Peter Greig-Smith (1922–2003) was a British plant ecologist, founder of the discipline of quantitative ecology in the United Kingdom. He had a deep influence across the world on vegetation studies and plant ecology, mostly from his book Quantitative Plant Ecology, first published in 1957 and a must-read for multiple generations of young ecologists. In 1952, Greig-Smith was hired by the University College of North Wales at Bangor, Wales (now Bangor University), where he spent the rest of his academic career. His book Quantitative Plant Ecology spread the ideas of rigorous, quantitative ecological methods among the world’s research community, and made his lab at Bangor a magnet for plant biologists with an interest in mathematical and statistical methods. A stout supporter of Bangor’s graduate program in ecology, he recruited and advised many students and research collaborators from all continents.
A prominent member of the British Ecological Society, he was first elected to council in 1957, and in 1961 he became honorary secretary. He held this post until 1964, when he resigned to become editor of the Journal of Ecology, the society’s scientific publication. He remained as editor until 1968. In 1977, he was elected vice president of the society, and served as president in 1978 and 1979.
Early work
Greig-Smith went to school in Birmingham, United Kingdom, before going to Downing College, Cambridge, where he was inspired by the teachings of Alexander S. Watt, the first British botanist to seriously study the phenomenon of pattern and scaling in plant communities from a quantitative perspective. Watt’s influence can be seen in Greig-Smith’s obituaries of him.Greig-Smith, Peter. 1982. A. S. Watt, F.R.S.: A biographical note. In The plant community as a working mechanism: Produced as a tribute to A. S. Watt. Edited by E. I. Newman, pp. 9–10. Special Publication of the British Ecological Society 1. Oxford: Blackwell Scientific.Greig-Smith, Peter. 1990. Alexander Stuart Watt: 21 June 1892 – 2 March 1985. Biographical Memoirs of Fellows of the Royal Society 35): pp. 404–423.
In 1944, after graduating from Cambridge, Greig-Smith went to the Imperial College’s Field Station at Slough, where he studied the effect of herbicides on oilseed crops under the direction of Geoffrey E. Blackman, founder of the science of selective herbicides in Britain. The Slough herbicide tests, part of Britain’s war effort, involved the planting of large-scale experiments with random block and nested designs, an experience that played a central role in nurturing Greig-Smith's interest in statistical methods and experimental plot designs.
In 1945, he moved to the University of Manchester as lecturer in botany. At Manchester, he cultivated a wide array of research interests, ranging from the taxonomy of liverworts and stinging nettles, the causes of spatial pattern in dune plants, the quantitative morphology and tussock formation of Ammophila arenaria (marram grass), and the biogeographic distribution of liverworts to understand the fate of vegetation in the British Isles during the last glaciation.
Dune ecology Greig-Smith’s first published paper dealt with the varying growth forms of the dune grass Ammophila arenaria in calcareous sand dunes on the Isle of Harris, Outer Hebrides.Greig-Smith, Peter, Alan R. Gemmell, and Charles H. Gimingham. 1947. Tussock formation in Ammophila arenaria (L) Link. New Phytologist 46(2): 262–268. This paper was followed two years later by a detailed description of the dune's plants,Gimingham, Charles H., Alan R. Gemmell, and Peter Greig-Smith. 1949. The vegetation of a sand-dune system in the Outer Hebrides. Transactions and Proceedings of the Botanical Society of Edinburgh 35(1): 82–96. using quantitative methods to describe the plant community. These early papers already mark his interest for quantification of natural phenomena. In this morphometric analysis of Ammophila arenaria, for example, Greig-Smith measured in detail the internode length of the plant’s rhizomes and noted the species’ tendency to form tussock-like clumps in older dunes and longer, exploratory rhizomes in unstable, younger dunes.
Taxonomy and biogeography Some of Greig-Smith’s early work was devoted to taxonomy and biogeography, trying to bring a quantitative approach to these branches of the biological sciences. Much of his taxonomic work was devoted to the evolutionary taxonomy and ecology of some liverworts (Lejeuneaceae).Greig-Smith, Peter. 1948b. Potamolejeunea Holtii (spruce) nov.comb. Transactions of the British Bryological Society 1(2): 108–109.Greig-Smith, Peter. 1948c. Leaf cell size in Lejeunea flava (Swartz) Nees. Transactions of the British Bryological Society 1(2): 110–112.Greig-Smith, Peter. 1953. Notes on Lejeuneaceae: I. Microlejeunea diversiloba (spr.) K.M. Transactions of the British Bryological Society 2(2): 285–288.
In 1950, he published a synthesis paper on the taxonomy and biogeography of liverworts,Greig-Smith, Peter. 1954. Notes on Lejeuneaceae: II. A quantitative assessment of criteria used in distinguishing some British species of Lejeunea. Transactions of the British Bryological Society 2(3): 458–469. as an early attempt to shed some light on the question of the effects of Quaternary glaciations on the history of the British flora.Greig-Smith, Peter. 1950. Evidence from hepatics on the history of the British flora. Journal of Ecology 38(2): 320–344. He tried to evaluate how many of the extant British liverworts had survived these glaciations in situ and how many, in contrast, were postglacial migrants from southern latitudes. This work as done almost two decades before the theory of glaciation refugia was proposed by Jürgen Haffer for the Amazon.Haffer, Jürgen. 1969. Speciation in Amazonian forest birds. Science 165(3889): 131–137.
Greig-Smith also wrote a synthesis of the natural history of nettles of the United Kingdom with a detailed analysis of the plant communities where they prospered.Greig-Smith, Peter. 1948a. Biological flora of the British Isles: Urtica L. Journal of Ecology 36(2): 339–355.
Tropical ecology In 1940, Paul Richards, Sir Arthur Tansley, and Alexander Watt published a seminal paperRichards, Paul W., Arthur G. Tansley and Alexander S. Watt. 1940. The recording of structure, life form and flora of tropical forest communities as a basis for their classification. Journal of Ecology 28 (1): 224–239. on the classification of tropical rainforests revealing a fascination with the complexity of tropical vegetation that was halted by the war, but that Watt passed down later to Greig-Smith, his student at Cambridge.
In July 1948, Greig-Smith received a research grant from the Colonial Office to visit the island of Trinidad to study tropical rainforests. Encouraged by E. Ashby, an early pioneer of quantitative methods who was then professor of botany at Manchester, he left the university for six months to study secondary succession in Trinidad, in collaboration with colleagues of the Imperial College of Tropical Agriculture on the island. In order to try to unravel and explain this complexity, he developed novel statistical methods to test hypotheses on the association between species and the spatial pattern of saplings and adult trees in the field. An avid naturalist, he also developed in Trinidad a passion for orchids, which led him later to start cultivating his own orchid collection in the United Kingdom. After his visit to Trinidad in 1948, Greig-Smith drafted two detailed papers on secondary tropical forests.Greig-Smith, Peter. 1952a. Ecological observations on degraded and secondary forest in Trinidad, British West Indies: 1. General features of the vegetation. Journal of Ecology 40 (2): 283–330.Greig-Smith, Peter. 1952b. Ecological observations on degraded and secondary forest in Trinidad, British West Indies: 2. Structure of the communities. Journal of Ecology 40(2): 316–330.
The first study included a number of techniques that at that time were novel in vegetation analysis, such as the species-area plots developed by Ronald A. Fisher and Henry Gleason, size-specific survivorship, and quantitative floristic tables.
In the second Trinidad paper, he also used nested analyses of variance (ANOVAs), comparisons with the Poisson distribution, and variance-to-mean ratios to test and understand spatial patterns in the trees of the secondary forest. He concluded his paper questioning Frederic Clement’s concept of plant communities as complex supra-organisms, and in support of Gleason's individualistic model of plant communities.
Bangor
In 1952, Greig-Smith moved to the University College of North Wales at Bangor, where he spent the rest of his career. After the publication of his book Quantitative Plant Ecology in 1957, which rapidly spread among the research community and became a success, Bangor became a magnet for plant biologists with an interest in quantitative methods. Greig-Smith was keenly interested in the MSc program in Ecology that Professor Paul W. Richards had developed at Bangor, and advised many Masters' students from where he recruited many doctoral students and research collaborators. The place was a haven for academic exchange: Laszlo Orlóci, a Professor of Ecology at the University of Western Ontario, Canada,Orlóci, László. 2013. Multivariate Analysis in Vegetation Research. Springer, Netherlands. 276 pp. recalls in his biographical memoriesOrlóci, László. 2019. Looking back: A life of statistical ecology. 3rd ed. Scada Publishing, Canada. Online Edition: https://www.amazon.com/dp/1796403148 (edited transcript of Orlóci’s talk at a symposium in his honor held at the University of Guelph, Guelph, Ontario, Canada, on 27 June 2012. Also available at: https://www.researchgate.net/publication/334432585) that during 1964 alone, Greig-Smith's lab at Bangor received the visit of Robert Sokal and Peter Sneath, founders of the discipline of numerical taxonomy, and David Goodall, William T. Williams, and Mike Dale, who were creating initial content for a solid school of statistical ecology in Australia.
Pattern and scaling in ecological sciences
Greig-Smith was fascinated by the phenomenon of plant spatial pattern, defined by the question: why are some plants regularly spaced while others are densely clumped? He realized that plants could be clumped at a certain scale, but that those clumps, in turn, could be randomly, or even regularly distributed at larger scales. Patterns and processes in ecology, he concluded, can have different behaviors at different scale. Strongly influenced by his early experience in agricultural experiments using random blocks and nested designs,Greig-Smith, Peter. 1952. The use of random and contiguous quadrats in the study of the structure of plant communities. Annals of Botany 16 (2): 293–316 he sought to answer this question by using, as his core tool, a series of quadrats nested within each other at different scales.
Much of Greig-Smith's published work was devoted to advancing the theory and methods of plant pattern, using his nested-quadrat approach as a basic tool. His fundamental idea that the perceived properties of ecological phenomena are scale-dependent set the foundations for the development of a theory of scaling in ecology, a central concept in modern climate change and global sustainability studies.
Dunes as model ecosystems