Darwin and the Galápagos

Charles Darwin's historic visit to the Galápagos Islands in 1835 represents a landmark in the annals of science. But contrary to the legend long surrounding Darwin's famous Galápagos visit, he continued to believe that species were immutable for nearly a year and a half after leaving these islands. This delay in Darwin's evolutionary appreciation of the Galápagos evidence is largely owing to numerous misconceptions that he entertained about the islands, and their unique organic inhabitants, during the Beagle voyage. For example, Darwin mistakenly thought that the Galápagos tortoise–adult specimens of which he did not collect for scientific purposes–was not native to these islands. Hence he apparently interpreted reports of island-to-island differences among the tortoises as analogous to changes that are commonly undergone by species removed from their natural habitats. As for Darwin's finches, Darwin initially failed to recognize the closely related nature of the group, mistaking certain species for the forms that they appear, through adaptive radiation, to mimic. Moreover, what locality information he later published for his Galápagos finch specimens was derived almost entirely from the collections of three other Beagle shipmates, following his return to England. Even after he became an evolutionist, in March of 1837 (when he discussed his Galápagos birds with the eminent ornithologist John Gould), Darwin's theoretical understanding of evolution in the Galápagos continued to undergo significant developments for almost as many years as it took him to publish the Origin of Species (1859). The Darwin-Galápagos legend, with its romantic portrait of Darwin's 'eureka-like' insight into the Galápagos as a microcosmic 'laboratory of evolution', masks the complex nature of scientific discovery, and, thereby, the real nature of Darwin's genius.     

PREDICTING MICROEVOLUTIONARY RESPONSES TO DIRECTIONAL SELECTION ON HERITABLE VARIATION

Microevolution of quantitative traits in the wild can be predicted from a knowledge of selection and genetic parameters. Testing the predictions requires measurement of the offspring of the selected group, a requirement that is difficult to meet. We present the results of a study of Darwin's finches on the Galápagos island of Daphne Major where this requirement is met. The study demonstrates microevolutionary consequences of natural selection.     

Geology of Galápagos

This paper reviews the origins and geological history of the Galápagos Islands. The islands arose from a 'hot-spot'. The oceanic crust on which the islands are built can be no more than 10 million years old, and the islands themselves have been in existence at least 3.3 million years.
The Galápagos are among the most active volcanic groups in the world, and the physical nature of the islands is dominated by lava structures of various ages, including lava tubes or tunnels, which have been of particular interest to biologists. Weathering of the rock to produce soil has generally been slow, particularly in the drier parts.     

Low MHC variation in the endangered Galápagos penguin (<Emphasis Type="Italic">Spheniscus mendiculus</Emphasis>)

The major histocompatibility complex (MHC) is one of the most polymorphic regions of the genome, likely due to balancing selection acting to maintain alleles over time. Lack of MHC variability has been attributed to factors such as genetic drift in small populations and relaxed selection pressure. The Galápagos penguin (Spheniscus mendiculus), endemic to the Galápagos Islands, is the only penguin that occurs on the equator. It relies upon cold, nutrient-rich upwellings and experiences severe population declines when ocean temperatures rise during El Niño events. These bottlenecks, occurring in an already small population, have likely resulted in reduced genetic diversity in this species. In this study, we used MHC class II exon 2 sequence data from a DRB1-like gene to characterize the amount of genetic variation at the MHC in 30 Galápagos penguins, as well as one Magellanic penguin (S. magellanicus) and two king penguins (Aptenodytes patagonicus), and compared it to that in five other penguin species for which published data exist. We found that the Galápagos penguin had the lowest MHC diversity (as measured by number of polymorphic sites and average divergence among alleles) of the eight penguin species studied. A phylogenetic analysis showed that Galápagos penguin MHC sequences are most closely related to Humboldt penguin (Spheniscus humboldti) sequences, its putative sister species based on other loci. An excess of non-synonymous mutations and a pattern of trans-specific evolution in the neighbor-joining tree suggest that selection is acting on the penguin MHC.     

Inbreeding depression in the partially self-incompatible endemic plant species <Emphasis Type="Italic">Scalesia affinis</Emphasis> (Asteraceae) from Galápagos islands

A previous study showed that some individuals of the tetraploid Galápagos endemic Scalesia affinis were able to produce offspring after selfing. The present study compares the fitness of self-pollinated offspring with the fitness of cross-pollinated offspring. Germination success, seedling survival, and four different growth parameters was measured. In most of the studied characters selfed offspring were significantly inferior to outcrossed progeny. The effect was very clear in germination and survival. Outcrossed embryos were 3.4 times more likely to germinate than those that were selfed-fertilized, and the mortality was 84% higher among selfed individuals. Also, there was no genetic variation in inbreeding depression. The present study is based on material from a large population on Isabela Island, Galápagos. At other localities in the archipelago, populations have been through recent dramatic bottlenecks due to the grazing of introduced mammals. Considering the significant inbreeding depression found in the large population and the presence of a partial self-incompatibility system, these small populations are likely to be highly vulnerable and their future survival critically threatened.     

Characterisation of Rubus niveus: a prerequisite to its biological control in oceanic islands

This study is intended to inform the search for a biocontrol solution to stem the spread of Rubus niveus, a morphologically-diverse and invasive Rubus species which has become an aggressive invader in areas of the world outside its native range. The fragile biodiversity of islands can be especially threatened by this plant – exemplars of this are the Galápagos archipelago and Hawaii. This report focuses on the morphological and molecular work carried out to establish the homogeneity or intra-specific variation of R. niveus populations within the Galápagos Islands, as well as their kinship with plants of the same name found in Asia. The premise of this study is that genetic closeness between these populations offers increased probability that pathogens of R. niveus in its native range will also provide effective control in the target environment. Increased knowledge of the genetic make-up of R. niveus in both its native and naturalised locations will also help to assess the feasibility of using organisms which have been found to be effective against other Rubus species, but not hitherto considered for R. niveus itself.