Spetner & Milton's ignorance of natural history

While close examination of the anti-evolutionists’ writings reveals broad ignorance of evolutionary theory, for some of the writers, their claims reveal a severe lack of knowledge of the natural history of organisms as well.

Dr. Lee Spetner makes several claims in his book, Not By Chance!, that are abundantly refuted by ecological research and could be rebutted by anyone with good knowledge of the natural history of a range of organisms. Similar claims that are similarly false are echoed by Richard Milton in Shattering the Myths of Darwinism. Spetner claims (p. 15) that 'Predator animals do not overexploit prey populations. They live only on the income from their resources and preserve the principal for posterity.' Apparently Spetner thinks that this claim would undermine neo-Darwinian theory, although in fact it would not. The question is whether predators are an agent of selection on their prey, and it is not necessary that they overexploit the prey in order to act as an agent of selection. Spetner also thinks that organisms rarely compete, and therefore seldom experience the ‘struggle for existence’ that might lead to natural selection on a population. It is worth quoting from Spetner (p. 16) at length to understand the magnitude of his error:

Later discoveries, however, have shown that animal populations do not follow Malthus’s prognosis for the human race. Darwin erred in the insight that led him to his theory of evolution. Animal populations generally do not hug the brink of disaster. Population size is not controlled by starvation, disease or predation (Wynne-Edwards 1986). … Moreover, predator animals do not overexploit prey populations. Man seems to the only predator that overexploits its resources and drives them to extinction. Animal predators generally take only the excess of the prey population (Wynne-Edwards 1986). They live, so to speak, only on the income from their resources and preserve the principal for posterity. Wynne-Edwards (1986) has suggested that animals generally manage their food resources by controlling their own numbers. Populations are kept in check not by the extrinsic forces of mass starvation or disease, but by intrinsic forces built into the animals themselves. ... Plants do not proliferate in a field to the point where they become overcrowded. They do not engage in a ‘struggle for existence’ where natural selection would preserve the strong and destroy the weak.

Spetner repeats the claim later in his book (p. 199): 'Plants and animals hardly ever increase their numbers so much that they exhaust their resources. They stop increasing well before they deplete their habitat.'

Unfortunately, Spetner is not alone among the anti-evolutionists in his claims about presumed scarcity of competition and predation having any role in natural selection. Richard Milton, in Shattering the Myths of Darwinism, makes the following absurd claims:

Talk of survival immediately conjures up a lurid vision of competition between the various forms of animal life in a hostile world: competition for scarce resources of food and living space, of ‘nature red in tooth and claw’ as Tennyson pictured it for his enthralled Victorian readers. In reality such competition is very rarely found in nature. One conservative estimate is that there are at least 22,000 common species of fish, amphibians, reptiles, mammals, and birds. In addition there are said to be at least 1 million common insect species. Some of the thousands of species -- notably humans -- do compete aggressively, killing competitors for living space and food. But the species that do are very much in the minority. The overwhelming majority of creatures do not fight, do not kill for food and do not compete aggressively for space in a way that results in the ‘loser’ dying out.’ (p. 124-125) ‘The majority of carnivores do not feed on prey that they themselves have just killed but rather are scavengers or carrion feeders. This includes legendary hunters such as lions or sharks who frequently eat as a result of not their own direct efforts but those of another lion or shark.

There are two profound errors in Spetner’s and Milton’s claims, one of which was hinted at above. The first error is that it is NOT necessary for organisms to exhaust their resources in order for them to compete. Selection can act via competition as long as the organism actually compete; this can occur prior to resources being completely exhausted. Of course, neither Spetner nor Milton clarify what he means by 'exhaust' or 'deplete'. Indeed, rarely in nature would every last bit of a resource be depleted by the consumer organisms; most organisms have some minimal threshold of resource availability below which they cannot sustain themselves, thus competition can readily occur at resource levels above absolute zero (Tilman 1981). These writers’ second major error is the simple fact that many, many cases are known where organisms DO severely deplete their resources -- either as competitors competing for some nonliving resource, or as predators driving their prey to scarcity. Both of these examples demonstrate Spetner’s severe lack of knowledge about natural history. Below we will expound on the second point.

Do organisms deplete their resources to levels that impose constraints on survival and reproduction (the main components of fitness)? Spetner & Milton say 'no', but extensive basic and applied ecological research says 'yes'. Literally hundreds of studies have demonstrated the effects of depleted light levels on growth of trees in dense stands (Mohler et al. 1978). Smaller individuals usually have stunted growth and often die for lack of light. In fact, the widespread observation that plants die at a predictable rate in dense stands that are growing, has become known as the ‘self-thinning law’ of plant ecology (White 1980, Weller 1990). This commonly-observed mortality pattern is a result of competition among the neighboring plants, which have depleted their resources to a level that the smallest individuals cannot survive. Numerous studies have also documented similar patterns among animals. A classic example is the study by Branch (1975) of limpets on the coast of South Africa. Branch studied limpets at naturally-occuring densities that spanned a tenfold range, from 125 to 1225 per square meter; as density increased, limpet size decreased, such that there was a asymptote in total limpet biomass per square meter. Branch further demonstrated a strong correlation between limpet size and reproductive output, showing that the competitive effect at high densities translated into fitness differences among the limpets. Numerous other studies document the role of competition in dense populations, as a result of reduced resource availability (Begon & Mortimer 1991). Spetner’s and Milton’s claim regarding the scarcity of competition is flat wrong.

The claim that predators do not suppress prey populations is also wrong. Many studies that involve the removal or addition of a species to an ecosystem document drastic changes in the abundance of their prey, whether that prey be plants or other animals. For example, the recent reintroduction of wolves to Yellowstone National Park has had a direct negative effect on the density and survival of elk (Ripple & Beschta 2004, Hebblewhite et al. 2005). The Yellowstone wolves have also caused decreases in abundance of smaller predators, probably through a combination of predation and competition (Crabtree & Sheldon 1999). Nelson & Mech (2006) describe how wolves completely extirpated deer from a 3000 km2 section of northeastern Minnosota by the mid-1970s, and have prevented recolonization by deer for the ensuing 30 years. In Yellowstone, the areas where the elk no longer graze heavily have experienced drastic increases in vegetation abundance, demonstrating that the elk were suppressing plant abundance; this three-level chain of effect (predator-plant eater-plants) is called a trophic cascade in ecological circles, and has been documented in a number of ecosystems (Pace et al. 1999, Schmitz et al. 2000). Such cascading effects that occur two or three levels down the food chain from an apex predator can only appear if prey species abundances are controlled by the consumer animals that are higher on the food chain. The conclusion is that predators (including herbivores) often dramatically reduce their prey (plant or animal) abundance, in direct refutation of Spetner’s and Milton’s claims to the contrary.

Finally, Milton’s claim that even charismatic, apex predators are actually predominantly scavengers is flat wrong. As usual, Milton provides nothing to back up his baseless claim. A very cursory examination of the relevant literature reveals the falsehood of his assertion. Funston et al. (1998) studied hunting and feeding behavior of lions in Kruger National Park of South Africa, and reported that they 'acquire most of their food by hunting rather than scavenging.' They documented that among three groups of lions, the percentage of carcasses scavenged rather than captured ranged from 2 to 40%. Stander (1992) conducted a similar study in Etosha National Park in Namibia and reported that 'lions scavenged rarely'. In the boreal forest of northern Scandinavia, the European lynx diet was studied by Odden et al. (2006), and they found that 'most of the diet was obtained by predation, although we did document several cases of scavenging.' Karanth (1993) documented predation and diet in India of three carnivores: tigers, leopards, and dholes, and found that 'carrion was inconsequential in the diets' of all three predators. Stahler et al. (2006) studied wolves in Yellowstone National Park and found that they scavenge only when they are unsuccessful in capturing live prey for several days, which was rare because a pack on average killed a prey animal every 2-3 days. In another study in northern Scandinavia, Linnell & Strand (2002), report that for the diminutive arctic fox, 'there is little evidence for the importance' of scavenging. As far back as 1972, Kruuk documented that even the spotted hyena, widely believed to be predominantly a scavenger, got only 1/3 of their diet by scavenging in the Serengeti, and less elsewhere.

The examples cited above resoundingly refute both aspects of Spetner’s and Milton’s claims. Competition and predation need not be as intense as these authors imply, and the existence of strong effects of competitors on each other, and predators on prey abundances, decidely disproves such claims.

Literature Cited

Begon, M., and M. Mortimer. 1991. Population Ecology, A Unified Study of Animals and Plants. Third Edition. Sinauer Associates, Sunderland, MA.

Branch, G.M. 1975. Intraspecific competition in Patella cochlear Born. Journal of Animal Ecology 44: 263-281.

Crabtree, R.L. and J.W. Sheldon. 1999. Coyotes and canid coexistence in Yellowstone. Pp. 429-440 in T.W. Clark, A.P. Curlee, S.C. Minta, and P.M. Kareiva, editors, Carnivores in Ecosystems: the Yellowstone Experience. Yale University Press, New Haven, CT.

Funston, P.J., M.G.L. Mills, H.C. Biggs, and P.R.K. Richardson. 1998. Hunting by male lions: ecological influences and socioecological implications. Animal Behavior 56: 1333-1345.

Hebblewhite, M., C.A. White, C.G. Nietvelt, J.A. McKenzi, T.E. Hurd, J.M. Fryxell. 2005. Human activity mediates a trophic cascade caused by wolves. Ecology 86: 2135-2144.

Karanth, K.U. 1993. Predator-prey relationships among large mammals of Nagarahole National Park (India). PhD thesis, Mangalore University.

Kruuk, H. 1972. The Spotted Hyena. University of Chicago Press, Chicago, IL.

Linnell, J.D.C. and O. Strand. 2002. Do arctic foxes Alopex lagopus depend on kills made by large predators? Wildlife Biology 8: 69-75.

Mohler, C., P.L. Marks, and D.G. Sprugel. 1978. Stand structure and allometry of trees during self-thinning of pure stands. Journal of Ecology 66: 599-614.

Nelson, M.E., and L.D. Mech. 2006. A 3-decade dearth of deer (Odocoileus virginianus) in a wolf (Canis lupus)-dominated ecosystem. American Midland Naturalist 155: 373-382.

Odden, J., J.D.C. Linnell, and R. Andersen. 2006. Diet of Eurasian lynx, Lynx lynx, in the boreal forest of southeastern Norway: the relative importance of livestock and hares at low roe deer density. European Journal of Wildlife Research 52: 237-244.

Pace, M.L., J.J. Cole, S.R. Carpenter, and J.F. Kitchell. 1999. Trophic cascades revealed in diverse ecosystems. Trends in Ecology & Evolution 14: 483-488.

Ripple, W.J., and R.L. Beschta. 2004. Wolves and the ecology of fear: can predation risk structure ecosystems? BioScience 54: 755-766.

Schmitz, O.J., P.A. Hamback, and A.P. Beckerman. 2000. Trophic cascades in terrestrial systems: a review of the effects of carnivore removals on plants. American Naturalist 155: 141-153.

Stahler, D.R., D.W. Smith, and D.S. Guernsey. 2006. Foraging and feeding ecology of the gray wolf (Canis lupus): lessons from Yellowstone National Park, Wyoming, USA. Journal of Nutrition 24: 1923s-1926s.

Stander, P.E. 1992. Foraging dynamics of lions in a semiarid environment. Canadian Journal of Zoology 70: 8-21.

Tilman, D. 1981. Resource Competition and Community Structure. Princeton University Press, Princeton, NJ.

White, J. 1980. Demographic factors in populations of plants. Pp. xx-xx in O.T. Solbrig, editor, Demography and Evolution in Plant Populations. Blackwell Scientific, Oxford, UK.