Saturday, January 02, 2010

To survive and reproduce in good times and bad

jte asks:

Is there reason to believe, or evidence to support, that the forms of evolution occurring among species during a period of abundant resources is different from the forms of evolution occurring among species during a period of deficient resources?

It seems that a lot of the argument in evolutionary theory is that it takes a lot of energy to grow extra and useless appendages or what have you, so if they really are useless, you'd expect them to evolve away. But if resources are abundant--energy is not a particularly limiting factor--do you then get a scenario in which all kinds of wacky and useless appendages appear and are not attritioned away? Which gives those appendages time to hang around enough to be available when the environment changes and all of a sudden they are useful and confer an advantage?

Or something like that?

I wouldn't go so far as to say the "forms of evolution" are different. In good times and bad evolution acts through natural selection, genetic drift, mutation and all the same basic mechanisms. Rather I would say that selection acts of different traits, or favors different forms of those traits, depending on if times are good or bad. One excellent example of this has been documented by Peter and Rosemary Grant in long term studies of Darwin's Finches on the Galapagos Islands. The climate in the Galapagos is impacted hugely by the El Nino/La Nina climate cycles. In some parts of the cycle, the islands are cool and damp, vegetation grows lush, and there are lots of big seeds to be had. In other parts of the cycle, it is very hot and very dry and only the desert plants with their tiny little seeds are producing. In the good years, the finches with the big bills can eat lots of big seeds, and reproduce like mad. In only a few years the population of one finch species is dominated by big-billed finches. Then when the rains stop, the population starts to crash, and the finches with the little bills good for extracting and opening small seeds are much more likely to survive. After a few years of that, the population of that same species is again dominated by small billed finches. This isn't individuals developing differently depending on the food supply, this is just massive, cyclical natural selection driving the population's genetic make-up around in circles.

On a much larger time scale, generalists are much more likely to survive large extinction events, while specialists often dominate in habitats that have been very stable for millions of years. Consider which of each of these pairs of species is in greater danger of extinction?

German Cockroaches or Lord Howe Island Woodeating Cockroaches
The Black Rat or the Salt-Marsh Harvest Mouse
The Common Pigeon or the Mariana Fruit Dove
The Common Raccoon or the Cozumel Raccoon
Goats or Alpine Ibex
Humans or Sumatran Orangutans

In each case the generalist are doing fine, while their specialized relatives can't cope with change. The fossil record shows multiple examples of large groups going extinct when the coprolites hit the fan, but one or two very generalized species in those groups making it through and giving rise to many new species. The amazing thing is that over and over most of those new species are specialists, evolving to be increasingly good at dominating increasingly narrow sets of resources. Give Rattus rattus a few tens of millions of years and no other mammals on the planet, and they would evolve into many thousands of separate species, filling a vast array of niches, and most of those species would be specialists. If another great collapse came, the ones most likely to make it through would again be the super generalized rat.

As far as the "useless appendages" argument goes, remember that even when resources are abundant, there is still the race to see who can convert those resources into the most offspring the fastest. Plus, the ideal situation rarely lasts very long. Usually within a few generations the population of predators has increased, the food supply has diminished, or population density has gotten so high that pathogens are spread easily. Exponential growth is not to be underestimated. So with the possible exception of humans over the last couple of hundred years, it is almost never the case that a population goes on growing for many generations without selection knocking back those who spend their energy recklessly.

That said, there are traits that are advantageous in bad time and costly in good times, or the opposite. Sometimes species evolve plasticity, such as the ability to grow a thicker coat when the winter is colder, but not waste the protein in mild winters. And sometimes, like Darwin's finches, they just evolve back and forth. The camel's hump is probably something of a hindrance when water and food are plentiful, but it bears that cost because more often than not things will get dry again, and that hump will save its life. If camels lived in an environment where it didn't get dry for some thousands of years, they might end up sans hump, looking more like big llamas. Or they might just die out, vanquished by cows and goats. Camels, after all, are specialists.

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