. E., Pisani, D., Tarver, J. E., Ruta, M., Sakamoto, M., Hone, D. W. E., Jennings, R. & Benton, M. J., 2008. Dinosaurs and the Cretaceous Terrestrial Revolution. Proceedings of the Royal Society B, 275, 2483-2490.
     Tree and XML files (zip; 1Mb; published version)
     Tree and XML files (zip; 1Mb; 2011 version with correction of minor errors - recommended)
     Reduced Baum and Ragan MRP matrix (NEXUS)
     Reduced Purvis MRP matrix (NEXUS)
     Subsampling code
     Research group supertree page

Brusatte, S. L., Benton, M. J., Ruta, M. and Lloyd, G. T., 2008. Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science, 321, 1485-1488.


Ruta, M., Pisani, D., Lloyd, G. T. & Benton, M. J., 2007. A supertree of Temnospondyli: cladogenetic patterns in the most species-rich group of early tetrapods. Proceedings of the Royal Society B, 274, 3087-3095.

Last updated 7th November 2008.