[Al Uy asked to me to write a pseudo-popular piece about the adaptive radiation of Darwin's finches for a forthcoming "incipientspecies.org" website. I decided to use this blog to present a first (and as yet overlong and too technical) draft of my contribution.]
An image of the Galápagos Islands and their topography. Image from Wikimedia Commons.
The Galapagos Islands were formed by magma welling up from a hotspot on the ocean floor to build underwater mountains, some of which broke the surface and became islands. When the first island formed, more than 8 million years ago, it had no terrestrial life given its 900+ km separation from the mainland. With time, however, various plants and animals either flew, drifted, or were carried to the islands. One of those colonists, arriving approximately 1.5 million years ago, was a bird – presumably a flock of them – that probably looked something like a modern-day grassquit.
The Darwin’s finch ancestor may have looked this Black-faced Grassquit (Tiaris bicolor). Photo by C.J. Sharp on Wikimedia Commons.
The colonizing finches increased in abundance and spread across the various Galapagos islands. As they did so, they encountered different conditions. Some islands were very low and dry. Some islands were high and wet. Some had many insects, some had few. Some had certain types of plants, some had other types. Each of these different sets of conditions meant that a different way of feeding would be optimal at different locations. These different ways of feeding had a classic evolutionary target – the beak of the finch.
The diversity of pliers. Images from http://coilku.blogspot.ca/
The diversity of bird beaks and their functions. Image from L. Shyamal on Wikimedia
One representation of the Darwin's finch radiation (Grant 1986: Ecology and Evolution of Darwin's finches)
Crude depiction of the distribution of different finch species on the different islands. From A Field Guide to the Birds of Galapagos by Michael Harris. Collins.
First, the previously described situation in which populations in different places show adaptation to different food types means that species coming into secondary contact have already specialized on somewhat different food types, thus reducing competition. This initial divergence can increase following secondary contact due to selection against individuals that have traits/diets/behaviors most similar to the other species, and that therefore experience the highest competition. The resulting process of “character displacement” will then further reduce competition and promote species coexistence.
David Lack's classic demonstration of character displacement. Image from Ricklefs' (1996) Economy of Nature.
One of my stickleback experiment field sites on Vancouver Island.
1. Darwin’s finches are “imperfect generalists”
As described earlier, a critical mechanism by which two young Darwin’s finch species coexist when they come into secondary contact is through adaptation to different resources. The assembly of a community of finches thus depends critically on the extent to which different species partition their resources. One of our first goals was to understand how this partitioning took place, so – starting in 2003 – we began what would become a long program of simply walking around our field sites, finding birds, identifying them (through binoculars) to species, and determining on what they were feeding. This task has been greatly facilitated by the fact that Darwin’s finches are very tame. In the early stages of this work, we were quite surprised to see that, contrary to our initial naïve expectations, most of the species seemed to be feeding on pretty much the same things. Where was this niche partitioning that was supposedly so critical to the adaptive radiation?
Those first years were very wet, with lots of plant reproduction, lots of seeds, and lots of insects. But then a major drought occurred and, for several years, plant reproduction was minimal and so seed and insect abundances declined dramatically. Fortunately, we had continued to record what the finches were eating throughout this period. During these drought years, we found that the different species increasingly diverged to use different resources – and niche overlap decreased accordingly. Thus, with 5 years of feeding observation data spanning wet and dry years, we were able to conclude: These results together suggest that the ground finches are ‘imperfect generalists’ that use overlapping resources under benign conditions (in space or time), but then retreat to resources for which they are best adapted during periods of food limitation. These conditions likely promote local and regional coexistence (De Leon et al. 2014). This finding that niche overlap decreased in years when little rain fell fit well with earlier observations that niche overlap decreased during the dry (as opposed to wet) seasons within a year.
2. The adaptive radiation is ongoing
During that early walk in 2002 with Peter and Rosemary, I asked them what they thought would be one of the most interesting questions to investigate on Santa Cruz, where we were planning to work. They suggested trying to understand the causes and consequences of the hyper-variable population of medium ground finches, Geospiza fortis. It turns out that this species is more variable on Santa Cruz than anywhere else – indeed, they are so variable that a paper by Hugh Ford in 1973 argued they were undergoing sympatric speciation. Just the next year, team member Anthony Herrel came into our dorm room to show us some data from the birds we had captured that year. The histogram of beak sizes was bimodal – just like Hugh Ford had reported. This result was inspiring because it suggested that the population might be in the midst of splitting into separate species, a rare event that would enable us to formally test the mechanisms thought to promote the adaptive radiation of Darwin’s finches. (It is difficult to test such mechanisms when species are already well established.)
Clockwise from top left: The large and small beak morphs, the demonstration of bimodality, assortative mating, and disruptive selection
3. Human influences on adaptive radiation
Given a population of finches seemingly in the midst of splitting into separate species, none of the above results were surprising – yet we did have a surprise coming. The bimodal population of finches described above was not the same population previously described by Hugh Ford. For the latter population, found at “Academy Bay” immediately adjacent to the main tourist town of Puerto Ayorra, we couldn’t find evidence for bimodality in any of our new samples. Even though the population was still quite variable, it just didn’t show the dip in the frequency distribution of beak size that Ford had reported and that we were finding at El Garrapatero, just 7 km away. It seemed that Ford’s population had lost its bimodality between then and now, and we became curious as to just when that collapse had occurred. At this point, we rounded up all of the previous data available for G. fortis from Santa Cruz. Peter and Rosemary and their collaborators provided much of it, including data from David Snow’s collections in 1963–1964. The most fun for me, however, was to find Hugh Ford’s contact information online and send him an email:
Another critical frontier is to examine how finches influence the evolution of plant traits and assembly of plant communities – work we are now starting with Marc Johnson, Nancy Emery, and Sofia Carvajal. Much work remains to be done, and I am curious to find out just how Darwin’s finches will continue reshaping my life and career.