There are many different ecological constraints that are collected in the literature under the umbrella of ‘food limitation’ when ecologists try to explain the causes of population changes or conservation problems. ‘Sockeye salmon in British Columbia are declining in abundance because of food limitation in the ocean’. ’Jackrabbits in some states in the western US are increasing because climate change has increased plant growth and thus removed the limitation of their plant food supplies.’ ‘Moose numbers in western Canada are declining because their food plants have shifted their chemistry to cope with the changing climate and now suffer food limitation”. My suggestion here is that ecologists should be careful in defining the meaning of ‘limitation’ in discussing these kinds of population changes in both rare and abundant species.
Perhaps the first principle is that it is the definition of life that food is always limiting. One does not need to do an experiment to demonstrate this truism. So to start we must agree that modern agriculture is built on the foundation that food can be improved and that this form of ‘food limitation’ is not what ecologists who are interested in population changes in the real world are trying to test. The key to explain population differences must come from resource differences in the broad sense, not food alone but a host of other ecological causal factors that may produce changes in birth and death rates in populations.
‘Limitation’ can be used in a spatial or a temporal context. Population density of deer mice can differ in average density in 2 different forest types, and this spatial problem would have to be investigated as a search for the several possible mechanisms that could be behind this observation. Often this is passed off too easily by saying that “resources” are limiting in the poorer habitat, but this statement takes us no closer to understanding what the exact food resources are. If food resources carefully defined are limiting density in the ‘poorer’ habitat, this would be a good example of food limitation in a spatial sense. By contrast if a single population is increasing in one year and declining in the next year, this could be an example of food limitation in a temporal sense.
The more difficult issue now becomes what evidence you have that food is limiting in either time or space. Growth in body size in vertebrates is one clear indirect indicator but we need to know exactly what food resources are limiting. The temptation is to use feeding experiments to test for food limitation (reviewed in Boutin 1990). Feeding experiments in the lab are simple, in the field not simple. Feeding an open population can lead to immigration and if your response variable is population density, you have an indirect effect of feeding. If animals in the experimentally fed area grow faster or have a higher reproductive output, you have evidence of the positive effect of the feeding treatment. You can then claim ‘food limitation’ for these specific variables. If population density increases on your feeding area relative to unfed controls, you can also claim ‘food limitation of density’. The problems then come when you consider the temporal dimension due to seasonal or annual effects. If the population density falls and you are still feeding in season 2 or year 2, then food limitation of density is absent, and the change must have been produced by higher mortality in season 2 or higher emigration.
Food resources could be limiting because of predator avoidance (Brown and Kotler 2007). The ecology of fear from predation has blossomed into a very large literature that explores the non-consumptive effects of predators on prey foraging that can lead to food limitation without food resources being in short supply (e.g., Peers et al. 2018, Allen et al. 2022).
All of this seems to be terribly obvious but the key point is that if you examine the literature about “food limitation” look at the evidence and the experimental design. Ecologists like medical doctors at times have a long list of explanations designed to sooth the soul without providing good evidence of what exact mechanism is operating. Economists are near the top with this distinguished approach, exceeded only by politicians, who have an even greater art in explaining changes after the fact with limited evidence.
As a footnote to defining this problem of food limitation, you should read Boutin (1990). I have also raved on about this topic in Chapter 8 of my 2013 book on rodent populations if you wish more details.
Allen, M.C., Clinchy, M. & Zanette, L.Y. (2022) Fear of predators in free-living wildlife reduces population growth over generations. Proceedings of the National Academy of Sciences (PNAS), 119, e2112404119. doi: 10.1073/pnas.2112404119.
Boutin, S. (1990). Food supplementation experiments with terrestrial vertebrates: patterns, problems, and the future. Canadian Journal of Zoology 68(2): 203-220. doi: 10.1139/z90-031.
Brown, J.S. & Kotler, B.P. (2007) Foraging and the ecology of fear. Foraging: Behaviour and Ecology (eds. D.W. Stephens, J.S. Brown & R.C. Ydenberg), pp. 437-448.University of Chicago Press, Chicago. ISBN: 9780226772646
Krebs, C.J. (2013) Chapter 8, The Food Hypothesis. In Population Fluctuations in Rodents. University of Chicago Press, Chicago. ISBN: 978-0-226-01035-9