Email: Toby Pennington
Phone: +44 (0)131 248 2818
Fax: +44 (0)131 248 2901
Systematics
My own taxonomic research, and that of PhD students has focused largely upon genera of the ecologically and economically important legume family (Leguminosae), including Andira, Dussia, Geoffroea, Cyclolobium, Lathyrus, Berlinia, Ateleia, Cyathostegia. Current legume PhD students are studying Senna in Arabia, neotropical Vicia and Amicia. Other PhD students have worked on Scaphium (Sterculiaceae), Cyrtandra (Gesneriaceae) and Garcinia (Guttiferae). These projects involved the production of fundamental monographic taxonomic accounts and the use of molecular systematics to resolve species relationships and to investigate the nature of tropical plant species.
The legume research expanded to include a molecular systematic study of relationships amongst the putatively primitive genera of the legume subfamily Papilionoideae, which formed part of an international effort to elucidate the phylogeny of the Leguminosae. Results from this study radically challenged conventional models of the evolution of the characteristic "pea" flower of the papilionoid (pea-flowered) legumes. This led to the development of a PhD project that demonstrated that legume homologues of the Antirrhinum floral symmetry gene cycloidea are involved in the control of floral symmetry in legumes.
Applied research: inventory and capacity building
More applied research has been centred in Peru. Work there has focused on two UK Government Darwin Initiative Projects "Tree diversity and agroforestry development in the Peruvian Amazon" and "Tree diversity, agroforestry development and reafforestation in the Peruvian Andes" (£244K; one UK staff, four Peruvian staff, one Peruvian MSc student). The first Darwin Initiative project provided a non-technical identification guide in Spanish to 140 tree species from the Peruvian Amazon with economic potential, and built the capacity of the Peruvian National Forest Herbarium. The second Darwin project continued this capacity building work, and will provide a companion identification guide to 130 tree species with economic uses from the Peruvian Andes. The focus of these Darwin projects on agroforestry systems was diversified in collaboration with the International Centre for Research in Agroforestry (ICRAF), a partner in the Darwin work, by examining genetic variation in the legume Inga edulis. This species is widely used in agroforestry systems across the Neotropics, and the project (Scottish Executive Environment and Rural Affairs Department (SEERAD) Flexible Fund, £125K; one postdoc), demonstrated that genetic variation in cultivated populations is almost as high as that in wild populations.
Time spent in Peru via has afforded opportunities for inventory work in dry forests found in inter-Andean valleys, building on RBGE's long history of research in neotropical dry forests and savannas. Collections made include several new species and new records for Peru. Given the high threat to these areas their conservation is paramount. In collaboration with Reynaldo Linares, these conservation issues have been addressed in two ways. First, by demonstrating that Peruvian dry forests have high endemism and diversity compared to many other neotropical dry forests and merit urgent conservation. Second, by developing an online floristic checklist of woody dry forest plants in Peru as a tool for both identification and conservation prioritisation.
Biogeography and environmental change
My final research area, which draws together aspects of all the above research projects, is the investigation of the historical construction of tropical floras, their biogeography, and how they have reacted to past environmental change. Molecular phylogenies calibrated with a temporal dimension offer a new means of dating the origin of plant species that form the structural framework of ecosystems. They can therefore address fundamental and unanswered questions relating to the explanation of high tropical species diversity such as whether species have accumulated slowly over geological time, or more recently in response to climatic changes.
Phylogenetic work carried out as part of the Inga project showed that all 300 species of this large genus had evolved remarkably recently, raising the possibility that much plant species diversity in the world's richest rain forests may have resulted from recent speciation. In contrast, via a Leverhulme Trust funded project "Biogeography of South American seasonally dry forests" (£100K; one postdoc), we have demonstrated that species in five unrelated genera characteristic of dry forests are more ancient. These studies have been followed by a series of related publications with emphasis on taxonomic groups that are research foci at RBGE such as Begonia, Cyrtandra (Gesneriaceae), and Renealmia (Zingiberaceae). These have contributed to a rapidly changing global view of the assembly of species-rich continental biomes that acknowledges a far greater role for long-distance transoceanic dispersal in the generation of pantropical distributions and the plausibility of recent (Quaternary) tropical plant speciation. Results of similar studies in different species-rich continental ecosystems were synthesised at a Royal Society of London Discussion Meeting, "Plant Phylogeny and the Origin of Major Biomes", which I co-organised in 2004. Our dry forest research led to the organisation of an international conference "Tropical Savannas and Seasonally Dry Forests: Ecology, Diversity and Development", held at RBGE in September 2003. The symposium within this meeting has resulted in a book: "Neotropical seasonally dry forests and savannas: plant diversity, biogeography and conservation" (CRC Press, 2006).
Other duties
I am Head of the Tropical Diversity section at RBGE. I attend the University of Edinburgh/RBGE MSc in Plant Taxonomy and Biodiversity steering committee. I teach Phylogenetics, Plant Geography and Leguminosae to MSc and undergraduate students at the University of Edinburgh
RESEARCH GROUP
Researchers (core funded RBGE staff not listed)
UK:
Reynaldo Linares (Darwin Initiative Scholar 2004-2005)
Peruvian seasonally dry tropical forests
Colin Pendry (post-doctoral researcher 1998-2001)
Historical biogeography of South American seasonally dry tropical forests
James Richardson (post-doctoral researcher 1999-2000)
Molecular systematics of the economically important neotropical genus Inga
Tania Durt (Darwin research fellow 2001-2002)
Tree diversity and agroforestry development in the Peruvian Amazon
Peru:
Dr Carlos Reynel (Professor, Universidad Nacional Agraria, La Molina, Lima, Peru; 2000-2006; 30%): Peruvian floristics
Dr Terry Pennington (Consultant; Darwin Initiative; 2000-2006; 30%): Peruvian floristics
Mr José-Luis Marcelo (Assistant professor, Universidad Nacional Agraria, La Molina, Lima, Peru; 2005-2006, 30%): Peruvian floristics
Mr Aniceto Daza (Technician, Universidad Nacional Agraria, La Molina, Lima, Peru; 2000-2006; 30%): Peruvian floristics
Ms Rocio Ravello (Technician, Universidad Nacional Agraria, La Molina, Lima, Peru; 2000-2006, 30%): Peruvian floristic
Sabbatical and other research visitors:
Dr Haroldo Cavalcante de Lima, Jardim Botanico do Rio de Janeiro, Brazil (one year sabbatical, 2009-2010)
Dr Susana Magallón, Universidad Nacional Autonoma de México (one year sabbatical, 2009-2010)
João Iganci, Universidade Federal do Rio Grande do Sul, Brazil (six month PhD ‘sandwich' placement funded by Brazilian Government)
PhD students:
Paulina Hechenleitner, University of Aberdeen (2009-2012)
Systematics and biogeography of neotropical Vicia (Leguminosae)
Supervisors: David Burslem (Aberdeen), Gwilym Lewis (Kew), Greg Kenicer, Toby Pennington
Euridice Honorio, University of Leeds (2009-2012)
Integrating ecological and molecular information of widespread tree species in Western Amazonia
Supervisors: Oliver Phillips (Leeds), Toby Pennington
Faten Filimban, University of Edinburgh (UoE) (2009-2012)
Systematics and biogeography of Senna (Leguminosae) in Arabia
Supervisors: Barbara MacKinder, Tony Miller, Michael Moeller, Toby Pennington
Katharine Fitzpatrick, UoE (2007-2010)
Multi-proxy based Late Quaternary environmental reconstruction in lowland tropical South America
Supervisors: Francis Mayle (UoE), FA Street-Perrott (Swansea University), Toby Pennington
Macarena Cardenas, Open University (2007-2010)
Climate change impacts on hyper-diverse Amazonian ecosystems
Supervisors: William D. Gosling, Sarah Sherlock, Vincent Gauci (OU), Toby Pennington, Imogen Poole (Utrecht)
Tiina Sarkinen, Department of Plant Sciences, University of Oxford (2006-2009)
Systematics of Amicia (Leguminosae) and plant diversification in the Andes
Supervisors: Colin Hughes, John Pannell (Oxford), Toby Pennington
Bhaskar Adhikari, University of Edinburgh (2006-2009)
Investigating the evolution of large genera: how did Berberis speciate in Nepal?
Supervisors: Richard Milne (UoE), Colin Pendry, Toby Pennington
Bronwen Witney, Dept. Geography, University of Edinburgh (UoE) (2005-2008; NERC)
Palaeoecology of the Pantanal: Late Quaternary vegetation and climate change in Eastern Boliviva
Supervisors: Francis Mayle (UoE), Toby Pennington
Huw Jones Dept. Geography, University of Edinburgh (2005-2008; NERC)
Characterisation of neotropical savanna ecosystems by their modern pollen rain.
Supervisors: Francis Mayle (UoE), Toby Pennington, Jim Ratter
Sofia Caetano, University of Geneva (2003-2007; Swiss National Science Foundation)
South American seasonally dry tropical forests: are they current refugia or the result of long distance dispersal?
Supervisors: Yamama Naciri (Geneva), Rodolphe Spichiger (Geneva), Toby Pennington
Greg Kenicer, Institute of Cell and Molecular Biology (ICMB), University of Edinburgh (2002-2006; self-funding)
Systematics and biogeography of Lathyrus section Orobus (Leguminosae)
Supervisors: Toby Pennington, Mark Watson
Nazre Saleh, ICMB, University of Edinburgh (2002-2006; Government of Malaysia)
Systematics of Garcinia section Garcinia (Guttiferae)
Supervisors: Toby Pennington, Mark Newman
Gemma Bramley, ICMB, University of Edinburgh (2001-2003; NERC)
Local, regional and monographic approaches to Cyrtandra (Gesneriaceae)
Supervisors: Quentin Cronk, Mary Mendum, Toby Pennington
Barbara MacKinder, ICMB, University of Edinburgh (2001-2006; RBG Kew)
Systematics and biogeography of Berlinia (Leguminosae - Caesalpinioideae)
Supervisors: Toby Pennington, David Harris
Helene Citerne, ICMB, University of Edinburgh (2001-2004; Carnegie Foundation)
Examination of cyc-like genes in the Leguminosae
Supervisors: Toby Pennington, Quentin Cronk, Enrico Coen (John Innes Institute)
Peter Wilkie, University of Aberdeen (2000-2006; RBGE)
Systematics of Asian Sterculiaceae
Supervisors: Toby Pennington, Chris Wilcock (University of Aberdeen)
Elizabeth Moylan: Department of Plant Sciences, University of Oxford (1998-2001)
Systematics of Hemigraphis (Acanthaceae)
Supervisors: Toby Pennington, Robert Scotland (University of Oxford), Paula Rudall (RBG Kew)
Helen Ireland: Department of Plant Sciences, University of Reading (1997-2000)
Systematics of Ateleia and Cyathostegia (Leguminosae-Swartzieae)
Supervisors: Toby Pennington, Brian Schrire (RBG Kew), Frank Bisby (University of Reading)
MSc students:
Cathy King, RBGE MSc student (2009) Biogeography of Dussia (Leguminosae)
Supervised with: Tiina Sarkinen
Sarah Cody, RBGE MSc student (2008) The use of plant and animal dated phylogenies to explore the effects of the closure of the Panama Isthmus
Supervised with: James Richardson
Euridice Honorio, RBGE MSc student (2006) Floristic similarity, soils and distance in the Amazon basin. Supervised with: Sam Bridgewater (Natural History Museum)
Tiina Sarkinen, RBGE MSc student (2005) Transatlantic biogeography of Renealmia (Zingiberaceae). Supervised with: Mark Newman
Rhonda Ridley, RBGE MSc student (2004) Biogeography and systematics of Lathyrus (Leguminosae). Supervised with: Greg Kenicer
James Tosh, RBGE MSc student (2004) Biogeography and systematics of Irvingiaceae. Supervised with: David Harris
Anton Russell, RBGE MSc student (2004) The 50kb chloroplast DNA inversion as a taxonomic character in Leguminosae. Supervised with Michelle Hollingsworth
Cindy Skema RBGE MSc student (2003) Phylogeny and biogeography of Andira Lam: a case study in Neotropical speciation. Supervised with: Michelle Hollingsworth
Josh Clayton RBGE MSc student (2003) Pantropical genera: systematics and biogepgrpahy. A pilot study for the pantropical tree Manilkara. Supervised with: Vanessa Plana, Michelle Hoillingsworth, Alex Ponge
Reynaldo Linares Palomino, RBGE MSc student (2002) A floristic and phytogeographic analysis of Peruvian seasonally dry forests. Supervised with: Sam Bridgewater
Simon Queenborough, RBGE MSc student (2001) The floristics and phytogeography of Central American tropical forests: a study based upon the deciduous forests of Belize. Supervised with: Sam Bridgewater and Peter Wilkie
Michael Pirie, RBGE MSc student (2000) Systematics and Biogeography of Centrolobium (Leguminosae). Supervised with: Bente Klitgaard (RBG Kew) and Colin Pendry
Helen Cortes Burns, RBGE MSc student (2000) A taxonomic and biogeographic study of Socotran Indigofereae. Supervised with: Brian Schrire (RBG Kew) and Tony Miller
Shirin Rezai, RBGE MSc student (2000) Biogeography and systematics of Basistemon (Scrophulariaceae). Supervised with Colin Pendry
Richard Howarth MSc (by research), Department of Geography, University of Edinburgh (1999) Conservation and land-use in neotropical montane cloud forests: an analysis of vegetation succession in the Venezuelan Cordillera de la Costa Central. Supervised with: Colin Pendry, Peter Furley (University of Edinburgh)
Barbara MacKinder, RBGE MSc student (1999) Systematics of Berlinia (Leguminosae-Caesalpinioideae). Supervised with: Gwilym Lewis (RBG Kew)
Freya Cooper, RBGE MSc student (1998) A field guide to the woody papilionoid legumes of the threatened Brazilian cerrado. Supervised with: Sam Bridgewater
Josephine Welsh, RBGE MSc student (1997) Morphological and molecular cladistic analysis of the Bencomia alliance (Rosaceae-Poteriae): adaptive radiation on the Macronesian islands. Supervised with: Richard Bateman and Quentin Cronk
Andrew Bell, RBGE MSc student (1996) Molecules, colleters and persistent petiole bases: a search for new characters in the genus Sorbus (Rosaceae, Maloideae). Supervised with: Crinan Alexander, Stephen Spongberg (Arnold Arboretum)
Helen Ireland, RBGE MSc student (1996) Taxonomy, biogeography and molecular systematics of Geoffroea (Leguminosae). Supervised with: Brian Schrire (RBG Kew)
Carlota Quilambo, University of Reading MSc student (1996) Anatomical survey of Dussia (Leguminosae) leaves. Supervised with: David Cutler and Charles Stirton (both RBG Kew)
Birgit Gemeinholzer, RBGE MSc student (1995). A survey of fruit anatomy in the genus Andira (Leguminosae)
Other students
Ben Mosse, University of Edinburgh undergraduate student (2000) Molecular systematics and biogeography of Loxopterygium (Anacardiaceae)
Lizabeth Caddick, Gatsby Charitable Foundation summer student (1995) Molecular identification of Andira hybrids
PUBLICATIONS
(a) Refereed research papers
69. Linares, R., Oliveira Filho A.T. & PENNINGTON, R.T. (2011, in press). Neotropical Seasonally Dry Forests: Diversity, Endemism and Biogeography of Woody Plants. In R. Dirzo, H. Mooney, G. Ceballos, H. Young (eds). Latin American Seasonally Dry Tropical Forests. Island Press
68. Meir, P. & Pennington, R.T. (2011, in press). Climate change and seasonally dry tropical forests. In R. Dirzo, H. Mooney, G. Ceballos, H. Young (eds). Latin American Seasonally Dry Tropical Forests. Island Press
67. Cardenas, M.L., Gosling, W.D., Sherlock, S.C., Poole, I., PENNINGTON, R.T. & Mothes P. (accepted). The response of vegetation at the Andean flank in western Amazonia to Pleistocene climate change. Science 331: 1055-1058
66. Barlow, J., Ewers, R.M., Anderson L., Aragao, L.E.O.C., Baker, T., Boyd E., Feldpausch T., Gloor E., Hall A., Malhi Y., Milliken W., Mulligan M., Parry L., PENNINGTON R.T., Peres C.A., Phillips O., Roman-Cuesta R.M., Tobias J. & Gardner T.A.. (2011, in press) Using learning networks to understand complex systems: a case study of biological, geophysical and social research in the Amazon. Biological Reviews
65. PENNINGTON, R.T., Daza A & Lavin M. (2011, in press). Poissonia eriantha (Leguminosae) From Cuzco, Peru: An Overlooked Species Underscores a Pattern of Narrow Endemism Common to Seasonally Dry Neotropical Vegetation Poissonia. Systematic Botany
64. Milliken, W., Zappi D., Sasaki D., Hopkins M. & PENNINGTON R.T. (2011). Amazon vegetation: how much don’t we know and how much does it matter? Kew Bulletin 65: 1-19.
63. MacKinder, B. & PENNINGTON R.T. (2011). A monograph of Berlinia (Leguminosae: Caesalpinioideae). Systematic Botany Monographs 91 (117 pp.)
62. PENNINGTON, R.T. & Dick, C.W. (2010). Diversification of the Amazonian flora and its relation to key geological and environmental events: a molecular perspective. In C. Hoorn, H. Vonhof, F. Wesselingh (eds). Amazonia, Landscape and Species Evolution: a Look into the Past. pp. 373-385. Blackwell.
61. PENNINGTON R.T., Lavin M, Hughes C., Sarkinen T, Lewis G, Klitgaard B. (2010). Differing diversification histories in the Andean biodiversity hotspot. Proceedings of the National Academy of Sciences, USA 107: 13783-13787.
60. Cody, S., Richardson, J.E., Rull, V., Ellis, C. & PENNINGTON, R.T. (2010). The Great American Biotic Interchange Revisited. Ecography 33: 1-7.56. Honorio E.N., Baker, T.R., Phillips, O.L., Pitman, N.C.A., PENNINGTON R.T. et al. (2009). Integrating regional and continental scale comparisons of tree composition in Amazonian terra firme forests. Biogeosciences Discussions 6: 1-31.
59. Ireland, H.E., Kite G.C., Veitch, N.C., Chase, M.W., Schrire, B., Lavin, M., PENNINGTON, R.T. (2010). Biogeographic, ecological, and morphological structure of a phylogenetic analysis of Ateleia (Swartzieae-Leguminosae) derived from combined molecular and morphological/chemical data. Botanical Journal of the Linnean Society 162: 39-53
58. Smith, G.F., Figueiredo, E., PENNINGTON, R.T., Davila, P. (2009). Getting that grant: how to convince an evaluation panel that your proposal is worthy of funding. Taxon, 58: 675-677. A. 2.36
57. Simon, M., Grether, R., de Queiroz L.P., Skema C., PENNINGTON, R.T. & C.E. Hughes . (2009) Recent assembly of the Cerrado, a neotropical plant diversity hotspot, by in-situ evolution of adaptations to fire. Proceedings of the National Academy of Sciences, USA 106: 20359-20364.
55. PENNINGTON, R.T., Lavin, M & Oliveira-Filho A. (2009). Plant diversity, evolution and ecology in the tropics: perspectives from seasonally dry tropical forests. Annual Review of Ecology, Evolution and Systematics 40: 437-457.
54. Kursar, T.A., Dexter, K.G., Lokvam, J., PENNINGTON R.T., Richardson, J.E., Weber, M.G., Murakami, E, Drake, C., McGregor, R. & Coley, P.D. (2009). The importance of plant-herbivore interactions for diversification and coexistence in the tropical tree genus Inga. Proceedings of the National Academy of Sciences, USA. doi: 10.1073/pnas.0904786106
53. Hollingsworth, M.L., Clark, A., Forrest, L.L., Richardson, J., PENNINGTON, R.T., Long, D.G., Cowan, R., Chase, M.W., Gaudeul, M., Hollingsworth, P.M. (2009). Selecting barcoding loci for plants: evaluation of seven candidate loci with species-level sampling in three divergent groups of land plants. Molecular Ecology Resources, 9: 439-457.
52. Pirie, M.D., B.B. Klitgaard & R.T. Pennington. (2009). Revision and biogeography of Centrolobium. Systematic Botany 34: 345-359
51. Honorio, E.N. R.T. PENNINGTON, L.A. Freitas, G. Nebel & T.R. Baker. (2008). Análisis de la composición florística de los bosques de Jenaro Herrera, Loreto, Perú. Revista Peruana de Biologia.
50. PENNINGTON, R.T. & Wojciechowski, M.J. (2008). The status of Sophora. The Plantsman 7: 186-189
49. Caetano, S., D. Prado, R.T. PENNINGTON, S. Beck, A. Oliveira-Filho, R. Spichiger and Y. Naciri. 2008. The history of Seasonally Dry Tropical Forests in eastern South America: inferences from the genetic structure of the tree Astronium urundeuva (Anacardiaceae). Molecular Ecology 17:3147-3159.
48. Dawson, I.K., Hollingsworth, P., Doyle, J.J., Kresovich, S., Weber, J.C., Sotelo Montes, C., Pennington, T.D., PENNINGTON, R.T. (2008). Origins and genetic conservation of tropical trees in agroforestry systems: a case study from the Peruvian Amazon. Conservation Genetics 9: 361-372.
47. PENNINGTON, R.T. in Flora Neotropica Monograph 100. Proteaceae (Prance, G.T., Edwards, K., Plana, V. & PENNINGTON, R.T.): accounts of Gevuina, Embothrium, Oreocallis and Lomatia, pp.22-36.
46. Särkinen, T.E., Newman, M.F., Maas, P.J.M., Maas, H., Poulsen, A.D., Harris, D.J., Richardson, J.E., Clark, A., Hollingsworth, M. & PENNINGTON, R.T. (2007). Recent oceanic long-distance dispersal and divergence in the amphi-Atlantic rain forest genus Renealmia L.f. (Zingiberaceae). Molecular Phylogenetics and Evolution 44: 968-980.
45. PENNINGTON, R.T., Richardson, J.A. & Lavin M. (2006, in press). Insights into the historical construction of species-rich biomes from dated plant phylogenies, phylogenetic community structure and neutral ecological theory. New Phytologist (invited Tansley Review) 172: 605-616.
44. Citerne, H., PENNINGTON, R.T. & Cronk, Q.C.B. (2006). An apparent reversal in floral symmetry in the legume Cadia is a homeotic transformation. Proceedings of the National Academy of Sciences, USA 103: 12017-12020
43. PENNINGTON, R.T., Ratter, J.A. & Lewis, G.P. (2006). An overview of the plant diversity, biogeography and conservation of neotropical savannas and seasonally dry forests. In R.T. Pennington, G.P. Lewis & J.A. Ratter (eds). Neotropical savannas and seasonally dry forests: plant diversity, biogeography and conservation. pp. 1-29. CRC Press, Florida.
42. Naciri, Y., Caetano, S., PENNINGTON, R.T., Prado, D. & Spichiger, R. (2006). Population genetics and inference of ecosystem history: an example using two neotropical seasonally dry forest species. In R.T. Pennington, G.P. Lewis & J.A. Ratter (eds). Neotropical savannas and seasonally dry forests: plantdiversity, biogeography and conservation. pp. 417-432. CRC Press, Florida.
41. Wilkie, P., Ponge, A., PENNINGTON, R.T., Cheek, M., Bayer, C. & Wilcock, C. (2006). A phylogenetic analysis of subfamily Sterculioideae (Malvaceae /Sterculiaceae - Sterculieae) using the chloroplast gene ndhF. Systematic Botany 31: 160-170.
40. Cronk, Q.C.B., Ojeda, I. & PENNINGTON, R.T. (2005). Legume comparative genomics: progress in phylogenetics and phylogenomics. Current Opinion in Plant Biology 9: 1-5.
39. Coley, P. D., Lokvam, J., Rudolph, K., Bromberg, K., Wright, L., Dvorett, D., Ring, S., Ponge, A., Baptiste, C., Pennington, R. T. & Kursar, T. A. (2005). Divergent defensive strategies of young leaves in two Neotropical species of Inga. Ecology 86: 2633-2643
38. Naciri-Graven, Y., Caetano, S., Prado, D., PENNINGTON, R.T. & Spichiger, R. (2005). Development and characterization of 11 microsatellite markers in a widespread neotropical seasonally dry forest tree species, Geoffroea spinosa Jacq. (Leguminosae). Molecular Ecology Notes 5: 542-545.
37. PENNINGTON, R.T., Stirton, C.S. & Schrire, B.D. (2005). Sophoreae. In Lewis, G.P., Schrire, B.D, MacKinder, B. & Lock, J.M. (eds). Legumes of the World pp. 227-249. Royal Botanic Gardens, Kew.
36. Kenicer, G., Kajita, T., PENNINGTON, R.T. & Murata, J. (2005). Systematics and biogeography of Lathyrus based upon internal transcribed spacer and cpDNA sequence data. American Journal of Botany 92: 1199-1209
35. PENNINGTON, R.T., Lavin, M., Prado, D.E., Pendry C.A. & Pell, S. (2005). Climate change and speciation. In Tropical forests and global atmospheric change (eds. Y. Malhi and O.L. Phillips), pp. 199-214. Oxford University Press
34. Hollingsworth, P. M., Dawson, I. K., Goodall-Copestake, W. P., Richardson, J. E., Weber, J. C., Sotelo Montes, C., Pennington, R.T. (2005). Do farmers reduce genetic diversity when they domesticate tropical trees? A case study from Amazonia. Molecular Ecology 14: 497-501.
33. PENNINGTON, R.T., Richardson, J.E. & Cronk, Q.C.B. (2004). Plant phylogeny and the origin of major biomes: introduction and synthesis. Philosophical Transactions of the Royal Society (Biological Sciences) 359: 1455-1465.
32. PENNINGTON, R.T. & Dick, C.W. (2004). The role of immigrants in the assembly of the Amazonian tree flora. Philosophical Transactions of the Royal Society (Biological Sciences) 359: 1611-1622.
31. Lavin, M., Schrire, B., Lewis, G., PENNINGTON, R.T., Delgado-Salinas, A., Thulin, M., Hughes, C. & Wojciechowski, M.F. (2004). Metacommunity process rather than continental tectonic history better explains geographically structured phylogenies in legumes. Philosophical Transactions of the Royal Society (Biological Sciences) 359: 1509-1522.
30. Cortés-Burns, H., Schrire, B. D., PENNINGTON, R. T. & Miller, A. G. (2004). A taxonomic revision of Socotran Indigofereae Benth. (Leguminosae - Papilionoideae) with insights into the phytogeographical links of the Socotran Archipelago. Nordic Journal of Botany 22: 693-711.
29. Plana, V., Gascoigne, A., Forrest, L.L., Harris, D., & PENNINGTON, R.T. (2004). Pleistocene and pre-Pleistocene Begonia speciation in Africa. Molecular Phylogenetics and Evolution 31: 449-461.
28. PENNINGTON, R.T., Lavin, M. Prado, D.E., Pendry, C.A., Pell, S. & Butterworth, C.A. (2004). Historical climate change and speciation: Neotropical seasonally dry forest plants show patterns of both Tertiary and Quaternary diversification. Philosophical Transactions of the Royal Society (Biological Sciences) 359: 515-538.
27. Bramley, G.L.C., PENNINGTON, R.T., Zakaria, R., Sudarmiyati Tjitrosoedirdjo, S., & Cronk, Q.C.B. (2004). Assembly of tropical plant diversity on a local scale: Cyrtandra (Gesneriaceae) on Mount Kerinci, Sumatra. Biological Journal of the Linnean Society 81: 49-62.
26. PENNINGTON, R. T., Pendry, C.A., Goodall-Copestake, W., O'Sullivan, S. (2004). Phylogenetic analysis of Ruprechtia. In Pendry, C.A. A monograph of Ruprecthia (Polygonaceae). Systematic Botany Monographs 67: 12-17.
25. Kite, G.C. and PENNINGTON, R.T. (2003). Quinolizidine alkaloid status of Styphnlobium and Cladrastis (Leguminosae). Biochemical Systematics and Ecology 31: 1409-1416.
24. Linares-Palomino, R., PENNINGTON, R.T. & Bridgewater, S. (2003). The phytogeography of seasonally dry tropical forests in Equatorial Pacific South America. Candollea 58: 473-499.
23. PENNINGTON, R.T. (2003). A monograph of Andira (Leguminosae-Papilionoideae). Systematic Botany Monographs 64 (145pp.; illus. 44 b+w).
22. Citerne, H., Luo, D., PENNINGTON, R.T., Coen, E. & Cronk, Q.C.B. (2003). A Phylogenomic investigation of CYC-like TCP genes in Leguminosae. Plant Physiology 131: 1042-1053.
21. Bridgewater, S., PENNINGTON, R.T., Reynel, C., Daza, A. & Pennington, T.D. (2003). A preliminary floristic and phytogeographic analysis of the woody flora of seasonally dry forests in northern Peru. Candollea 58: 129-148.
20. Lewis, G.P., Knudsen, J.T., Klitgaard, B., & PENNINGTON, R.T. (2003). The floral scent of Cyathostegia mathewsii (Benth.) Schery and preliminary observations on its reproductive biology. Biochemical Systematics and Ecology 31: 951-962.
19. Moylan, E.C., PENNINGTON, R.T. & Scotland, R.W. (2002). Taxonomic account of Hemigraphis Nees (Strobilanthinae-Acanthaceae) from the Philippines. Kew Bulletin 57: 769-825.
18. PENNINGTON, R.T. (2002). Proposal to change the authorship of Andira nom. cons. (Leguminosae-Papilionoideae) and to conserve it with a conserved type. Taxon 51: 385-386.
17. Warwick, M. & PENNINGTON, R.T. (2002). A revision of Cyclolobium (Leguminosae-Papilionoideae). Edinburgh Journal of Botany 59: 247-258.
16. Richardson, J.E., PENNINGTON, R.T., Pennington, T.D. & Hollingsworth, P.M. (2001). Rapid diversification of a species-rich genus of Neotropical rain forest trees. Science 293: 2242-2245
15. PENNINGTON, R.T., Lavin, M., Ireland, H.E., Klitgaard, B. & Preston, J. (2001). Phylogenetic relationships of primitive papilionoid legumes based upon sequences of the chloroplast intron trnL. Systematic Botany 26: 537-556.
14. Lavin, M., PENNINGTON, R. T., Klitgaard, B.B., Sprent, J.I., de Lima, H.C. & Gasson, P.E. (2001). The dalbergioid legumes (Fabaceae): Delimitation of a pantropical monophyletic clade. American Journal of Botany 88: 503-533.
13. PENNINGTON, R.T. & Gemeinholzer, B. (2000). Cryptic clades, fruit wall morphology and biology of Andira (Leguminosae-Papilionoideae). Botanical Journal of the Linnean Society 134: 267-286.
12. Ireland, H., PENNINGTON, R.T. & Preston, J. (2000). Molecular Systematics of the Swartzieae. In: P. Herendeen & A. Bruneau (editors). Advances in legume systematics. Part 9. pp. 217-232. Royal Botanic Gardens, Kew.
11. PENNINGTON R.T., Klitgaard, B., Ireland, H.E., and Lavin, M. (2000). New insights into floral evolution of basal Papilionoideae from molecular phylogenies. In: P. Herendeen & A. Bruneau (editors). Advances in legume systematics. Part 9. pp. 233-248. Royal Botanic Gardens, Kew.
10. PENNINGTON, R.T., Lewis, G.P., Marsh, M. (2000). Andira inermis subsp. inermis (Leguminosae-Papilionoideae). Curtis Botanical Magazine 17: 188-194.
9. Lavin, M., Thulin, M., Labat, J-N. & PENNINGTON, R.T. (2000). Africa, the odd man out: molecular biogeography of dalbergioid legumes (Fabaceae) suggests otherwise. Systematic Botany 25: 449-467.
8. PENNINGTON, R.T. (2000). Introduction In Scotland, R.W. & Pennington, R.T. (eds.) Homology in Systematics: Coding Characters for Phylogenetic Analysis. pp. 1-9. Taylor and Francis, London.
7. PENNINGTON, R.T., Prado, D.A. & Pendry, C. (2000). Neotropical seasonally dry forests and Pleistocene vegetation changes. Journal of Biogeography 27: 261-273.
6. Ireland, H. & PENNINGTON, R.T. (1999). A revision of Geoffroea Leguminosae-Dalbergieae. Edinburgh Journal of Botany 56: 329-347.
5. Zamora, N, PENNINGTON, R.T & Stirton, C.S. (1999). Dussia atropurpurea (Leguminosae), a new species from Central America and notes on sarcotesta coloration in Dussia systematics. Edinburgh Journal of Botany 56: 175-180.
4. PENNINGTON, R.T, Aymard, G & Cuello, N. (1997). A new species of Andira from the Venezuelan Guayana. Novon 7(1): 72-74.
3. PENNINGTON, R.T. (1996). Molecular and morphological data provide resolution at different hierarchical levels in Andira. Systematic Biology 45: 496-515.
2. PENNINGTON, R.T. (1995). Cladistic analysis of chloroplast DNA restriction site characters in Andira Leguminosae: Dalbergieae). American Journal of Botany 82: 526-534.
1. PENNINGTON, R.T. & de Lima, H.C. (1995). Two new species of Andira from Bahia Brazil, and the influence of dispersal in determining their distributions. Kew Bulletin 50: 557-566.
(b) Books and volumes authored or edited
6. Marcelo Peña, J.L., PENNINGTON R.T. & Reynel, C (2010). Guia ilustrada de la flora leñosa de los bosques estacionalmente secos de Jaén. Tarea Gráfica Educativa, Perú. 286pp. illus 13 b+w, 338 colour.
5. Reynel, C., Pennington, T.D., PENNINGTON, R.T., Marcelo, J. & Daza, A. (2006, in press). Arboles útiles del ande peruano (Useful Trees of the Peruvian Andes). Tarea Gráfica Educativa, Perú. c. 450 pp: illus 130 b+w, 20 colour.
4. PENNINGTON, R.T., Lewis, G. & Ratter, J.A. (eds.) (2006). Neotropical savannas and dry forests: plant diversity, biogeography and conservation. CRC Press, Florida. 484 pp
3. PENNINGTON, R.T., Richardson, J.E. & Cronk, Q.C.B. (eds). (2004). Plant phylogeny and the origin of major biomes. Philosophical Transactions of the Royal Society, Series B 359: 1453-1656.
2. Reynel, C., Pennington, T.D., PENNINGTON, R.T., Daza, A. & Flores, C. (2003). Árboles útiles de la Amazonia Peruana y sus usos (Useful Trees of the Peruvian Amazon). Tarea Gráfica Educativa, Perú. 509 pp: illus 150 b+w, 20 colour.
1. Scotland, R.W. & PENNINGTON, R.T. (eds.) (2000). Homology in Systematics: Coding Characters for Phylogenetic Analysis. Taylor and Francis, London. 288pp: illus. 58 b+w.
(c) Web
3. Ratter, J.A., Bridgewater S., Milliken W., Pullan M., Oliveira-Filho, A., Ribeiro F., & PENNINGTON R.T. (2011). http://cerrado.rbge.org.uk (website for floristic surveys and general botanical information about Brazilian cerrado vegetation; includes downloadable data and publications)
2. Linares-Palomino, R. (supervised by PENNINGTON, R.T. (10%), Hughes, C.E., Pennington, T.D. and Ratter, J.A.). 2005. Annotated Checklist of the woody plants in Peruvian seasonally dry forests. http://rbg-web2.rbge.org.uk/dryforest/database.htm
1. Reynel, C. and PENNINGTON, R.T. (40%) 2006. www.darwintreediversity.org.pe (website for Darwin work in Peru; includes downloadable publications)
(d) In press/accepted/submitted refereed research papers
1. PENNINGTON, R.T. & Duno de Stefano, R. (submitted). Andira In Flora ilustrada de la Península de Yucatán Mexicana, ed. G. Carnevali.