Certhidea Research Articles

Evolutionary divergence of body size and wing and leg structure in relation to foraging mode in Darwin’s Galapagos finches

Abstract The wings, legs, and tail in Darwin’s finches show many clear adaptations to different types of locomotion used during foraging. We use size scaling to analyse how various characters vary with body mass to clarify dimensional relationships. The selective advantage of a character is judged in terms of energy savings. The wing aspect ratio (4.6–4.9) is very low, so the energy costs for flight are high. Low body mass, low wing loading, and short arm wings in the warbler finch, small tree finch, and small ground finch promote agility and manoeuvrability among vegetation, along with short wings in the warbler finch. Evolution towards a shorter arm wing seems to be favoured in the smaller finch species. Long legs, long toes, and long curved claws are adaptations for climbing/clinging locomotion without tail support (woodpecker finch, small and large tree finches, cactus finch but having short legs). Selection for longer legs seems to act towards a lengthening of the tarsometatarsus. The climbing technique in the woodpecker finch is described. We discuss how the diversification in the beaks relates to the locomotion organs.

Plant invasion causes alterations in Darwin's finch feeding patterns in Galápagos cloud forests

Invasive species pose a major threat to forest biodiversity, particularly on islands such as the Galapágos. Here, invasive plants are threatening the remnants of the unique cloud forest and its iconic Darwin's finches. We posit that food web disturbances caused by invasive Rubus niveus (blackberry), have contributed to the rapid decline of the insectivourous green warbler finch (Certhidae olivacea). We compared the birds' dietary changes in long-term management, short-term management and unmanaged areas. We measured C:N ratios, and δ15N‑nitrogen and δ13C‑carbon values in both consumer tissues (bird-blood) and food sources (arthropods), as indicators of resource use change, and collected mass abundance, and arthropod diversity data. We characterised the birds' diets using isotope mixing models. The results revealed that finches in (blackberry-invaded) unmanaged areas foraged more on abundant, yet lower quality, arthropods present in the invaded understory. This suggests that blackberry encroachment leads to a decrease in food source quality with physiological consequences for green warbler finch chicks. Results also implied that blackberry control has a short-term impact on food source quantity, which led to a decrease in chick recruitment that we observed in our previous studies; despite this, in the long-term, these managed systems show signs of recovery within three years of restoration.

More extinctions on the Galápagos Islands? An unsuccessful search for 4 landbirds on Floreana

Previous intensive surveys on Floreana, the smallest inhabited island of the Galápagos archipelago, in 2015 and 2016, produced reliable population estimates for all landbirds but failed to find 4 species that formerly occurred on the island: Gray Warbler Finch (Certhidea fusca), Vegetarian Finch (Platyspiza crassirostris), Little Vermilion Flycatcher (Pyrocephalus nanus), and Galápagos Rail (Laterallus spilonota). We revisited the island in February 2018 to extend our search to additional areas but again found none of the 4 lost species. As there is no longer any suitable habitat, we are certain that Galápagos Rail is now extinct on Floreana. It is possible that the remaining 3 species still survive in pockets of habitat that have not yet been surveyed. The very distinct subspecies of Gray Warbler-Finch (C. f. ridgwayi) especially merits further investigation.

Timing of infestation influences virulence and parasite success in a dynamic multi-host\u2013parasite interaction between the invasive parasite, Philornis downsi, and Darwin\u2019s finches

Recently commenced host–parasite interactions provide an excellent opportunity to study co-evolutionary processes. Multi-host systems are especially informative because variation in virulence between hosts and temporal changes provides insight into evolutionary dynamics. However, empirical data under natural conditions are scarce. In the present study, we investigated the interaction between Darwin’s finches and the invasive fly Philornis downsi whose larvae feed on the blood of nestlings. Recently, however, the fly has changed its behavior and now also attacks incubating females. Two sympatric hosts are affected differently by the parasite and parasite load has changed over time. Our study observed a reversal of trends described two decades ago: while, currently, small tree finches (Camarhynchus parvulus) experience significantly higher parasite load than warbler finches (Certhidea olivacea), this was the opposite two decades ago. Currently, fledging success is higher in warbler finches compared to small tree finches. Our data indicate that not only intensity but also timing of infestation influences hosts’ reproductive success and parasite fitness. During incubation, prevalence was higher in warbler finches, but once chicks had hatched, prevalence was 100% in both species and parasite load was higher in small tree finches. Furthermore, our results suggest faster development and higher reproductive success of P. downsi in small tree finch nests. A change in host preference driven by larvae competition could have led to the reversal in parasite load.

Weed management increases the detrimental effect of an invasive parasite on arboreal Darwin's finches

The detrimental effects of invasive parasites on hosts often increase under poor environmental conditions. Both natural fluctuations in environmental conditions and habitat management measures can temporarily cause adverse environmental effects. In this study, we investigated the interaction between the invasive parasitic fly Philornis downsi, control of invasive plants and precipitation on the breeding success of Darwin's finches. Introduced plant species have invaded a unique forest on the Galapagos island of Santa Cruz, which is a key habitat for Darwin's finches. The Galapagos National Park Directorate applies manual control and herbicides to combat this invasion. We hypothesized that these measures led to a reduction in the arthropod food supply during chick rearing, which in turn caused mortality in chicks that were already weakened by parasitism. We compared food availability in three study sites of varying degrees of weed management. To assess the interaction of parasitism and weed management, we experimentally reduced P. downsi abundance in nests in all three study sites. The key result suggests that there is an increase in the adverse effect of parasitism under unfavourable habitat conditions. As predicted, we found a negative additive effect of parasitism and weed management on the breeding success of the insectivorous warbler finch (Certhidea olivacea), but not on the omnivorous small tree finch (Camarhynchus parvulus), which was strongly affected by parasitism independently of weed management. Management measures can temporarily decrease environmental quality and reduce resistance/tolerance to invasive species. This should be considered when applying management measures to habitats of vulnerable species.

The impact of invasive plant management on the foraging ecology of the Warbler Finch (Certhidea olivacea) and the Small Tree Finch (Camarhynchus parvulus) on Gal\xe1pagos

In recent decades, arboreal Darwin’s Finches have suffered from a dramatic population decline, which has been attributed to parasitism by the invasive botfly Philornis downsi. However, changes to their primary habitat caused by invasive plant species may have additionally contributed to the observed population decline. The humid cloud forest on Santa Cruz Island is a stronghold of arboreal Darwin’s Finches but has been invaded by blackberry (Rubus niveus). In some areas, manual control and herbicide application are used to combat this invasion, both causing a temporary removal of the entire understory. We hypothesized that the removal of the understory reduces the availability of arthropods, which are a main food source during chick rearing. We compared the foraging behaviour of Warbler Finches (Certhidea olivacea) and Small Tree Finches (Camarhynchus parvulus) at three study sites that varied in the degree of R. niveus invasion and the length of time since the last herbicide application. We used prey attack rate and foraging success as an index for food availability and predicted a lower attack rate and foraging success in areas that had recently been sprayed with herbicides. We found that both the invasion and the management of R. niveus influenced microhabitat use, foraging substrate and prey choice in both species. Contrary to our hypothesis, we did not find a lower attack rate or foraging success in the area with recent herbicide application. This may be explained by the finding that both species mainly foraged in the canopy but also used dead plant structures of the understory of the recently controlled area that resulted from the invasive plant management.

Host-specific associations affect the microbiome of Philornis downsi, an introduced parasite to the Galápagos Islands.

The composition and diversity of bacteria forming the microbiome of parasitic organisms have implications for differential host pathogenicity and host-parasite co-evolutionary interactions. The microbiome of pathogens can therefore have consequences that are relevant for managing disease prevalence and impact on affected hosts. Here, we investigate the microbiome of an invasive parasitic fly Philornis downsi, recently introduced to the Galápagos Islands, where it poses extinction threat to Darwin's finches and other land birds. Larvae infest nests of Darwin's finches and consume blood and tissue of developing nestlings, and have severe mortality impacts. Using 16s rRNA sequencing data, we characterize the bacterial microbiota associated with P.downsi adults and larvae sourced from four finch host species, inhabiting two islands and representing two ecologically distinct groups. We show that larval and adult microbiomes are dominated by the phyla Proteobacteria and Firmicutes, which significantly differ between life stages in their distributions. Additionally, bacterial community structure significantly differed between larvae retrieved from strictly insectivorous warbler finches (Certhidea olivacea) and those parasitizing hosts with broader dietary preferences (ground and tree finches, Geospiza and Camarhynchus spp., respectively). Finally, we found no spatial effects on the larval microbiome, as larvae feeding on the same host (ground finches) harboured similar microbiomes across islands. Our results suggest that the microbiome of P.downsi changes during its development, according to dietary composition or nutritional needs, and is significantly affected by host-related factors during the larval stage. Unravelling the ecological significance of bacteria for this parasite will contribute to the development of novel, effective control strategies.

Invasive Parasites, Habitat Change and Heavy Rainfall Reduce Breeding Success in Darwin's Finches

Invasive alien parasites and pathogens are a growing threat to biodiversity worldwide, which can contribute to the extinction of endemic species. On the Galápagos Islands, the invasive parasitic fly Philornis downsi poses a major threat to the endemic avifauna. Here, we investigated the influence of this parasite on the breeding success of two Darwin's finch species, the warbler finch (Certhidea olivacea) and the sympatric small tree finch (Camarhynchus parvulus), on Santa Cruz Island in 2010 and 2012. While the population of the small tree finch appeared to be stable, the warbler finch has experienced a dramatic decline in population size on Santa Cruz Island since 1997. We aimed to identify whether warbler finches are particularly vulnerable during different stages of the breeding cycle. Contrary to our prediction, breeding success was lower in the small tree finch than in the warbler finch. In both species P. downsi had a strong negative impact on breeding success and our data suggest that heavy rain events also lowered the fledging success. On the one hand parents might be less efficient in compensating their chicks' energy loss due to parasitism as they might be less efficient in foraging on days of heavy rain. On the other hand, intense rainfalls might lead to increased humidity and more rapid cooling of the nests. In the case of the warbler finch we found that the control of invasive plant species with herbicides had a significant additive negative impact on the breeding success. It is very likely that the availability of insects (i.e. food abundance)is lower in such controlled areas, as herbicide usage led to the removal of the entire understory. Predation seems to be a minor factor in brood loss.

The evolutionary history of Darwin's finches: speciation, gene flow, and introgression in a fragmented landscape.

Many classic examples of adaptive radiations take place within fragmented systems such as islands or mountains, but the roles of mosaic landscapes and variable gene flow in facilitating species diversification is poorly understood. Here we combine phylogenetic and landscape genetic approaches to understand diversification in Darwin's finches, a model adaptive radiation. We combined sequence data from 14 nuclear introns, mitochondrial markers, and microsatellite variation from 51 populations of all 15 recognized species. Phylogenetic species-trees recovered seven major finch clades: ground, tree, vegetarian, Cocos Island, grey and green warbler finches, and a distinct clade of sharp-beaked ground finches (Geospiza cf. difficilis) basal to all ground and tree finches. The ground and tree finch clades lack species-level phylogenetic structure. Interisland gene flow and interspecies introgression vary geographically in predictable ways. First, several species exhibit concordant patterns of population divergence across the channel separating the Galápagos platform islands from the separate volcanic province of northern islands. Second, peripheral islands have more admixed populations while central islands maintain more distinct species boundaries. This landscape perspective highlights a likely role for isolation of peripheral populations in initial divergence, and demonstrates that peripheral populations may maintain genetic diversity through outbreeding during the initial stages of speciation.

A CENTURY OF GENETIC CHANGE AND METAPOPULATION DYNAMICS IN THE GALÁPAGOS WARBLER FINCHES (CERTHIDEA)

Populations that are connected by immigrants play an important role in evolutionary and conservation biology, yet we have little direct evidence of how such metapopulations change genetically over evolutionary time. We compared historic (1894-1906) to modern (1988-2006) genetic variation in 11 populations of warbler finches at 14 microsatellite loci. Although several lines of evidence suggest that Darwin's finches may be in decline, we found that the genetic diversity of warbler finches has not generally declined, and broad-scale patterns of variation remained similar over time. Contrary to expectations, inferred population sizes have generally increased over time (6-8%) as have immigration rates (8-16%), which may reflect a recent increase in the frequency and intensity of El Niño events. Individual island populations showed significant declines (18-19%) and also substantial gains (18-20%) in allelic richness over time. Changes in genetic diversity were correlated with changes in immigration rates, but did not correspond to population size or human disturbance. These results reflect the expected stabilizing properties of whole metapopulations over time. However, the dramatic and unpredictable changes observed in individual populations during this short time interval suggests that care should be taken when monitoring individual population fragments with snapshots of genetic variation.

Scaling and shear transformations capture beak shape variation in Darwin's finches.

Evolution by natural selection has resulted in a remarkable diversity of organism morphologies that has long fascinated scientists and served to establish the first relations among species. Despite the essential role of morphology as a phenotype of species, there is not yet a formal, mathematical scheme to quantify morphological phenotype and relate it to both the genotype and the underlying developmental genetics. Herein we demonstrate that the morphological diversity in the beaks of Darwin's Finches is quantitatively accounted for by the mathematical group of affine transformations. Specifically, we show that all beak shapes of Ground Finches (genus Geospiza) are related by scaling transformations (a subgroup of the affine group), and the same relationship holds true for all the beak shapes of Tree, Cocos, and Warbler Finches (three distinct genera). This analysis shows that the beak shapes within each of these groups differ only by their scales, such as length and depth, which are genetically controlled by Bmp4 and Calmodulin. By measuring Bmp4 expression in the beak primordia of the species in the genus Geospiza, we provide a quantitative map between beak morphology and the expression levels of Bmp4. The complete morphological variation within the beaks of Darwin's finches can be explained by extending the scaling transformations to the entire affine group, by including shear transformations. Altogether our results suggest that the mathematical theory of groups can help decode morphological variation, and points to a potentially hierarchical structure of morphological diversity and the underlying developmental processes.

Darwin's Finches as Seed Predators and Dispersers

We investigated seed predation and dispersal by nine Darwin's finch (Geospizinae) species, including species previously regarded as seed predators or insectivores. All nine ate fruit. Eight commonly discarded seeds, acting as short-distance dispersers. Seven defecated viable seeds, being therefore potential longer-distance dispersers. Fruit comprised the majority of dietary items in all four Geospiza species and Small Tree Finch (Camarhynchus parvulus). All except Common Cactus Finch (G. scandens) defecated intact seeds. The highest proportions of feces containing viable seeds were of Small Ground Finch (G. fuliginosa) and the “insectivorous” species Woodpecker Finch (Camarhynchus pallidus) and Warbler Finch (Certhidea olivacea). These two may be more important endochorous dispersers than other species that eat more fruit but are better seed predators. Intact seeds were found in 23% of fecal samples; 50% of the samples with intact seeds had viable seeds. The most frequently encountered intact seeds were of the tiny-seeded, fleshy-fruited Miconia robinsoniana and a sedge, Kyllinga brevifolia. Dry fruit, such as sedges and grasses, were usually crushed but occasionally found intact in feces, and might frequently be dispersed long distances. Seeds of larger-seeded, fleshy-fruited species were often discarded, but may be commonly dispersed short distances. Dispersal of plants by finches is probably more common than suggested by their predominant feeding patterns of seed predation and insectivory, and rare long-distance dispersal could be important for plant evolution and spread of introduced plants in the Galapagos Islands.

The calmodulin pathway and evolution of elongated beak morphology in Darwin's finches

A classic textbook example of adaptive radiation under natural selection is the evolution of 14 closely related species of Darwin's finches (Fringillidae, Passeriformes), whose primary diversity lies in the size and shape of their beaks. Thus, ground finches have deep and wide beaks, cactus finches have long and pointed beaks (low depth and narrower width), and warbler finches have slender and pointed beaks, reflecting differences in their respective diets. Previous work has shown that even small differences in any of the three major dimensions (depth, width and length) of the beak have major consequences for the overall fitness of the birds. Recently we used a candidate gene approach to explain one pathway involved in Darwin's finch beak morphogenesis. However, this type of analysis is limited to molecules with a known association with craniofacial and/or skeletogenic development. Here we use a less constrained, complementary DNA microarray analysis of the transcripts expressed in the beak primordia to find previously unknown genes and pathways whose expression correlates with specific beak morphologies. We show that calmodulin (CaM), a molecule involved in mediating Ca2+ signalling, is expressed at higher levels in the long and pointed beaks of cactus finches than in more robust beak types of other species. We validated this observation with in situ hybridizations. When this upregulation of the CaM-dependent pathway is artificially replicated in the chick frontonasal prominence, it causes an elongation of the upper beak, recapitulating the beak morphology of the cactus finches. Our results indicate that local upregulation of the CaM-dependent pathway is likely to have been a component of the evolution of Darwin's finch species with elongated beak morphology and provide a mechanistic explanation for the independence of beak evolution along different axes. More generally, our results implicate the CaM-dependent pathway in the developmental regulation of craniofacial skeletal structures.

Comparative landscape genetics and the adaptive radiation of Darwin's finches: the role of peripheral isolation

We use genetic divergence at 16 microsatellite loci to investigate how geographical features of the Galápagos landscape structure island populations of Darwin's finches. We compare the three most genetically divergent groups of Darwin's finches comprising morphologically and ecologically similar allopatric populations: the cactus finches (Geospiza scandens and Geospiza conirostris), the sharp-beaked ground finches (Geospiza difficilis) and the warbler finches (Certhidea olivacea and Certhidea fusca). Evidence of reduced genetic diversity due to drift was limited to warbler finches on small, peripheral islands. Evidence of low levels of recent interisland migration was widespread throughout all three groups. The hypothesis of distance-limited dispersal received the strongest support in cactus and sharp-beaked ground finches as evidenced by patterns of isolation by distance, while warbler finches showed a weaker relationship. Support for the hypothesis that gene flow constrains morphological divergence was only found in one of eight comparisons within these groups. Among warbler finches, genetic divergence was relatively high while phenotypic divergence was low, implicating stabilizing selection rather than constraint due to gene flow. We conclude that the adaptive radiation of Darwin's finches has occurred in the presence of ongoing but low levels of gene flow caused by distance-dependent interisland dispersal. Gene flow does not constrain phenotypic divergence, but may augment genetic variation and facilitate evolution due to natural selection. Both microsatellites and mtDNA agree in that subsets of peripheral populations of two older groups are genetically more similar to other species that underwent dramatic morphological change. The apparent decoupling of morphological and molecular evolution may be accounted for by a modification of Lack's two-stage model of speciation: relative ecological stasis in allopatry followed by secondary contact, ecological interactions and asymmetric phenotypic divergence.

Habitat selection and ecological speciation in Galápagos warbler finches ( Certhidea olivacea and Certhidea fusca )

We investigated phylogeographic divergence among populations of Galápagos warble finches. Their broad distribution, lack of phenotypic differentiation and low levels of genetic divergence make warbler finches an appropriate model to study speciation in allopatry. A positive relationship between genetic and geographical distance is expected for island taxa. Warbler finches actually showed a negative isolation by distance relationship, causing us to reject the hypothesis of distance-limited dispersal. An alternative hypothesis, that dispersal is limited by habitat similarity, was supported. We found a positive correlation between genetic distances and differences in maximum elevation among islands, which is an indicator of ecological similarity. MtDNA sequence variation revealed monophyletic support for two distinct species. Certhidea olivacea have recently dispersed among larger central islands, while some Certhidea fusca have recently dispersed to small islands at opposite ends of the archipelago. We conclude that females have chosen to breed on islands with habitats similar to their natal environment. Habitat selection is implicated as an important component of speciation of warbler finches, which is the earliest known divergence of the adaptive radiation of Darwin's finches. These results suggest that small populations can harbour cryptic but biologically meaningful variation that may affect longer term evolutionary processes.

Extinction behind our backs: the possible fate of one of the Darwin’s finch species on Isla Floreana, Galápagos

Without regular monitoring a rare species may slip into extinction unnoticed. We report a possible case from the Galápagos archipelago. The warbler finch ( Certhidea fusca) has not been recorded by scientists on Isla Floreana in recent years, and we have been concerned about its status. On a two-day visit to prime habitat in the breeding season of 2004 we used playback of warbler finch song and calls recorded on another island to stimulate an approach of local birds. We failed to find a single warbler finch, whereas we encountered numerous small tree finches ( Camarhynchus parvulus), medium tree finches ( C. pauper) and yellow warblers ( Dendroica petechia). Searches in 1979, 1983, 1997 and 1999 were also unsuccessful. Our continuing failure to find any warbler finches indicates the species must be extremely rare on the island, if not actually extinct. We discuss possible reasons for their demise and draw attention to other species that could be heading in the same direction.

Lack of Premating Isolation at the Base of a Phylogenetic Tree

Darwin's finches in the Galápagos archipelago are an unusual example of adaptive radiation in that the basal split separates two lineages of warbler finches (Certhidea olivacea and Certhidea fusca) believed until recently to be only one species. The large genetic difference between them contrasts with their similarity in plumage, size, shape, and courtship behavior. They differ in song, which is a key factor in premating isolation of other sympatric Darwin's finches. We conducted playback experiments to see whether members of the population of C. olivacea on Santa Cruz Island would respond to songs of C. fusca from two islands, Genovesa and Pinta, and songs of C. olivacea from another island (Isabela). Another set of experiments was performed, using the same playback tapes, with C. fusca on Genovesa. Some members of both populations responded to all playbacks; therefore, the hypothesis of complete premating isolation on the basis of song is rejected. Discrimination between songs of the two lineages was inconsistent. We conclude that premating barriers to interbreeding among the tested populations have not arisen in the 1.5-2.0 m.yr. of their geographical isolation on different islands. This contrasts with strong premating barriers between more recently derived sympatric species. Early learning of song associated with morphology is later used in mate recognition. This explains why sympatric species that are vocally and morphologically distinct yet genetically less differentiated than Certhidea do not interbreed, whereas the Certhidea lineages that are genetically well differentiated but vocally and morphologically similar have no apparent premating barrier. We discuss this unusual situation in terms of the forces that have produced similarities and differences in song, morphology, and ecology and their relevance to phylogenetic and biological species concepts. Neither principles nor details are unique to Darwin's finches, and we conclude by pointing out strong parallels with some continental birds.

A New Perspective on The Evolutionary History of Darwin's Finches

The 13 species of finches found on the Galapagos Islands, and the one species from Cocos Island, together collectively known as Darwin's finches, are one of the better-studied groups of birds. Much has been learned about their ecology, behavior, and the short-term effects of natural selection (Lack 1947, Bowman 1961, Grant 1999). The four species of tree finch (Camarhynchus), six species of ground finch (Geospiza), the tool-using Woodpecker Finch (Cactospiza pallida), the Vegetarian Finch (Platyspiza crassirostris), the Warbler Finch (Certhidea olivacea), and the finch on Cocos Island (Pinaroloxias inornata) appear to fill different ecological roles via differentiation of bill size and shape (Grant and Grant 2002). In fact, most authors consider Darwin's finches to be a classic example of an adaptive radiation, owing to the great diversity in bill form and ecological habit that presumably evolved in a relatively short time. For example, Petren et al. (1999:321) noted, Species in this group show adaptive variation in beak size, beak shape and body size that is more typical of differences among [taxonomic] families of birds... (boldface added). Taxonomic History.-Darwin originally collected the finches from the Galapagos, not realizing what an evolutionary gold mine they would become (Sulloway 1982). In fact, he was not careful about labeling specimens as to the island from which a specimen was obtained, thus obscuring taxonomic boundaries. Subsequent work with better-labeled specimens revealed taxonomically significant patterns of variation. Gould (1837) recognized that different species existed in the collections from the Darwin expedition. However, Gould was unable to produce a stable classification. Since Gould's time, many of the major figures in avian taxonomy have published differing classifications. The succession of revisionary efforts attests to the difficult nature of classifying phenotypic variation among the finches. Although the species show considerable morphological diversity, many in fact are notoriously difficult to identify in the field (especially females and immatures). Populations from different species and islands overlap in morphometric space (Grant 1981).

Phylogenetics of Darwin's Finches: Paraphyly in the Tree-Finches, and Two Divergent Lineages in the Warbler Finch

The Galapagos Darwin's finches (Geospizinae) have been classified as three major groups based on morphology and behavior: ground-finches, tree-finches, and the Warbler Finch (Certhidea olivacea). Little is known about the evolutionary relationships within and among these groups, which is partly due to the lack of a phylogeny based on molecular sequence data. We used mitochondrial sequence data to reconstruct a phylogeny of Darwin's finches. These data show that within the tree-finches, only one genus is conclusively monophyletic, and another is conclusively paraphyletic. It may be appropriate to uphold the classification of the tree-finches into two genera. The Warbler Finch complex is paraphyletic, as revealed by two divergent genetic lineages contained within this species. Stochastic lineage sorting within relatively recently diverged species and interspecific and intergeneric hybridization are the two most likely explanations for the sharing of haplotypes among taxa.

Phylogeny of Darwin's finches as revealed by mtDNA sequences.

Darwin's finches comprise a group of passerine birds first collected by Charles Darwin during his visit to the Galápagos Archipelago. The group, a textbook example of adaptive radiation (the diversification of a founding population into an array of species differentially adapted to diverse environmental niches), encompasses 14 currently recognized species, of which 13 live on the Galápagos Islands and one on the Cocos Island in the Pacific Ocean. Although Darwin's finches have been studied extensively by morphologists, ecologists, and ethologists, their phylogenetic relationships remain uncertain. Here, sequences of two mtDNA segments, the cytochrome b and the control region, have been used to infer the evolutionary history of the group. The data reveal the Darwin's finches to be a monophyletic group with the warbler finch being the species closest to the founding stock, followed by the vegetarian finch, and then by two sister groups, the ground and the tree finches. The Cocos finch is related to the tree finches of the Galápagos Islands. The traditional classification of ground finches into six species and tree finches into five species is not reflected in the molecular data. In these two groups, ancestral polymorphisms have not, as yet, been sorted out among the cross-hybridizing species.