Dr Robert Mill: Gymnosperm Systematist

email: Robert Mill

Phone: +44 (0)131 248 2935

Fax: +44 (0)131 248 2901

Research interests

After many years working on RBGE's Flora projects, particularly Turkey, Arabia, Bhutan, and China, I began monographic research on the southern hemisphere conifer families Araucariaceae and Podocarpaceae. I am currently researching the biology and systematics of Araucaria and revising genera of Podocarpaceae. I have also recently been involved in a project that has helped to unravel some of the taxonomic problems in Chinese and Himalayan species of Taxus.

Current research

  • Systematics of Podocarpaceae using traditional and modern techniques
  • Cuticular micromorphology as a systematic tool in Podocarpaceae
  • Molecular phylogeny of Podocarpus, particularly of South America
  • Systematics and biology of Araucaria



MILL RR, Ruhsam M, Thomas P, MF Gardner &  PM Hollingsworth (2017) Araucaria goroensis (Araucariaceae), a new Monkey Puzzle from New Caledonia, and nomenclatural notes on Araucaria muelleri. Edinburgh Journal of Botany:doi: 10.1017/S0960428617000014.


MILL RR (2016) A monographic revision of Retrophyllum (Podocarpaceae) Edinburgh Journal of Botany. 73(2):171-261

Ruhsam M, Clark A, Finger A, Wulff AS, MILL RR, Thomas P, Gardner MF, Gaudeul M, Ennos RA and PM Hollingsworth (2016). Hidden in plain view: cryptic diversity in the emblematic Araucaria of New Caledonia. American Journal of Botany 103(5): 888-898


MILL RR (2015) A monographic revision of the genus Podocarpus (Podocarpaceae): III. The species of the Central America and Northern Mexico Bioregion Edinburgh Journal of Botany 72(2): 243-341

MILL RR (2015) A monographic revision of the genus Podocarpus (Podocarpaceae): II. The species of the Caribbean Bioregion Edinburgh Journal of Botany 72(1): 61-185

Quiroga MP, Mathiasen P, Iglesias A, MILL RR & AC Premoli (2015) Molecular and fossil evidence disentangle the biogeographical history of Podocarpus, a key genus in plant geography. Journal of Biogeography (early view): doi:10.1111/jbi.12630


Kranitz ML, Biffin E, Clark A, Hollingsworth ML, Ruhsam M, Gardner MF, Thomas P, MILL RR, Ennos RA, Gaudeul M, Lowe AJ and Hollingsworth PM (2014). Evolutionary Diversification of New Caledonian Araucaria. PloS ONE 9: e110308

MILL RR (2014) A monographic revision of the genus Podocarpus (Podocarpaceae): I. Historical Review. Edinburgh Journal of Botany 71(3): 309-360


Möller M, Gao LM, MILL RR, Liu J, Zhang DQ, Poudel RC and Li DZ. (2013) A multidisciplinary approach reveals hidden taxonomic diversity in the morphologically challenging Taxus wallichiana complex. Taxon 62 (6) 1161-1177


Mao K, Milne RI, Zhang L, Peng Y, Liu J, THOMAS P, MILL RR and S Renner (2012) Distribution of living Cupressaceae reflects the breakup of Pangea. PNAS 109: 7793–7798

MILL, R.R. and Whiting, M. (2012) Podocarpus orarius (Podocarpaceae), a new species from the Solomon Islands and a taxonomic clarification of Podocarpus spathoides from Malaysia. Gard. Bull. Singapore 64 (1) 171-193


Christenhusz, M.J.M., Reveal, J.L., Farjon, A., Gardner, M.F., MILL, R.R. & Chase, M.W.  (2011) A new classification and linear sequence of extant gymnosperms. Phytotaxa 19: 55–70

Stark Schilling, D. M. & MILL, R. R. (2011) Cuticle micromorphology of Caribbean and Central American species of Podocarpus (Podocarpaceae). International Journal of Plant Sciences 172 (5): 601-631


MILL, R. R. (2010) [1938] Proposal to reject the name Torreya bogotensis (Gymnosp., incertae sedis, Podocarp.). Taxon 59(3): 975-976 

MILL, R. R. & Stark Schilling, D. M. (2010) Typification and nomenclature of Podocarpus angustifolius Griseb. and Podocarpus aristulatus Parl. (Podocarpaceae). Taxon, 59(3): 935-956.

Stark Schilling, D. M. & MILL, R. R. (2010). [1937] Proposal to conserve Podocarpus aristulatus Parl. (Podocarpaceae) with a conserved type. Taxon, 59(3): 973-975


MILL, R. R. & Stark Schilling, D. Cuticular micromorphology of Saxegothaea Lindl. (Podocarpaceae). Bot. J. Linn. Soc., 159: 58-67


Gao, L.-M., Möller, M., Zhang, X.-M., Hollingsworth, M.L., Liu, J., MILL, R. R., Gibby, M., Li D.-Z. (2007). High variation and strong phylogeographic pattern among cpDNA haplotypes in the Taxus wallichiana complex (Taxaceae) in China and North Vietnam. Molecular Ecology 16: 4684-4698

Möller, M., Gao, L.-M., MILL, R. R., Li D.-Z, Hollingsworth, M.L. & Gibby, M. (2007). Morphometric analysis of the Taxus wallichiana-complex based on herbarium material. Bot. J. Linn. Soc. 155: 307-335.


MILL, R. R.(2006). A new name for Podocarpus rotundus de Laub. (Podocarpaceae). Edin. J. Bot. 63: 81-83.

MILL, R. R. & Farjon, A. (2006). (1710). Proposal to conserve the name Xanthocyparis against Callitropsis Oerst. (Cupressaceae). Taxon 55: 229-231.



Brummitt, R. K., MILL, R. R. & Farjon, A. (2004). The significance of ‘it' in the nomenclature of three Tasmanian conifers: Microcachrys tetragona and Microstrobos niphophilus (Podocarpaceae), and Diselma archeri (Cupressaceae). Taxon 53(2): 529-539.

MILL, R. R. & Hill, R. S. (2004). Validations of the names of seven Podocarpaceae macrofossils. Taxon 53: 1043-1046.

MILL, R. R., Möller, M. Glidewell, S. M., Masson, D. & Williamson, B. (2004). Comparative anatomy and morphology of fertile complexes of Prumnopitys and Afrocarpus species (Podocarpaceae) as revealed by histology and NMR imaging, and their relevance to systematics. Bot. J. Linn. Soc. 145: 295-316.

MILL, R. R. & Weston, P. (2004). (1613-1614) Proposals to reject the names Polypodiopsis and Polypodiopsis muelleri (Plantae vasculares, incertae sedis). Taxon 53(1): 203-205.

MILL, R. R. (2004). (1612) Proposal to conserve Podocarpus affinis Seem. (Recent Podocarpaceae) over Podocarpus affinis A. Massal. (fossil Podocarpaceae). Taxon 53(1): 201-202

Muller, E. M., Barker, N. P. & MILL, R. R. (2004). A Yellowwood by any other name: Molecular systematics and the taxonomy of Podocarpus and the Podocarpaceae in Southern Africa. South African Journal of Science 100: 629-632.



Collins, D., MILL, R. R., & Möller, M. (2003). Species separation of Taxus baccata, T. canadensis, and T. cuspidata (Taxaceae) and origins of their reputed hybrids inferred from RAPD and cpDNA data. Amer. J. Bot. 90: 175-182.

MILL, R. R. (2003). Towards a biogeography of Podocarpaceae. Acta Horticulturae (ISHS) 615: 137-147.

MILL, R. R. (ed.) (2003). Proceedings of the International Conifer Conference 1999. Being Acta Horticulturae (ISHS) 615. 485 pp.


Glidewell, S.M., Möller, M., Duncan, G., MILL, R.R., Masson, D. & Williamson, B. (2002). NMR imaging as a tool for noninvasive taxonomy: comparison of female cones of two Podocarpaceae. New Phytologist 154: 197-207. 

Herbert, J., Hollingsworth, P. M., Gardner, M. F., MILL, R. R., Thomas, P. I. & Jaffré, T. (2002). Conservation genetics and phylogenetics of New Caledonian Retrophyllum (Podocarpaceae) species. New Zealand J. Bot. 40(2): 175-188.

MILL, R.R. (2002). Proposal to conserve Podocarpus rotundus de Laub. (Recent Podocarpaceae) over Podocarpus rotundus Bocharn. (fossil Podocarpaceae). Taxon 51: 185-186.

MILL, R.R. & Quinn, C. J. (2002). Prumnopitys andina reinstated as the correct name for "lleuque", the Chilean conifer recently renamed P. spicata (Podocarpaceae). Taxon 50: 1143-1154.

Sinclair, W. T., MILL, R. R., Möller, M., Gardner, M. F., Woltz, P., Jaffré, T., Preston, J., Hollingsworth, M. & Ponge, (2002). A. Evolutionary relationships of the New Caledonian heterotrophic conifer, Parasitaxus usta (Podocarpaceae), inferred from chloroplast trnL-F intron / spacer and nuclear rDNA ITS2 sequences. Plant Syst. Evol. 233: 79-104.


Masson, D., Glidewell, S.M., Möller, M., MILL, R.R. & Williamson, B. (2001). Non-destructive examination of herbarium material for taxonomic studies using NMR imaging. Edinb. J. Bot. 58: 1-14.

MILL, R.R. (2001). A new sectional combination in Nageia Gaertn. (Podocarpaceae). Edin. J. Bot. 58: 499-501.

MILL, R.R. Möller, M., Glidewell, S.M., Masson, D., Christie, F. & Williamson, B. (2001). Morphology, anatomy and ontogeny of female cones in Acmopyle pancheri (Brongn. & Gris) Pilg. (Podocarpaceae). Annals of Botany 88: 55-67.


Möller, M., MILL, R.R., Glidewell, S.M., Masson, D., Williamson, B. & Bateman, R.M. (2000). Comparative biology and taxonomic significance of the pollination mechanisms in Acmopyle pancheri and Phyllocladus hypophyllus. Annals of Botany 86: 149-158.


Farjon, A. & MILL, R.R. (1388-1389) Proposals to conserve the current spelling of two generic names of conifers. Taxon 48: 151-154.

Fu Liguo, Li Nan & MILL, R.R. (1999). Ginkgoaceae, Araucariaceae, Pinaceae (except Abies and Picea), Taxodiaceae, Podocarpaceae, Cephalotaxaceae and Taxaceae. In Wu Zheng-yi & Raven, P.H., Flora of China 4: 8-25, 33-44, 52, 54-61, 78-96. Science Press, Beijing & Missouri Botanical Garden Press, St. Louis.

Fu Li-kuo, MILL, R.R. & Turland, N.J. (1999). Validation of the name Cephalotaxus latifolia (Cephalotaxaceae), a species from southeast China. Novon 9(2): 185-186.

MILL, R.R. & THOMAS, P. (1999). Plant portraits: 370. Falcatifolium taxoides. Curtis's Botanical Magazine 16(3): 199-211.

MILL, R.R. (1999). A new species of Larix from south-east Tibet and other nomenclatural notes on Chinese Larix. Novon 9(1): 79-82.

MILL, R.R. (1999). A new combination in Nageia (Podocarpaceae). Novon 9(1): 77-78.

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Dr Robert Mill

The Royal Botanic Garden Edinburgh is a charity (registration number SC007983)