Conifer Conservation Publications


  • MILL RR, RUHSAM M, THOMAS P, MF GARDNERPM 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.


  • Buckley BM, Stahle DK, Luu, HT, Wang SY, Nguyen TQT, THOMAS P, Le CN, Ton TM, Bui TH & VT Nguyen (2016). Central Vietnam climate over the past five centuries from cypress tree rings. Climate Dynamics doi:10.1007/s00382-016-3297-y
  • Gao LM, Yan L, Phan LK, Yan LJ, THOMAS P, Phan L, M Moller and DZ Li (2016) DNA barcoding of East Asian Amentotaxus (Taxaceae): potential new species and implications for conservation Journal of Systematics and Evolution. doi: 10.1111/jse.12207
  • 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
  • RUHSAM M, Rai HS, Mathews S, Ross TG, Graham SW, Raubeson LA, Mei W, THOMAS P, GARDNER MF, Ennos RA, HOLLINGSWORTH PM. (2015) Does complete plastid genome sequencing improve species discrimination and phylogenetic resolution in Araucaria? Mol Ecol Resour. 2015 Jan 22. doi: 10.1111/1755-0998.12375. [Epub ahead of print]
  • Sakaguchi S, Worth JRP, Jordan GJ, THOMAS P & M Ito (2015) Development of nuclear microsatellite markers for the Tasmanian endemic conifer Diselma archeri Hook.f. (Cupressaceae). Conservation Genetic Resources 7: 369-372
  • Sakaguchi S, Worth JRP, Jordan GJ, THOMAS P & M Ito (2015) Development of nuclear and mitochondrial microsatellite markers for the relictual conifer genus Athrotaxis (Cupressaceae). Conservation Genetic Resources 7: 477-481


  • GARDNER, M.F.  (2014) Conifers under threat. Chartered Forester Winter Issue 2014/15: 16-18. [This article first appeared in the Winter 2014/15 issue of Chartered Forester, the magazine for members of the Institute of Chartered Foresters (ICF), and is reproduced by kind permission of the ICF.]
  • Gaudeul M, GARDNER MF, THOMAS P, Ennos RA, HOLLINGSWORTH PM. (2014) Evolutionary dynamics of emblematic Araucaria species (Araucariaceae) in New Caledonia: nuclear and chloroplast markers suggest recent diversification, introgression, and a tight link between genetics and geography within species. BMC Evol Biol. 5(14)171. doi: 10.1186/s12862-014-0171-6.
  • 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


  • Biffin E, Brodribb TJ, Hill RS, THOMAS P & Lowe AJ (2012) Leaf evolution in Southern Hemisphere conifers tracks the angiosperm ecological radiation Proc. R. Soc. B 279: 341-348
  • Gaudeul M, Rouhan G, GARDNER MF and HOLLINGSWORTH PM (2012) AFLP markers provide insights into the evolutionary relationships and diversification of New Caledonian Araucaria species (Araucariaceae). American Journal of Botany 99: 68–81
  • Kettle, CJ, RA Ennos, T Jaffre, S McCoy, T Le-Borgne, MF GARDNER and PM Hollingsworth. (2012) Importance of demography and dispersal for the resilience and restoration of a critically endangered tropical conifer Araucaria  nemorosa. Diversity and Distributions 18: 248-259
  • 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 RR 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
  • Poudel RC, Möller M, Gao L-M, Ahrends A, Baral SR, Liu J, THOMAS P and DZ Li (2012) Using Morphological, Molecular and Climatic Data to Delimitate Yews along the Hindu Kush-Himalaya and Adjacent Regions. PLoS ONE 7(10):e46873.doi:10.1371/journal.pone.0046873


  • Chou YW, THOMAS P, Ge XJ, LePage B & Wang CW (2011) Refugia and phylogeography of Taiwania in East Asia. Journal of Biogeography.  DOI: 10.1111/j.1365-2699.2011.02537.x
  • Christenhusz MJM, Reveal JL, Farjon A, GARDNER MF, MILL RM & Chase MW  (2011) A new classification and linear sequence of extant gymnosperms. Phytotaxa 19: 55–70
  • Iason GI, Moore B, Russell J, HOLLINGSWORTH PM, Ennos R (2011) Reappraisal of our zones of genetic conservation of Caledonian Scots pinewood. In: Woodland Biodiversity (ed. Iason GI), pp. 18-19. Macaulay Land Use Research Institute, Aberdeen.
  • Little DP, THOMAS P, Nguyen Tien Hiep, Phan Ke Loc (2011) Before it had a name: Diagnostic characteristics, geographic distribution, and the conservation of Cupressus tonkinensis (Cupressaceae). Brittonia 63(2): 171-196
  • Stark Schilling DM & MILL RR (2011) Cuticle micromorphology of Caribbean and Central American species of Podocarpus (Podocarpaceae). International Journal of Plant Sciences  172 (5): 601-631
  • THOMAS P (2011) A review of the distribution and conservation status of Taxus in the Himalayas, China and Southeast Asia. Spanish Journal of Rural Development 2 (Special edition 2): 35-42
  • THOMAS P & LePage B (2011 ) The end of an era?—The conservation status of redwoods and other members of the former Taxodiaceae in the 21st century. Jpn. J. Histor. Bot. 19 (1–2): 89-100


  • MILL RR (2010) [1938] Proposal to reject the name Torreya bogotensis (Gymnosp., incertae sedis, Podocarp.). Taxon 59(3): 975-976
  • MILL RR & Stark Schilling DM (2010) Typification and nomenclature of Podocarpus angustifolius Griseb. and Podocarpus aristulatus Parl. (Podocarpaceae). Taxon, 59(3): 935-956.
  • Stark Schilling, DM & MILL RR (2010) [1937] Proposal to conserve Podocarpus aristulatus Parl. (Podocarpaceae) with a conserved type. Taxon, 59(3): 973-975


  • Gosling PG, McCartan SA, Ives LM, Cunningham VJ, SQUIRREL J and THOMAS P (2009) Preliminary observations on fruit handling, seed germination and chloroplast inheritance of an Amentotaxus hybrid arising at the Royal Botanic Garden Edinburgh from A. argotaenia (F) x A. formosana (M). Sibbaldia 6: 101-114
  • MILL RR & Stark Schilling D (2009) Cuticular micromorphology of Saxegothaea Lindl. (Podocarpaceae). Bot. J. Linn. Soc., 159: 58-67
  • THOMAS P (2009) The Araucariaceae: discovery, introduction and cultivation in the United Kingdom. In Proceedings of International Araucariaceae Symposium: Auckland New Zealand 14-17 March 2002: International Dendrology Society. 4, Dendrology and amenity: 231-240.


  • Amin Shah, De-Zhu Li, MÖLLER M, Lian-Ming Gao, HOLLINGSWORTH M & GIBBY M (2008) Delimitation of Taxus fuana Nan Li & R.R. Mill (Taxaceae) based on morphological and molecular data. Taxon 57(1): 211-222.
  • KETTLE CK, Ennos RA, Jaffre T, GARDNER MF & HOLLINGSWORTH PM (2008) Cryptic genetic bottlenecks during restoration of an endangered tropical conifer. Biological Conservation 141:1953-1961. 


  • Adams RP, THOMAS P & Rushforth K (2007) The leaf essential oils of the new conifer genus, Xanthocyparis : Xanthocyparis vietnamensis and X. nootkatensis. The Journal of essential oil research 19 (1): 30-33.
  • Farjon A & THOMAS P (2007) Taiwania cryptomerioides - an overview biogeography and conservation. In: The International Symposium on Taiwania cryptomerioides, 8-10 December 2007, Nantou County, Taiwan. Taiwan: Experimental Forest, College of Bio-resources and Agriculture, National Taiwan University. 9-17 pp.
  • Gao, L-M, MÖLLER M, Zhang X-M, HOLLINGSWORTH ML, Liu J, MILL RR, 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
  • KETTLE CJ, HOLLINGSWORTH PM, Jaffré T, Moran B, and Ennos RA (2007) Identifying the early genetic consequences of habitat degradation in a highly threatened tropical conifer, Araucaria nemorosa de Laub. Molecular Ecology 16: 3581-3591.
  • MÖLLER M, Gao, L-M, MILL RR, Li D-Z, HOLLINGSWORTH ML & GIBBY M (2007) Morphometric analysis of the Taxus wallichiana-complex based on herbarium material. Bot. J. Linn. Soc. 155: 307-335.
  • NEWMAN M, Sounthone Ketphanh, Bouakhaykhone Svengsuksa, THOMAS P, Khamphone Sengdala, Vichith Lamxay & ARMSTRONG K (2007) A checklist of the vascular plants of Lao PDR. Royal Botanic Garden Edinburgh, Forest Reseach Center, University of Lao PDR, IUCN Lao PDR, Vientiane. 394 pp.
  • NEWMAN M, THOMAS P, Soulivanh Lanorsavanh, Sounthone Ketphanh, Bouakhaykhone Svengsuksa and Vichith Lamxay (2007) New records of angiosperms and pteridophytes in the Flora of Laos. Edin. J. Bot. 64: 225-251.
  • Soto, D.P., C. Le Quesne, A. Lara & M.F. GARDNER (2007) Precarious conservation status of Pilgerodendron uviferum forests in their northern distribution in the Chilean Coastal Range. Bosque 28(3): 263-270.
  • THOMAS P, Khamphone Sengdala, Vichit Lamxay & Eanghourt Khou (2007) New records of conifers in Cambodia and Laos. Edin. J. Bot. 64(1): 37-44.


  • Farjon A, Bachman S, GARDNER MF, Luscombe D, Reynolds C & THOMAS P (2006) Conservation Assessments of Data Deficient (DD) Conifers Using Herbarium and Geographical Information System (GIS) data. Edinburgh: RBG Kew, Bedgebury Pinetum, RBGE. 311 pp.
  • GARDNER MF, HECHENLEITNER VP, THOMAS P, Echeverría C, Escobar B, BROWNLESS P & Martínez AC. (2006) Threatened Plants of Central and South Chile: Distribution, Conservation and Propagation. Universidad Austral de Chile & Royal Botanic Garden Edinburgh. 188 pp. (English language)
  • MILL RR (2006) A new name for Podocarpus rotundus de Laub. (Podocarpaceae). Edin. J. Bot. 63: 81-83.
  • MILL RR & Farjon A (2006) (1710). Proposal to conserve the name Xanthocyparis against Callitropsis Oerst. (Cupressaceae). Taxon 55: 229-231.
  • RAE D, CUBEY R, GARDNER MF, LATTA J & Walter K. (2006) Royal Botanic Garden Edinburgh Catalogue of plants 2006. Edinburgh: Royal Botanic Garden Edinburgh. 598 pp. 


  • GARDNER MF (2005) The International Conifer Conservation Programme. Plant Heritage 12: 23-26.  
  • HECHENLEITNER VP, GARDNER MF, THOMAS PI, Echeverría C, Escobar B, BROWNLESS P & Martínez CA. (2005) Plantas Amenazadas del Centro-Sur de Chile. Universidad Austral de Chile/ Real Jardín Botánico de Edimburgo. 187 pp. (Spanish language)


  • Brummitt, RK, MILL RR & 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.
  • Farjon A, THOMAS P & Nguyen Duc To Luu (2004) Conifer conservation in Vietnam: case reports of three potential ‘flagship species'. Oryx 38 (3): 257-266.
  • MILL RR (2004). (1612) Proposal to conserve Podocarpus affinis Seem. (Recent Podocarpaceae) over Podocarpus affinis A. Massal. (fossil Podocarpaceae). Taxon 53(1): 201-202.
  • MILL RR & Hill RS (2004) Validations of the names of seven Podocarpaceae macrofossils. Taxon 53: 1043-1046.
  • MILL RR, MÖLLER M, Glidewell SM, 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 RR & Weston P. (2004) (1613-1614) Proposals to reject the names Polypodiopsis and Polypodiopsis muelleri (Plantae vasculares, incertae sedis). Taxon 53(1): 203-205.
  • Muller EM, Barker NP & MILL RR. (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.
  • Nguyen Tien Hiep, Phan Ke Loc, Nguyen Duc To Luu, THOMAS P, Farjon A, Averyanov L  & J. Regalado Jr. (2004) Vietnam Conifers: Conservation Status Review 2004. Fauna & Flora International, Vietnam Programme, Hanoi. 128 pp.
  • Nguyen Duc To Luu & THOMAS P (2004) Cay La Kim Viet Nam (Conifers of Vietnam: an illustrated field guide). World Publishing House, Hanoi. 121 pp. (in Vietnamese and English)
  • Robertson A, HOLLINGSWORTH PM, KETTLE CJ, Ennos RA & GARDNER MF (2004) Characterisation of nuclear microsatellites in New Caledonian Araucaria species. Molecular Ecology Notes 4: 62-63. 


  • Collins D, MILL RR & 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.
  • GARDNER MF (2003) The International Conifer Conservation Programme. Acta Horticulturae (ISHS) 615: 405-409.
  • GARDNER MF  & Lara A. (2003) The Conifers of Chile: an overview of their Distribution and Ecology. Acta Horticulturae (ISHS) 615: 165 - 170.
  • Lara A, GARDNER MF, Vergara A & Escobar B. (2003) The Use and Conservation of Fitzroya cupressoides (Alerce) Forests in Chile. Acta Horticulturae (ISHS) 615: 381-385.

  • MILL RR (2003) Towards a biogeography of Podocarpaceae. Acta Horticulturae (ISHS) 615: 137-147.
  • MILL RR (ed.) (2003) Proceedings of the International Conifer Conference 1999. Being Acta Horticulturae (ISHS) 615. 485 pp.


  • Bekessy SA, D. Sleep, A. Stott, M. Menucinni, P. THOMAS, R.A. Ennos, M.A. Burgman, M.F. GARDNER, A.C. Newton (2002) Adaptation of Monkey Puzzle to arid environments reflected by regional differences in stable carbon isotype ratio and allocation to root biomass. Forest Genetics 9(1): 63-70. 
  • Glidewell SM, MÖLLER M, Duncan G, MILL RR, 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 PM, GARDNER MF, MILL RR, THOMAS PI & Jaffré T. (2002) Conservation genetics and phylogenetics of New Caledonian Retrophyllum (Podocarpaceae) species. New Zealand J. Bot. 40(2): 175-188.
  • MILL RR (2002) Proposal to conserve Podocarpus rotundus de Laub. (Recent Podocarpaceae) over Podocarpus rotundus Bocharn. (fossil Podocarpaceae). Taxon 51: 185-186.
  • MILL RR & Quinn CJ. (2002) Prumnopitys andina reinstated as the correct name for "lleuque", the Chilean conifer recently renamed P. spicata (Podocarpaceae). Taxon 50: 1143-1154.
  • Sinclair WT, MILL RR, MÖLLER M, GARDNER MF, Woltz P, Jaffré T, PRESTON J, HOLLINGSWORTH M & PONGE A. (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.


  • Allnutt T, Courtis J, GARDNER MF & Newton AC. (2001) Genetic variation in wild Chilean and cultivated British populations of Podocarpus salignus D.Don (Podocarpaceae). Edinb. J. Bot. 58: 459-473. 
  • Etisham-ul-Haq M, Allnutt TR, Smith-Ramírez C, GARDNER MF, Armesto JJ & Newton AJ. (2001) Patterns of Genetic Variation in in and ex situ populations of the threatened Chilean vine Berberidopsis corallina, detected using RAPD markers. Annals of Botany 87: 813-821.  
  • Masson D, Glidewell SM, MÖLLER M, MILL RR & Williamson B. (2001) Non-destructive examination of herbarium material for taxonomic studies using NMR imaging. Edinb. J. Bot. 58: 1-14.
  • MILL RR (2001) A new sectional combination in Nageia Gaertn. (Podocarpaceae). Edin. J. Bot. 58: 499-501.
  • MILL RR, MÖLLER M, Glidewell SM, 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 RR, Glidewell SM, Masson D, Williamson B & Bateman RM (2000) Comparative biology and taxonomic significance of the pollination mechanisms in Acmopyle pancheri and Phyllocladus hypophyllus. Annals of Botany 86: 149-158.
  • THOMAS P & WATSON MF. (2000) Data management for plant collections. A handbook of best practice. Edinburgh, Scotland: Royal Botanic Garden Edinburgh. 48 pp. 


  • Allnutt T, Newton AC, Lara A, Premoli A, Armesto J & GARDNER MF. (1999) Genetic variation in Fitzroya cupressoides (alerce), a threatened South American conifer. Jour. Molecular Ecology 8: 975-987.
  • Farjon A & MILL RR (1388-1389) Proposals to conserve the current spelling of two generic names of conifers. Taxon 48: 151-154.
  • Fraver S, González ME, Silla F, Lara A & GARDNER MF (1999) Composition and structure of remnant Fitzroya cupressoides forests of Southern Chile's Central Depression. Bull.Torrey Bot. Club 126(1): 49-57.
  • Fu Liguo, Li Nan & MILL RR (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 RR & Turland NJ (1999) Validation of the name Cephalotaxus latifolia (Cephalotaxaceae), a species from southeast China. Novon 9(2): 185-186.
  • GARDNER MF, THOMAS P, Lara A & Escobar B. (1999) Fitzroya cupressoides. Curtis's Botanical Magazine 16(3): 229-240.
  • GARDNER MF  (1999) Managing ex situ conifer conservation collections. In: S. Andrews, A.C. Leslie and C. Alexander (Editors). Taxonomy of Cultivated Plants: Third International Symposium, pp. 19-23. Royal Botanic Gardens, Kew  
  • MILL RR & THOMAS P. (1999) Plant portraits: 370. Falcatifolium taxoides. Curtis's Botanical Magazine 16(3): 199-211.
  • MILL RR (1999). A new species of Larix from south-east Tibet and other nomenclatural notes on Chinese Larix. Novon 9(1): 79-82.
  • MILL RR (1999) A new combination in Nageia (Podocarpaceae). Novon 9(1): 77-78.


  • Allnutt T, THOMAS P, Newton AC & GARDNER MF (1998) Genetic variation in Fitzroya cupressoides cultivated in the British Isles, assessed using RAPDS. Edinb. J. Bot. 55(3): 329-341. 

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