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.

Can't live with'em can't live without 'em.

Biologists and Ecologists have (mostly) learned to give at least some forethought to the consequences of introducing non-native species. The problems invasive species cause are often most sever on islands, where the native species often have limited evolutionary experience dealing with the relative of the invaders. For example, many island endemic bird species were flightless, as there were no land based predators to need to fly away from, so why bother building all that expensive escape equipment when there is nobody to escape from. Other island nesting birds, like those on Macquarie Island can fly (except the penguins) but don't seem to have the right tricks in their behavioral repertoires to escape from introduced predators.

A recent attempt to help these birds is making ecologists aware that one has to be very careful not only about introducing invasive species, but also about removing them. A commentary in Nature describes the story:

Cats were introduced to the Macquarie Island in 1818; sealers introduced rabbits 60 years later.

The rabbits tore through the island's vegetation. In 1968, the rabbit flea was introduced. Once that had established the lethal myxomatosis virus — which the flea spreads — was introduced in 1978.

Rabbit numbers crashed, but then the cats, which had previously eaten rabbits, switched their attentions to the island's birds.

But once the cats were gone, the few hardy rabbits that had survived both the cats and the myxomatosis emerged and began doing what rabbits do best — breeding and eating.

So the island has cats, rabbits, rabbit fleas, rabbit viruses as well as rats and mice, all introduced. The cats were bad for the birds because they ate them. The rabbits were introduced and were bad for the birds because they destroyed the vegetation, but good in that they distracted the cats. And the cats were at least partly good for the birds, because they ate the rabbits. Some kind of unstable plateau was reached. And then disease was introduced to reduce the rabbit population, leaving a whole bunch of hungry cats, which were bad for the birds. But then the cats were killed off, which was bad for the birds and everything else (except the rabbits, rats and mice) because for the first time there were rabbits without cats on the island. The rabbits, unchecked, ate through most of the island's plants. And this, of course, is a vast over simplification.

The Australian authorities who run Macquarie Island are now trying to figure out how to exterminate the rabbits, rats and mice without harming other native populations, such as the seals and sea lions that breed on the island. They estimate it will take tens of millions of dollars, and that may be optimistic. Few efforts to irradicate rodents from any land mass of decent size have succeeded. Macquarie Island, at 128 km², offers a lot of hiding places big enough for a rat. Miss one pregnant rat and in just few years you are back to square one. And who knows what effect the rats would have without rabbits around harboring disease and eating potential cover? Rats eat bird eggs, and have been known to finish off large bird colonies in just a few years.

The lesson learned is that just because introducing species is generally bad, removing those introduced species from an ecosystem that has started to adjust to their input isn't always good. At least it has to be done very carefuly.