Historical fragmentation of islands and genetic drift in populations of Galápagos lava lizards (Microlophus albemarlensis complex).

The formation of islands following a rise in sea level at the end of Pleistocene is expected to disrupt the equilibrium between genetic drift and gene flow in species with limited ability to disperse. Here, we test the hypothesis that genetic drift in isolation has caused the differentiation of Galápagos lava lizards (Microlophus albemarlensis complex) found on 12 islets that are likely to have been connected to a larger island, Isla Santa Cruz, during the late Pleistocene. Using 11 microsatellite loci, screened on 524 individuals from 17 localities distributed among and within 15 islands, we found marked differences in allelic richness and heterozygosity. Genetic differentiation was strong (global F(ST) = 0.44), with pairwise differences found among populations on islets being larger than differences among three localities sampled within Isla Santa Cruz. As expected under a scenario of drift in isolation, there was a positive correlation of genetic diversity with island size, no relationship between genetic and geographical distance and a strong negative correlation between heterozygosity and measures of genetic differentiation. We conclude that seawater is a significant barrier to gene flow in lava lizards on this timescale. Our results suggest that the shallow diversification of the M. albemarlensis complex is not due to recent gene flow and that genetic drift may have played a substantial role in observed patterns of phenotypic variation among islands.

Phenotypic divergence despite high levels of gene flow in Galápagos lava lizards (Microlophus albemarlensis).

The extent of evolutionary divergence of phenotypes between habitats is predominantly the result of the balance of differential natural selection and gene flow. Lava lizards (Microlophus albemarlensis) on the small island of Plaza Sur in the Galápagos archipelago inhabit contrasting habitats: dense vegetation on the western end of the island thins rapidly in a transitional area, before becoming absent on the eastern half. Associated with these habitats are phenotypic differences in traits linked to predator avoidance (increased wariness, sprint speed, and endurance in lizards from the sparsely vegetated habitat). This population provides an opportunity to test the hypothesis that reduced gene flow is necessary for phenotypic differentiation. There was no evidence of any differences among habitats in allele frequencies at six out of seven microsatellite loci examined, nor was there any indication of congruence between patterns of genetic variability and the change in vegetation regime. We infer that gene flow between the habitats on Plaza Sur must be sufficiently high to overcome genetic drift within habitats but that it does not preclude phenotypic differentiation.

Character congruence and phylogenetic signal in molecular and morphological data sets: a case study in the living Iguanas (Squamata, Iguanidae).

The lizard family Iguanidae comprises eight living genera distributed throughout the New and Old World, and includes several island endemics. We reconstruct phylogenetic relationships among these genera using 90 previously published morphological characters, to which we add a molecular (mtDNA sequence) data set that includes 742 nucleotides of the ND4 gene and the complete sequences of the histidine, serine, and leucine tRNAs (217 nucleotides). Trees were initially constructed separately from these three data sets, and then tested for significant conflict in topologies that would suggest the influence of different evolutionary processes. The three data sets were then combined, and a single tree was obtained from the total evidence that permitted identification of potential sources of character incongruence. Several additional analyses of the combined data sets were repeated with sequential deletion of successive classes of homoplastic characters, and we show that the same single tree topology is recovered in most cases. However, part of the tree structure collapses when the matrix of combined characters is completely purged of all homoplastic characters. We argue that this extreme results in an unacceptable loss of phylogenetic information, and we present a single phylogenetic hypothesis for all living genera of iguanas. We show that this hypothesis is significantly more parsimonious than either of two previously published trees, and we discuss the evolution and biogeography of the Iguanidae based on the preferred hypothesis.

A survey of the coccidian parasites of reptiles from islands of the Galápagos Archipelago: 1990-1994.

From 1990 through 1994, fecal samples were collected and examined for coccidian parasites from 26 giant land tortoises Geochelone nigra, from 715 lava lizards Tropidurus spp., from 139 land iguanas Conolophus subcristatus, and from 128 marine iguanas Amblyrhynchus cristatus, all of which inhabit various islands in the Galápagos Archipelago. None of the samples from A. cristatus or from C. subcristatus was infected with coccidia. Only 1 of 26 (4%) G. nigra was infected with a single Eimeria species that we describe here as new. A total of 262 of 715 (37%) individuals representing 3 species of Tropidurus discharged oocysts of 1-3 different coccidian species; these included 2 previously described species Eimeria tropidura and Isospora insularius, and an eimerian that we describe here as new. Additionally, 104 fecal samples from Tropidurus spp. were from 51 animals recaptured in either 2 or 3 yr; 21 had no infections in any year, 15 were infected at least once, 14 were infected in 2 yr, and only 1 was infected during 3 yr. No animal was recaptured and sampled during each of the 4 yr of this study. Of the 262 infected individuals, 30 (12%) had multiple coccidial infections at the time of collection (eimerian and isosporan, or 2 eimerians). Where determination of the sexes was possible in the lava lizards, there was no difference in prevalence rates between males (39%) and females (41%). Sporulated oocysts of the new eimerian from Tropidurus are ellipsoidal, 27.1 x 15.6 (25-31 x 14-18) microns, with a polar body, but without a micropyle or oocyst residuum; they contain ellipsoidal sporocysts, 11.8 x 6.7 (10-14 x 6-8) microns, without Stieda, sub-, or parastieda bodies, but with a sporocyst residuum. Sporulated oocysts of the new eimerian from G. nigra are ellipsoidal to ovoidal, 21.6 x 18.1 (18-25 x 16-20) microns, with a large polar body, but without a micropyle or oocyst residuum; they contain ellipsoidal sporocysts 10.7 x 7.0 (8-12 x 5-8) microns, with Stieda body but no sub- or parastieda bodies. Also present is a sporocyst residuum of medium to large granules randomly distributed among the sporocysts.