What Do Economics and Game Theory Have to Do With Dogs?
“[Darwin] pointed out how, in numberless animal societies, the struggle between separate individuals for the means of existence disappears, how struggle is replaced by cooperation ... He intimated that in such cases the fittest are not the physically strongest, nor the cunningest, but those who learn to cooperate so as to mutually support each other.” —Prince Peter Kropotkin, Mutual Aid: A Factor of Evolution
Huxley vs. Wallace, Competition vs Cooperation
It seems to me that evolutionary science took a huge wrong turn the day it was discovered that some chickens form pecking orders. Suddenly, the Darwinian idea that different species might compete with others over resources within a certain ecological niche was misinterpreted to mean that members of social animal groups were engaged in a constant power struggle for dominance over their fellow group members.
In his book Cooperation Among Mammals (1997), evolutionary biologist Lee Alan Dugatkin details the conflict that arose between early Darwinists Thomas Huxley—a hard-liner when it came to the idea of competition—and Henry Wallace, co-creator with Darwin of the theory of natural selection. Dugatkin writes, “While Darwin (1859) acknowledged that the struggle for existence is often metaphorical, insofar as it is often a struggle against the environment, Huxley … took a more extreme view.”
According to Dugatkin, “Huxley believed the animal world was on the same level of competition found in ancient gladiators. ‘Life is a continuous free fight,’ Huxley wrote, and went on to say that ‘the state war of each [animal] against the other was a normal state of existence.’” However, Wallace, in his book Darwinism (1891), argued the exact opposite, stating that ‘On the whole, the popular idea of the struggle for existence entailing misery and pain on the animal world is the very reverse of the truth.’”
The truth—as Darwin himself said—is that social animals “provide many little services for one another.” And it’s not just about prey species like elk finding safety in numbers, or group predators like wolves finding it easier to hunt large prey as a group. Dugatkin points out that goldfish tend to live longer when their group size increases. They also tend to grow faster when surrounded by more rather than less goldfish. If they were in constant competition with one another, that wouldn’t happen. Dugatkin also points out that social amphibians are able to regenerate lost tails more quickly and that social animals learn new tasks faster than solitary species do. All of this points to the likelihood that sociability and cooperation have a positive effect on survivability, while engaging in internecine battles or threats of aggression would have the opposite effect.
Game Theory and Evolutionary Strategies
In 1973, John Maynard Smith proposed the idea that game theory—which is most often associated with economics—would be very useful in defining a framework of animal contests and strategies into which Darwinian competition could be modeled. His position was that “players” of these evolutionary “games” don’t necessarily act in a rational manner, but have strategies that, when successful, produce higher levels of fitness in some organisms than their competitors. In this model, players do not choose their strategy or have the ability to change it, they are born with it and their offspring will inherit it.
However, no matter how well-developed and successful evolutionary game theory has become, it still explains natural selection through the lens of human thought processes (i.e., strategies) and through the concept of competition between species, as well as competition between members of the same social group, which is a huge mistake.
In their 1998 paper, “Animal Contests as Evolutionary Games,” Mike Mesterson-Gibbons and Eldridge Adams write: “It is not unusual for an exercise in game theory to remain partially inconclusive. On the one hand, game-theoretic models are valuable because they suggest ways to test new ideas. On the other, suggesting a test is not the same thing as conducting it, and the difficulties of doing so should not be underestimated.”
Finding new ways to conduct tests is what science is all about. Yet how useful is the information provided by these tests if the underlying premise isn’t valid? And if these tests are designed with “winners and losers” in mind, aren’t evolutionary game theorists loading the dice in favor of a pre-determined thesis, and thus actually creating contests between animals that might not otherwise exist?
Cheaters Never Prosper (At Least, Not for Long)
One of the ways that evolutionary game players are said to gain an advantage over others is through cheating or free-riding. Both “strategies” are a means of getting more while doing less. But now, a new review of dozens of key ecological studies has found very little evidence to support the belief that “cheating” is widespread. Rice University evolutionary ecologist Emily Jones, the study’s co-lead author: “We find that although there are numerous observations of low-quality partners, there is currently very little support that any of these meet our criteria to be considered cheaters.” She goes on to say, “A behavior is only cheating if it provides one partner with an advantage and also imposes a disadvantage on the other.”
In a similar vein, University of Pennsylvania researchers Alexander J. Stewart and associate professor Joshua B. Plotkin recently offered mathematical proof that the only evolutionary strategies in social animals that can succeed in the long term are generous ones. “Ever since Darwin,” Plotkin writes, “biologists have been puzzled about why there is so much apparent cooperation, and even flat-out generosity and altruism in nature. Our paper provides such an explanation.”
After simulating how some generous strategies would fare in an evolving population, Stewart and Plotkin crafted a mathematical proof showing that not only can generous strategies succeed they’re actually the only approaches that work over the long term.
“Our paper shows that no selfish strategies will succeed in evolution,” said Plotkin. “The only strategies that are evolutionarily robust are generous ones.” 
In another recent paper Christoph Adami of Michigan State writes, “For a short time and against a specific set of opponents, some selfish organisms may come out ahead. But selfishness isn’t evolutionarily sustainable.” 
Adami and his colleagues' research was based on testing a 2012 paper unveiling a newly discovered zero-determinant (or ZD) strategy, which supposedly gave selfish players a guaranteed advantage over others. But Adami et al had serious doubts as to whether this strategy would essentially eliminate cooperation and create a world full of selfish beings. So they used high-powered computing to run hundreds of thousands of games and found that zero-determinant game strategies can never be the product of evolution.
Says Adami, “We found that evolution will punish you if you’re selfish and mean.”
Lack of Cooperation in Wolf Packs?
Wolves are often cited as a prime example of cooperation in animals. Yet research shows that hunting success peaks at about ± 4 wolves. The larger a pack gets, the less successful they are.
In a 2011 article, McNulty, Smith, Mech, et al say there are 2 prevailing hypotheses about why this happens. The interference hypothesis proposes that hunting success is limited because individual predators impede each others’ actions. In the free-rider hypothesis, some pack members are said to consume more than their fair share of a resource, or shoulder less than their fair share of the labor costs. To see if this is true, they measured levels of participation by pack members during various stages of the hunt, using an ethogram, an objective scientific inventory of a set of behaviors.
The idea that hunting success was negatively impacted when some members withheld effort was generally borne out by the fact that the rate of decline was most apparent for the most dangerous task: capture, or biting and restraining prey. In other words, as pack size increased fewer wolves felt like going in for the kill.
“Our study suggests that [some] wolves in large groups (>4 hunters) withheld effort … and likely participated merely to be at hand when a kill was made.”
I think this is very unlikely. It would require a mental process called mental time travel, where an animal is able to hypothesize about possible future events, then form a devious mental plan on how to act in such a way so as to “fool” its pack members and thus derive a purely imaginary future benefit that may or may not materialize.
Since a wolf pack is a cohesive unit where members don’t tend to wander off much to engage in their own activities, whenever the pack goes hunting, all members (except for the very young), go along. And since pack hunting success levels off at anything greater than ± 4 wolves, those who are the most experienced, the most fearless, and the most driven would likely shoulder most of the work. And since according to the latest research, a behavior can only be considered cheating (or “free-riding,” which is a form of cheating) if it provides one member of the pack with an advantage while also causing a disadvantage to others—which doesn’t happen here. This means that these non-participating wolves should not be considered free-riders, but should be thought of as something like the pack’s bullpen. They’re happy to let the starting pitcher play the whole game if he can, but they’re also ready to come in and pitch during the late innings if needed.
They’re a team, after all.
Lee Charles Kelley
“Life Is an Adventure—Where Will Your Dog Take You?”
1. (“From extortion to generosity, evolution in the Iterated Prisoner’s Dilemma,” A. Stewart, J. Plotkin, University of Pennsylvania, July 25, 2013.)
2. (“Evolutionary instability of zero-determinant strategies demonstrates that winning is not everything,” Nature Communications, August 2013.) Adami et al.