Ten papers that shook my world – Solomon (1949) – quantifying predator efficiency

Solomon, M. E. (1949). The natural control of animal populations. Journal of Animal Ecology, 18, 1-35.*


According to Google Scholar there are 1149 (only 773 in 2013)citations to this paper, an average of 17.1 citations per year, compared with the 12.5 I reported back in 2013.  Although influential, it had a slow start, only 383 citations being recorded for it between 1949 and 1993. Since 2000 it has averaged about 48 citations a year (760 in total), 225 of those since 2013.** To the modern reader this paper comes across as wordy and discursive, more like a popular article than a scientific paper. This does not, however, mean that the science and the man behind the article were not first class. Journals had less pressure on their space in those days and scientists had more time to think and read. If only it were so now. Despite the relatively low number of citations, this paper has had an immense influence on the study of population dynamics, although I will have to confess, that for my generation who were undergraduates in the 1970s, Solomon’s little Study in Biology*** book, Population Dynamics, published in 1969, was our main, if not only, encounter with his work.

Solomon 1

Making sure that nobody could claim my copy of Population Dynamics

Solomon terms

Here Solomon introduces the term functional as in a density related response

Nowadays we remember the paper as the first one to formalise the term ‘functional response’ although the early citations to this paper are in reference to density dependence, competition, population regulation and population variability e.g. (Elton 1949; Glen 1954; Southwick 1955; Bakker 1963). Interestingly, one of the earlier papers to cite Solomon, (Burnett 1951) presented functional response curves but did not mention the term (Watt (1959)). To add further insult to injury, Holling (1959) in the same year, in his classic paper in which he described and numbered the types of functional responses did not even refer to Solomon, rather deferring to Watt’s paper (loc. cit.). Since then, with the likes of Varley, Gradwell and Hassell (1973) and luminaries such as Bob May (May 1978), this paper has been cited often, and justifiably, and continues to influence us to this day, including the author of this eulogy (Aqueel & Leather 2012). This paper as well as being  the first one to formalise the term ‘functional response’ was the first attempt to draw together the disparate conceptual strands of the first half of the twentieth century work on population dynamics in one coherent whole. Truly, a remarkable and very influential paper.



Aqueel, M. A., & Leather, S. R. (2012) Nitrogen fertiliser affects the functional response and prey consumption of Harmonia axyridis (Coleoptera: Coccinellidae) feeding on cereal aphids. Annals of Applied Biology, 160, 6-15.

Bakker, K. (1963) Backgrounds of controversies about population theories and their terminologies. Zeitschrift fur Angewandte Entomologie, 53, 187-208.

Burnett, T. (1951) Effects of temperature and host density on the rate of increase of an insect parasite. American Naturalist, 85, 337-352.

Elton, C. (1949) Population interspersion: an essay on animal community patterns. Journal of Ecology, 37, 1-25.

Glen, R. (1954) Factors that affect insect abundance. Journal of Economic Entomology, 47, 398-405.

Holling, C. S. (1959) Some characteristics of simple types of predation and parasitism. Canadian Entomologist, 91,385-398.

May, R. M. (1978) Host-parasitoid systems in patchy environments: A phenomenological model. Journal of Animal Ecology, 47, 833-844.

Solomon, M. E. (1969) Population Dynamics. Edward Arnold, London.

Southwick, C. H. (1955) The population dynamics of confined house mice supplied with unlimited food. Ecology, 36, 212-225.

Varley, G. C., Gradwell, G. R. & Hassell, M.P. (1973) Insect Population Ecology: An Analytical Approach. Blackwell Scientific Publications, Oxford.

Watt, K. E. F. (1959). A mathematical model for the effect of densities of attacked and attacking species on the numbers attacked. Canadian Entomologist, 91, 129-144.


Post script

A few months ago I was privileged to be given Robert Tillyard’s excellent The Insects of Australia and New Zealand (first published in 1926), by a former colleague of mine.


What made this even more special was that it had originally belonged to the great Maurice Solomon when he was a student, and contained some of his original annotations and revision notes.

Solomon combo



*This is an expanded and updated version of the article I wrote as part of the British Ecological Society’s Centenary celebrations in 2013

**It is probably wishful thinking, but I might be tempted to think that by writing about this influential but somewhat overlooked paper, I increased the number of citations, so had a positive influence 🙂

***The Studies in Biology series, published by the then Institute of Biology (Now Royal Society of Biology), were excellent little books and the series on plant physiology were the main reason that I passed my first year plant physiology module as an undergraduate at Leeds University. I am reliably informed that there are plans to revive the series next year.




Filed under Entomological classics, Ten Papers That Shook My World

3 responses to “Ten papers that shook my world – Solomon (1949) – quantifying predator efficiency

  1. Marlies

    Oh, I loved those little Studies on Biology – books as well! I still have three of them. One being the one by Mike Hassel: “The dynamics of competition en predation”. I used that for an postgraduate course in Leiden in the early ’80s. Good to hear about the plans to revive the series!


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