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Peter Greig-Smith

Jubilee Symposium for the 75th anniversary of the British Ecological Society at University College, London held on April 12-13 1988. Past Presidents gathered for a team photograph: Back row, left to right: A.D. Bradshaw, R.J. Berry, Peter Greig-Smith, A. Macfadyen, C.H. Gimingham, G. Dunnet, L.R.Taylor. Front row from left: C.H. Evans; P.R. Richards, J. Harley, R. Southwood.

Peter Greig-Smith (b. 1922–d. 2003) was a British plant ecologist, founder of the discipline of quantitative ecology in the United Kingdom. A stout supporter 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.

Mathematics in Ecology

The postwar period, and especially the 1950s, saw a rising quest for a truly quantitative approach to community ecology. Community description to that point had been predominantly qualitative and had been based mostly on attempts to extend the principles of Linnaean taxonomy into the classification of multispecies sets. Four innovative ecologists directed their efforts to bring the principles of statistics and mathematics to vegetation analysis: John T. Curtis and Robert H. Whittaker in the United States, and David W. Goodall and Peter Greig-Smith in the United Kingdom. J. T. Curtis, in collaboration with J. Roger Bray, developed the first method of community ordination and also developed the idea of ecological dissimilarity or “ecological distance measure” that later opened the way to the concept of niche overlap. Robert H. Whittaker was the proponent of gradient analysis to address fundamental questions of how plants use space and resources. He provided foundational elements for a quantitative theory of biodiversity and was most active in the areas of plant community analysis, succession, and productivity. David Goodall worked as a professor of agricultural botany at the University of Reading, where he developed a series of studies on plant sampling, distribution, and multivariate analysis. In 1956, however, Goodall immigrated to Australia, where he led a sixty-year-long and extremely fruitful career, becoming one of the world’s most renowned ecologists.

Greig-Smith, eight years younger than Goodall, became the champion of the quantitative approach in plant ecology in Britain. One of the founding fathers of quantitative plant ecology, 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 revised in 1964 and 1983), a must-read for multiple generations of young ecologists.

Early Work

Alexander S. Watt, Greig-Smith's mentor, and one of the most influential British ecologists of the 20th Century.

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 deep inspirational influence can be seen in Greig-Smith’s obituaries of him (Greig-Smith 1982, Greig-Smith 1990). The Biographical Memoirs of the Royal Society show that Watt, in turn, had studied at Cambridge under the mentorship of the legendary ecologist Sir Arthur G. Tansley (1871–1955), a direct academic descendant of T. H. Huxley and Charles Darwin (Tansley had studied with Francis W. Oliver [1864–1951] at the University College, London, who in turn had been mentored by Sir E. Ray Lankester [1847–1929] at Cambridge, a student of George Rolleston [1829–1881] at Oxford, who had been in turn mentored by Thomas Henry Huxley in London). 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 the legendary 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 (a dune grass), and the biogeographic distribution of liverworts to understand the fate of vegetation in the British Isles during the last glaciation.

Greig-Smith’s fondness and admiration for Alexander S. Watt, his first teacher and mentor, is patently recorded in the biographical note he authored for the publication of a book in tribute to Watt’s memory[1]. After Watt’s death, the Royal Society selected Greig-Smith, his most renowned student, to write his biographical memoir[2].

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[3]. This paper was followed two years later by a detailed description of the dune’s multi-species assemblage[4], where he presented a description of the dune vegetation on the Isle of Harris by using, again, quantitative methods to describe the plant community. These early papers already mark his interest for quantification of natural phenomena. In this detailed 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 pounceworts (Lejeunaceae, a family of liverworts)[5]. Apart from the more strictly taxonomic papers, in 1950 he published a brilliant synthesis paper on the taxonomy and biogeography of liverworts, as an early attempt to shed some light on the question of the effects of Quaternary glaciations on the history of the British flora[6]. He tried to evaluate how many of the extant British liverworts had survived Pleistocene glaciations in situ and how many, in contrast, were postglacial migrants from southern latitudes. Almost two decades before the theory of Pleistocene refugia was proposed by Jürgen Haffer for the Amazon[7], Greig-Smith was using plant distributions to understand the vegetation changes induced by Pleistocene glaciations, and setting a visionary direction for Quaternary paleobotany that few could value in its full significance at that time. As part of the Flora of the British Isles series, Greig-Smith drafted a detailed paper on the taxonomy, ecology, and biogeography of the nettles of the United Kingdom, containing a remarkable synthesis of the natural history of these stinging plants and a detailed analysis of the communities where they prosper[8].

Tropical Ecology. In 1940, in collaboration with Paul Richards, Arthur Tansley and Alexander Watt published a seminal paper[9] 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.

Like Darwin in the Brazilian Atlantic Forest, he was fascinated and amazed by the complexity of tropical forests, and in order to try to unravel and explain this complexity, he developed for the first time some 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[10]. Despite the largely descriptive goals of his study, the first study included a number of techniques that at that time were quite novel in vegetation analysis, such as the species-area plots developed by R. 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. Failing to find non-randomness in tree distribution, 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 MS program in ecology that Professor Paul W. Richards had developed at Bangor, and advised many students, from where he recruited many doctoral students and research collaborators. Most of the students, postdocs, and research visitors in his lab later became outstanding ecologists in their own right, such as Georg Grabherr (University of Vienna), Kenneth A. Kershaw (who moved to McMaster University, Hamilton, Ontario), Michael P. Austin (a postdoc at Bangor, studying tropical forests, who later moved to CSIRO at Canberra, Australia), Robert P. McIntosh (University of Notre Dame, South Bend, Indiana), László Orlóci (also a postdoc, later moved to the University of Western Ontario in London, Canada), Timothy F. H. Allen (University of Wisconsin–Madison), Mark O. Hill (Natural Environment Research Council, Swindon, UK), Michael J. Liddle (Monks Wood Experimental Station, Cambridgeshire, UK; later at Griffith University, Queensland, Australia), Michael D. Swaine (University of Aberdeen, Scotland), M. A. Pemadasa (University of Sri Lanka), David J. Gibson (Southern Illinois University, Carbondale), and Tewolde B. G. Egziabher (Environmental Protection Authority, Ethiopia), among many others. The place was a haven for academic exchange: Laszlo Orlóci recalls in his biographical memories[11] 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.

Intellectual Progeny

In Bangor, Greig-Smith took the view that it was up to his students and young postdocs to define the project, develop the scientific hypotheses, and do the work, provided that they included quantitative ecology, his own area of expertise, in their graduate research. With his characteristic shyness and humility, he rarely put his own name on his students’ research papers, so that his enormous influence is visible often only in the acknowledgements of published papers. However, some of the papers written by students and researchers while at Greig-Smith’s lab rank among the highest cited in ecology, and owe much of their originality to Greig-Smith’s mentorship[12].

Pattern and Scaling in Ecological Sciences

Above any other subject, Greig-Smith was fascinated by the phenomenon of plant spatial pattern. Why are some plants regularly spaced while others are densely clumped? — he asked himself. Strongly influenced by his early experience in agricultural experiments using random blocks and nested designs, 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. Although the quest to understand plant pattern at different scales might have been seen by many as a trivial pursuit, his ideas really set the foundations for the development of the concept of scaling in ecological phenomena, a central concept in modern climate change and global sustainability studies.

In the early 1950s, Greig-Smith started using graphs in which he plotted the variance in plant density versus the size of the sample plot, to find peaks that could identify clump size in aggregated distributions. In doing so, he realized that ecosystem properties are scale-dependent: a species can have a regular distribution at a given scale, but be highly clumped at another.

Dunes as Model Ecosystems. One of Greig-Smith’s early research papers, Gemmell, et al. (1953)[13], dealt with the formation of tussocks in Ammophila arenaria and included detailed observations on the clumped distribution of tillers, which form tussock-like clumps in older, fixed dunes but are regularly distributed in the younger, moving dunes. After this initial study, the spatial pattern of dunes and grasslands became a lifelong interest in his ecological research, published in a number of papers with collaborators and students[14]. One of Greig-Smith’s students, Andrew Morton, continued Greig-Smith’s pattern analysis studies in the dunes at Newborough Warren in Anglesey, Wales, near Bangor.

Pattern in Tropical Ecosystems. Below only the study of coastal dunes, Greig-Smith’s interests were strongly focused on tropical ecosystems. After his visit to Trinidad as a young researcher, he developed an interest for African savannas, on the possibility of using his methodology for pattern analysis to understand the importance of intraspecific competition in tropical, strongly rainfall-seasonal ecosystems[15].

Toward a Theory of Scaling. It did not take long for Greig-Smith to realize that many plants form clumps at a certain scale, but that at a larger scale the clumps themselves may be randomly, or even regularly, distributed. In different papers[16], he clearly showed the importance of scaling in the measurement of pattern. He understood that researchers sampling the same vegetation with quadrats of different size could reach diametrically opposed conclusions: a researcher using small quadrats would find that his quadrats harbor either clumps or no plants at all, and would conclude that the species pattern in space follows a clumped distribution. Similarly, a researcher using large quadrats would find a similar number of clumps in each quadrat and would conclude that the species distribution is markedly regular. In reality, both would be right because a species can have a strongly clumped distribution at a certain scale and a regular distribution at another. As president of the British Ecological Society, he wrote an address in in 1979[17], in which he summarized his ideas on the importance of scaling in ecological spatial pattern. Some of his students took his ideas and pursued them further: Mark Hill developed a rigorous mathematical theory for the analysis of pattern at a community level[18], while Kenneth Kershaw, after publishing in co-authorship with Greig-Smith, continued exploring for years the drivers of pattern at different scales.

Scaling in Modern Ecology. The fundamental concept that ecological phenomena could have different behaviors when measured at different scales became really important for understanding large-scale phenomena in the biosphere. The Theory of Scaling became a mainstream concept in ecological research in the late 1980s and 1990s, with the publication of two now-classic papers: Wiens 1989 and Levin 1992[19]. Greig-Smith, however, had figured out the main concepts four decades before, and both John Wiens and Simon Levin recognized him as a scientific pioneer.

Multivariate Classification and Ordination

Greig-Smith was among the first ecologists to understand that multivariate methods were destined to become important tools in quantitative plant ecology. His inclusion of multivariate classification and ordination in his book opened the door for the explosion of analytical approaches and methodological discussions that surged in the 1970s and 1980s. As with pattern analysis, Greig-Smith was ahead of his time: although the potential of multivariate methods—based on matrix algebra—was already recognized in the 1950s, mostly as a result of a growing need for more-rigorous methods in ecology, performing an analysis of a modestly sized floristic matrix of, say, fifty sites and one hundred species was a complicated and enormously time-consuming task. The generalized access to electronic computers, which began in the 1960s, opened the doors for the widespread use of these methods; Bangor’s first computer became available in 1964. Greig-Smith saw in multivariate methods an opportunity to analyze and understand the complex floristic composition of tropical forests, which had fascinated him since his visit to Trinidad in 1948. In 1963 he applied for a grant to investigate the use of quantitative methods to establish whether the highly diverse tropical rainforests showed any organized pattern, particularly in relation to environment, an open question at the time. With these funds, in 1964 he appointed Michael Austin as a postdoctoral research assistant to the project. Three seminal papers were published as a result of their collaboration, highlighting the potential of multivariate methods to analyze floristic patterns in complex, multispecies communities[20]. The power of his approach, merging multivariate descriptive methods with experimental hypothesis testing, is especially noticeable in the work of some of his students like Mike Austin (mentioned previously) and others[21]. Mike Austin later moved to Australia and became one of the country’s leading scientists.

Having laid the foundations of the discipline of multivariate methods, as a committed naturalist and field biologist Greig-Smith never took much interest in abstruse theoretical discussions, always expressing his concern about the risk of becoming too engaged with theoretical refinements of methodology and repeatedly stating his belief that numerical methods are worth developing only if they are to be used on real data in attempts to answer real questions with relevance in the field. The ultimate evidence, for him, was not in the computer output but in the field. In a synthesis paper published in 1980, he insisted that multivariate methods are simply hypothesis-generating procedures, and counseled students to do experimental tests addressing the insights from the classification and ordination analyses rather than taking the results of their analyses as an unchallengeable truth[22].

Novel Approaches to Multivariate Analysis. Greig-Smith’s later papers on multivariate methods, with Mike Swaine and Carlos Montaña[23], were devoted to exploring new and innovative applications for multivariate analysis, including the analysis of time-series data and the exploration of the niche preference of species within a community.

A Seminal Book: Quantitative Plant Ecology

Greig-Smith’s most influential work was his ground-breaking book Quantitative Plant Ecology, used by generations of students and young researchers around the world. The book, first published in 1957, enjoyed two subsequent editions in 1964 and 1982[24].

The success of Greig-Smith’s book triggered the publication of many other similar books, many of them by former Greig-Smith collaborators (such as Kershaw 1964 and Orlóci 1975), and nurtured an explosion of concepts and ideas around the quantitative treatment of ecological data, many of which arose in Canada (such as Pielou 1969) and Australia (such as Williams 1976)[25].

Holism and Reductionism in Ecology

In the 1950s and 1960s, at the time when the School of Plant Biology at Bangor consolidated as a global magnet for plant ecologists, an active scientific debate, excellently analyzed in Barbour (1996)[26], was taking place worldwide on the role of holism and reductionism as the main tools to understand ecological phenomena. In Britain, this debate centered mostly on the use of descriptive methods or the quest to understand patterns in nature, as opposed to the use of experimental methods or the search for ecological processes. In Bangor, in particular, the most vocal proponent of an experimental and reductionist approach to ecological research was John L. Harper, who often criticized quantitative vegetation scientists on their descriptive, rather than experimental, approach to research. Roy Turkington[27] recalls that Harper strongly favored the power of simple, direct experiments and distrusted complex multivariate methods, often arguing that ordination methods were “a formalized description of consequences.” “Nowhere is this holism more apparent than in the way ecology is taught in many schools and universities with its emphasis on complex systems and the ways in which they can be described,” wrote Harper in “After Description”[28], and he immediately added: “There is probably no other science in which students are taught by being dropped into the deep end of complexity.” The contrast between experimental ecology in simplified systems, with only one or a few species, against large-scale pattern-seeking methods led Peter Greig-Smith and John Harper into frequent discussions in the form of good-humored debates following departmental seminars in Bangor, which, as amusingly evoked in Hill (2003) and Matlack (2009)[29], Greig-were conducted amid legendary clouds of pipe smoke for the benefit of the students, who could see firsthand how academic arguments conducted both with rigor and humor can be very stimulating and deeply entertaining.

Greig-Smith, however, was not really a holist. From his very first papers on the tropical forests of Trinidad, he embraced Henry Gleason’s “individualistic” model of plant communities. Tim Allen, a Greig-Smith student in the 1960s, recalls that while in the 1980s “the use of descriptive multivariate methods degenerated into a ritual that community ecologists were expected to perform”[30], Greig-Smith insisted that ecological research should always have experimental tests addressing the insights gained and the hypotheses derived from the quantitative analysis, as in Morton (1974)[31]. Perhaps the main difference with Harper’s single-species, experimental approach lies in Greig-Smith’s interest in complex, multispecies systems, and his opinion that projects should start with observations and the search for patterns, before establishing hypotheses about the processes that drive ecosystem functions. In any case, what is undeniable is that those post-seminar discussions had a tremendous philosophical influence on the development of ecological theory in the last four decades of the 20th Century.

The Enduring Legacy of Peter Greig-Smith

The "Peter Greig-Smith" Orchid Collection at Treborth Botanical Gardens, Bangor, Wales.

Peter Greig-Smith was shy and somewhat introverted, and disliked pomp and show. Scientifically, he was a disciplinarian and somewhat of a traditionalist but was also a strong proponent of the separation between scientific positions and political or social points of view. In a letter to Michael Barbour, he wrote: “There may be parallels between ecology and politics but if so I suspect they are coincidental. I see no parallels between my political or social attitudes and my ecological views.” Possibly as a result of his work at Trinidad as a young researcher in 1953, he always had a kind and supportive attitude toward foreign students, and especially toward those from developing countries. Tewolde B. G. Egziabher[32] describes in eloquent and grateful terms his arrival from Ethiopia to Greig-Smith’s lab in Bangor. Greig-Smith maintained a lifelong interest in tropical ecology. He was a friend and follower of Paul W. Richards, CBE, a professor at Bangor and one of Britain’s most outstanding tropical ecologists. In his obituary of Richards, Greig-Smith recalled the establishment of the legendary MSc course in ecology at Bangor as a program fundamentally directed to meet the needs of students from the tropics[33]. After Trinidad, he visited Nigeria in 1966, where he established a long-lasting relationship with the University of Ife as an external examiner. His passion for tropical ecosystems was palpable in the time and love he devoted to his orchid collection, donated to Bangor’s Botanical Gardens, as described in Retallick (2011)[34].

Generous with his time both for students and colleagues, he was very active in scientific societies. In the 1960s and 1970s he served the British Ecological Society as its secretary (1961–1963), as editor of the Journal of Ecology (1964–1968), and as president (1978–1979), with John Maynard-Smith as his vice president. Always concerned about maintaining a separation between scientific results and political views, in his presidential viewpoint address in the Bulletin of the British Ecological Society, he advised younger ecologists not to become too specialized: “Insight so often comes from initially unconsidered information and ideas,” he wrote, urging colleagues to maintain a broad curiosity for the natural world[35]. His time as editor of the Journal of Ecology was the only period in the society’s recent history where only one editor was selected to shoulder the burden of the task, and he carried out the charge with grace and efficiency. The Bangor School of Plant Biology took a strong blow in the 1980s, when the British government introduced an early-retirement scheme for university staff in order to save money and confront the economic crisis of that time. Both John Harper and Peter Greig-Smith took early retirement, and, although they continued to lecture for some time, they were no longer responsible for the department, and their participation in the recruitment and teaching of graduate students declined. His students remember him as a caring mentor, with a dry but kind sense of humor. His core values were rigor, objectivity, and service to science. Under his leadership, quantitative ecology flourished in Britain and around the world. His intellectual descendants are scattered over all five continents, and the methods he championed now form a fundamental part of global ecological studies.

References

  1. ^ 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, 9–10. Special Publication of the British Ecological Society 1. Oxford: Blackwell Scientific.
  2. ^ Greig-Smith, Peter. 1990. Alexander Stuart Watt: 21 June 1892–2 March 1985. Biographical Memoirs of Fellows of the Royal Society 35):404–423.
  3. ^ Greig-Smith, Peter, Alan R. Gemmell, and Charles H. Gimingham. 1947. Tussock formation in Ammophila arenaria (L) Link. New Phytologist 46(2): 262–268.
  4. ^ 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.
  5. ^ Greig-Smith, Peter. 1948b. Potamolejeunea Holtii (spruce) nov.comb. Transactions of the British Bryological Society 1(2): 108–109. (A new taxonomic treatment for Holt’s pouncewort Lejeunea holtii). Greig-Smith, Peter. 1948c. Leaf cell size in Lejeunea flava (Swartz) Nees. Transactions of the British Bryological Society 1(2): 110–112. (A quantitative analysis of leaf cell size and its geographic correlations for the golden pouncewort Lejeunea flava). Greig-Smith, Peter. 1953. Notes on Lejeuneaceae: I. Microlejeunea diversiloba (spr.) K.M. Transactions of the British Bryological Society 2(2): 285–288. (A taxonomic treatment for the Irish pouncewort Lejeunea diversiloba). 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. (A pioneering attempt at using quantitative traits to classify the species of Lejeunea in the British Isles, developed almost twenty years before the advent of numerical taxonomy and cladistics).
  6. ^ Greig-Smith, Peter. 1950. Evidence from hepatics on the history of the British flora. Journal of Ecology 38(2): 320–344.
  7. ^ Haffer, Jürgen. 1969. Speciation in Amazonian forest birds. Science 165(3889): 131–137.
  8. ^ Greig-Smith, Peter. 1948a. Biological flora of the British Isles: Urtica L. Journal of Ecology 36(2): 339–355.
  9. ^ Richards, 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.
  10. ^ 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.
  11. ^ Orlóci, 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, Otario, Canada, on 27 June 2012. Also available at: https://www.researchgate.net/publication/334432585)
  12. ^ Some of the papers published by young researchers and students at Greig-Smith’s lab during the 1960s and 70s, which do not bear his name as coauthor but certainly are a result of his ideas and mentorship include the following; Austin, Michael P., and László Orlóci. 1966. Geometric models in ecology: II. An evaluation of some ordination techniques. Journal of Ecology 54(1): 217–227. Ezcurra, Exequiel, Miguel Equihua, and Jorge López-Portillo. 1987. The desert vegetation of El Pinacate, Sonora, Mexico. Vegetatio 71(1): 49–60. Hall, John B., and Michael D. Swaine. 1976. Classification and ecology of closed-canopy forest in Ghana. Journal of Ecology 64(3): 913–951. Hill, Mark O. 1973. Reciprocal averaging: An eigenvector method of ordination. Journal of Ecology 61(1): 237–249. Kershaw, Kenneth A. 1957. The use of cover and frequency in the detection of pattern in plant communities. Ecology 38(2): 291–299. McIntosh, Robert P. 1967. An index of diversity and relation of certain concepts to diversity. Ecology 48(3): 392–404. McIntosh, Robert P. 1974. Plant ecology, 1947–1972. In Special issue: 25 years of botany. Annals of the Missouri Botanical Garden 61(1): 132–165. Orlóci, László. 1966. Geometric models in ecology: I. The theory and application of some ordination methods. Journal of Ecology 54(1): 193–215.
  13. ^ Gemmell, Alan R., Peter Greig-Smith, and Charles H. Gimingham. 1953. A note on the behaviour of Ammophila arenaria (L.) Link, in relation to sand-dune formation. Transactions of the Botanical Society of Edinburgh 36(2): 132–136.
  14. ^ Gemmell, Alan R., Peter Greig-Smith, and Charles H. Gimingham. 1953. A note on the behaviour of Ammophila arenaria (L.) Link, in relation to sand-dune formation. Transactions of the Botanical Society of Edinburgh 36(2): 132–136 (A reanalysis of his dune data from the Hebrides, aimed at describing scale-dependent pattern). Gibson, David J., and Peter Greig-Smith. 1986. Community pattern analysis: A method for quantifying community mosaic structure. Vegetatio 66(1): 41–47 (David Gibson, one of Greig-Smith’s last PhD students, extended the idea of species-specific pattern analysis in nested samples to the whole plant community, by using ordination axes summarizing vegetation and soil data and by testing for spatial randomness in the multivariate scores). Greig-Smith, Peter. 1961. Data on pattern within plant communities: II. Ammophila arenaria (L) Link. Journal of Ecology 49(3): 703–708 (Once again, Greig-Smith used his Ammophila data from the Hebrides to prove the importance of detecting pattern and scaling to understand plant spatial dynamics). Liddle, Michael J., and Peter Greig-Smith. 1975a. A survey of tracks and paths in a sand dune ecosystem: I. Soils. Journal of Applied Ecology 12(3): 893–908 (Michael Liddle came to Bangor under a Natural Environment Research Council (NERC) fellowship to study the effect of anthropogenic trampling on natural ecosystems. In this paper, the first of a series of two, he jointly analyzes with Greig-Smith the effect of vehicle trampling on the soils of coastal dunes). Liddle, Michael J., and Peter Greig-Smith. 1975b. A survey of tracks and paths in a sand dune ecosystem: II. Vegetation. Journal of Applied Ecology 12(3): 909–930 (In the second paper, Liddle and Greig-Smith analyze the effect of vehicle trampling on the vegetation of coastal dunes. Liddle eventually moved to Queensland, Australia, where he became a recognized expert on the ecological effects of human trampling on fragile ecosystems such as grasslands and corals). Morton, Andrew J. 1974. Ecological studies of a fixed dune grassland at Newborough Warren, Anglesey: I. The structure of the grassland. Journal of Ecology 62(1): 253–260 (In this first paper on the spatial pattern of fixed dune grassland, Morton, a Greig-Smith doctoral student, used nested-plot pattern analysis to study the scaling effects of soil organic matter and dune landform on plant spatial clumping). Pemadasa, M. A., Peter Greig-Smith, and Peter H. Lovell. 1974. A quantitative description of the distribution of annuals in the dune system at Aberffraw, Anglesey. Journal of Ecology 62(2): 379–402 (Pemadasa, a Sri Lankan botanist studying with Greig-Smith, tested some of the ideas on dune succession—developed by Greig-Smith studying dune perennials in the Hebrides—with the annual plants at the Aberffraw dunes in Anglesey).
  15. ^ Greig-Smith, Peter, and Michael J. Chadwick. 1965. Data on pattern within plant communities: III. Acacia-Capparis semi-desert scrub in the Sudan. Journal of Ecology 53(2): 465–474 (For this paper Greig-Smith collaborated with Michael Chadwick, who had just returned to Cambridge from the University of Khartoum, to test the generalized tenet that desert plants showed a regular pattern as a result from competition for water—disproving the hypothesis). Later in his life, and using data from his visit to Nigeria in 1966, he published a new analysis of pattern in dryland African plants: Greig-Smith, Peter. 1991. Pattern in a derived savanna in Nigeria. Journal of Tropical Ecology 7(4): 491–502 (Although he found spatial aggregation at larger scales, at smaller scales he found a regular pattern, which he attributed to competition for water).
  16. ^ 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 (His first paper of the use of blocked design to understand scaling in plant spatial patterns). Greig-Smith, Peter. 1961. Data on pattern within plant communities: I. The analysis of pattern. Journal of Ecology 49(3): 695–702 (First of a series of three papers aimed at understanding the interaction between plants and their environment and testing hypotheses on the scale and intensity of pattern in plant succession, this paper contains a methodological synthesis of the problem and the analysis of spatial pattern in plants). Greig-Smith, Peter, and Kenneth A. Kershaw. 1958. The significance of pattern in vegetation. Vegetatio 8(3): 189–192 (A methodological note on the problem of plant pattern, published with Kenneth Kershaw, his student at Bangor). Greig-Smith, Peter, Kenneth A. Kershaw, and Derek J. Anderson. 1963. The analysis of pattern in vegetation: A comment on a paper by D. W. Goodall. Journal of Ecology 51(1): 223–229 (Greig-Smith, together with his collaborators Kershaw and Anderson, responded to David Goodall—a British ecologist working in Australia and a contemporary competitor of Greig-Smith as founder and champion of quantitative ecology—defending the nested-block sampling design in the study of plant spatial pattern).
  17. ^ Greig-Smith, Peter. 1979. Pattern in vegetation. Journal of Ecology 67(3): 755–779.
  18. ^ Hill, M. O. 1973. The intensity of spatial pattern in plant communities. Journal of Ecology 61(1): 225–235 (Mark Hill, a Greig-Smith student at Bangor, reused Greig-Smith’s Sudan vegetation data to demonstrate the stabilizing effect of data standardization on pattern estimation under random thinning of individuals).
  19. ^ Levin, Simon A. 1992. The problem of pattern and scale in ecology: The Robert H. McArthur Award Lecture. Ecology 73(6): 1943–1967 (Levin argued that the problem of pattern and scale is the central problem in ecology. Ecological phenomena occur at different scales, and understanding complex problems such as global environmental change requires the interfacing of phenomena that occur on very different scales of space, time, and ecological organization). Wiens, John A. 1989. Spatial scaling in ecology. Functional Ecology 3(4): 385–397 (Wiens’s main tenet was that ecological patterns and processes are scale dependent. Failing to understand these scaling relationships and comparing across ecosystems studied at arbitrary choice of scales may lead to contradictory results, as Greig-Smith had insisted decades before).
  20. ^ Greig-Smith, Peter, Michael P. Austin, and Timothy C. Whitmore. 1967. The application of quantitative methods to vegetation survey: I. Association-analysis and principal component ordination of rain forest. Journal of Ecology 55(2): 483–503 (An early study analyzing floristic variation in tropical forests of the Solomon Islands). Austin, Michael P., and Peter Greig-Smith. 1968. The application of quantitative methods to vegetation survey: II. Some methodological problems of data from rain forest. Journal of Ecology 56(3): 827–844 (One of the first papers analyzing the effect of standardization of species abundance and the completeness of floristic sampling on multivariate analyses). Austin, Michael P., Peter S. Ashton, and Peter Greig-Smith. 1972. The application of quantitative methods to vegetation survey: III. A re-examination of rain forest data from Brunei. Journal of Ecology 60(2): 305–324 (An extensive soil-vegetation analysis performed a decade earlier in Borneo is reexamined by using multivariate methods).
  21. ^ Ezcurra, E. 1987. A comparison of Reciprocal Averaging and Non-Centred Principal Component Analysis. Vegetatio 71(1): 41–48 (Ezcurra, a Greig-Smith student, showed the conceptual similarities between simple PCA and Hill’s reciprocal scoring approach, and applied the results to a large-scale survey of desert vegetation in the Pinacate, which eventually led to the declaration of this region as a UNESCO Biosphere Reserve and a World Heritage Site). Goldsmith, F. B. 1973a. The vegetation of exposed sea cliffs at South Stack, Anglesey: I. The multivariate approach. Journal of Ecology 61(3): 787–818 (In this first paper of a series of two, Goldsmith, a Greig-Smith student, made use of multivariate methods to describe floristic patterns and to generate ecological hypotheses on the dynamics of sea cliff vegetation). Goldsmith, F. B. 1973b. The vegetation of exposed sea cliffs at South Stack, Anglesey: II. Experimental studies. Journal of Ecology 61(3): 819–829 (In the second paper of the series, Goldsmith developed controlled experiments to test the differential effect of interspecific completion and marine salt spray on the establishment and survival of Armeria maritima, a maritime species, and Festuca rubra, an inland grass).
  22. ^ Greig-Smith, Peter. 1980. The development of numerical classification and ordination. Vegetatio 42(1–3): 1–9 (In this outstanding review paper, published in a special issue covering the 1979 symposium “Advances in Vegetation Science,” Greig-Smith made a historical synthesis of the development of multivariate methods in vegetation ecology).
  23. ^ Swaine, Michael D., and Peter Greig-Smith. 1980. An application of principal components analysis to vegetation change in permanent plots. Journal of Ecology 68(1): 33–41 (Instead of the traditional usage of multivariate methods to analyze species-by-sites matrices, Swaine and Greig-Smith used principal-components analysis to explore the time-series trajectory of the same sites sampled at different times). Montaña, Carlos, and Peter Greig-Smith. 1990. Correspondence analysis of species by environmental variable matrices. Journal of Vegetation Science 1(4): 453–460 (Carlos Montaña, a Mexican ecologist, and Greig-Smith used correspondence analysis to investigate the probabilistic preference of species for different ranges of environmental variation, i.e., the environmental niche of the species in a community).
  24. ^ Greig-Smith, Peter. 1957. Quantitative plant ecology. London: Butterworths (The first edition of Greig-Smith’s book was written before the advent of electronic computers, and most of the methods and analysis contained in it were designed to be resolved by hand, or by using mechanical calculators at best). Greig-Smith, Peter. 1964. Quantitative plant ecology. 2d ed. London: Butterworths (Although computing at that time was still restricted to mainframes and punched cards, the second edition of the book acknowledges the accelerated growth of multivariate methods and presents a separate chapter [chapter 7] solely devoted to classification and ordination. Greig-Smith, Peter. 1982. Quantitative plant ecology. 3d ed. Studies in Ecology 9. Berkeley: Univ. of California Press (Written at a time in which the use of mainframe computers was rapidly giving way to desktop devices, the third edition acknowledges the popularization of quantitative methods and presents substantial additions, trying to keep up with a discipline that was exploding both in content and diversity of approaches. Republished in 1992 by Blackwell Sci. Publications, Oxford).
  25. ^ Kershaw, Kenneth A. 1964. Quantitative and dynamic ecology. New York: American Elsevier (Kershaw, a student at Bangor in the 1950s, moved to Canada and published a book compiling his own experience on quantitative methods. Kershaw’s book, which also went through three editions, was widely used as a text for undergraduates and complemented Greig-Smith’s text, which was heavier in theory and mathematical derivations). Orlóci, László. 1975. Multivariate analysis in vegetation research. The Hague: W. Junk (Orlóci, a Hungarian-born Canadian ecologist, visited Bangor in the 1960s, working with Mike Austin on ordination methods. This book, published a few years after he left Bangor, is strongly based on his work and experience while visiting Greig-Smith’s lab). Pielou, Evelyn C. 1969. Introduction to mathematical ecology. New York: Wiley-Interscience (Pielou, a Canadian ecologist, published a series of highly cited books on quantitative methods, the first of which, published in 1970, touched many of the same subjects analyzed in Greig-Smith’s book). Williams, William T., ed. 1976. Pattern analysis in agricultural science. New York: Elsevier Scientific (Bill Williams, together with David Goodall and Mike Dale, visited Bangor in 1964, exploring new ideas that led to the creation of an outstanding school of quantitative ecology in Australia. This book summarizes the Australian experience in quantitative methods, with a very strong emphasis on multivariate analysis).
  26. ^ Barbour, Michael G. 1996. American ecology and American culture in the 1950s: Who led whom? Bulletin of the Ecological Society of America 77(1): 44–51 (A superb analysis of the debate between holism and reductionism that took place in the second half of the 20th century).
  27. ^ Turkington, Roy. 2009. Obituary: Professor John L. Harper FRS CBE (1925–2009). Journal of Ecology 97(5): 835–837 (A brilliant, humorous, and heartfelt obituary for John L. Harper, describing some of his witty quips, or “Harperisms,” criticizing ill-defined ideas and approaches in ecological research).
  28. ^ Harper, John L. 1982. After description. In The plant community as a working mechanism: Produced as a tribute to A. S. Watt. Edited by E. I. Newman, 11–25. Special Publication of the British Ecological Society 1. Oxford: Blackwell Scientific Publications (Harper’s strong call for a more experimental type of ecological research, criticizing community ecologists for their observational and descriptive approach to research).
  29. ^ Hill, Mark O. 2003. Peter Greig-Smith (1922–2003). Bulletin of the British Ecological Society 34(4): 10–11 (Hill, himself a former Greig-Smith student at Bangor, wrote a moving obituary shortly after his passing, rich in biographical detail). Matlack, Glenn R., ed. 2009. Memories of John L. Harper: A remembrance by his friends, students, and colleagues. 2d ed. Bangor, Wales: SPB (An excellent book, edited by Matlack, containing the memories of forty former students and collaborators of John Harper and describing in detail the rich and intense exchange of ideas that were taking place in Bangor during 1960–1990 period. Available online at http://people.virginia.edu/~ja8n/Publications/2011 Memories of John L Harper.pdf).
  30. ^ Allen, Timothy F. H., Amanda J. Zellmer, and C. J. Wuennenberg. 2005. The loss of narrative. In Ecological paradigms lost: Routes of theory change. Edited by Kim Cuddington and Beatrix Beisner, 333–370. Theoretical Ecology 2. Burlington, MA: Elsevier Academic (Tim Allen describes in this text his time at Greig-Smith’s lab at Bangor in the 1980s).
  31. ^ Morton, Andrew J. 1974. Ecological studies of a fixed dune grassland at Newborough Warren, Anglesey: II. Causal factors of the grassland structure. Journal of Ecology 62(1): 261–278 (Following Greig-Smith’s method of using community-level descriptive methods to establish hypotheses and then testing them experimentally, in this second paper Morton uses exclusion experiments and detailed soil analyses to analyze the causes of vegetation pattern).
  32. ^ Egziabher, Tewolde Berhan Gebre. 2002. The human individual and community in the conservation and sustainable use of biological resources. Paper presented as the Darwin Lecture 2002 at the Annual Darwin Initiative meeting, held in London on 14 March 2002 (Tewolde Egziabher, general manager at the Environmental Protection Authority, Addis Ababa, Ethiopia, recalls fondly his time at Bangor at Greig-Smith’s lab: “[in] 1966, just over 36 years ago, [at] Bangor, North Wales, I ‘read’ quantitative plant ecology for a Ph.D. under a pioneer in the field, Professor Peter Greig-Smith. On my way, I took 6 weeks travelling by bus, train, river barge, and even a taxi between two countries, Lebanon and Syria. I learnt a lot on the way, especially how people adapted to different conditions in their local communities. Of course I learnt even more in Bangor, about how plants adapted to different conditions, also in their local communities.” Available online at https://nyeleni.org/IMG/pdf/Tewolde_Darwin_Lecture2002.pdf).
  33. ^ Greig-Smith, Peter 1995. Obituary: Professor P. W. Richards. The Independent, 11 October 1995 (In his obituary of Paul W. Richards, CBE (b. 1908–d. 1995, a famed Bangor botanist who specialized in tropical rainforests), Greig-Smith recalls the establishment of the pioneering MSc course in ecology at Bangor, “with the needs of students from the tropics particularly in mind.”)
  34. ^ Retallick, Simon. 2011. Orchids at Treborth Botanic Garden. Bangoriad: Bangor University’s Alumni and Friends’ Magazine (Spring): 8 (In 2004, a year after his death, Greig-Smith’s outstanding living orchid collection was donated by his wife, Edna, to Bangor University’s Botanic Garden at Treborth. In his memory, it was named the “Peter Greig-Smith Collection.”)
  35. ^ Greig-Smith, Peter 1978. Presidential viewpoint. Bulletin of the British Ecological Society 9(1): 2–3 (In his address as President of the British Ecological Society, Greig-Smith analyzes the risks for ecologists in engaging in advocacy or lobbying, and blurring the distinction between scientific problems and issues of public policy. He also advises younger members of the society against overspecialization, urging them to maintain a broad curiosity for the natural world, natural history, and field observations, which he saw as the main drivers of ecological research questions).


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