Darwin's Finches Research Articles

Ecological and Phenotypic Diversification after a Continental Invasion in Neotropical Freshwater Stingrays.

Habitat transitions are key potential explanations for why some lineages have diversified and others have not-from Anolis lizards to Darwin's finches. The ecological ramifications of marine-to-freshwater transitions for fishes suggest evolutionary contingency: some lineages maintain their ancestral niches in novel habitats (niche conservatism), whereas others alter their ecological role. However, few studies have considered phenotypic, ecological, and lineage diversification concurrently to explore this issue. Here, we investigated the macroevolutionary history of the taxonomically and ecologically diverse Neotropical freshwater river rays (subfamily Potamotrygoninae), which invaded and diversified in the Amazon and other South American rivers during the late Oligocene to early Miocene. We generated a time-calibrated, multi-gene phylogeny for Potamotrygoninae and reconstructed evolutionary patterns of diet specialization. We measured functional morphological traits relevant for feeding and used comparative phylogenetic methods to examine how feeding morphology diversified over time. Potamotrygonine trophic and phenotypic diversity are evenly partitioned (non-overlapping) among internal clades for most of their history, until 20-16 mya, when more recent diversification suggests increasing overlap among phenotypes. Specialized piscivores (Heliotrygon and Paratrygon) evolved early in the history of freshwater stingrays, while later trophic specialization (molluscivory, insectivory, and crustacivory) evolved in the genus Potamotrygon. Potamotrygonins demonstrate ecological niche lability in diets and feeding apparatus; however, diversification has mostly been a gradual process through time. We suggest that competition is unlikely to have limited the potamotrygonine invasion and diversification in South America.

Where did the finch go? Insights from radio telemetry of the medium ground finch (Geospiza fortis).

Movement patterns and habitat selection of animals have important implications for ecology and evolution. Darwin's finches are a classic model system for ecological and evolutionary studies, yet their spatial ecology remains poorly studied. We tagged and radio‐tracked five (three females, two males) medium ground finches (Geospiza fortis) to examine the feasibility of telemetry for understanding their movement and habitat use. Based on 143 locations collected during a 3‐week period, we analyzed for the first time home‐range size and habitat selection patterns of finches at El Garrapatero, an arid coastal ecosystem on Santa Cruz Island (Galápagos). The average 95% home range and 50% core area for G. fortis in the breeding season was 20.54 ha ± 4.04 ha SE and 4.03 ha ± 1.11 ha SE, respectively. For most of the finches, their home range covered a diverse set of habitats. Three finches positively selected the dry‐forest habitat, while the other habitats seemed to be either negatively selected or simply neglected by the finches. In addition, we noted a communal roosting behavior in an area close to the ocean, where the vegetation is greener and denser than the more inland dry‐forest vegetation. We show that telemetry on Darwin's finches provides valuable data to understand the movement ecology of the species. Based on our results, we propose a series of questions about the ecology and evolution of Darwin's finches that can be addressed using telemetry.

Tetracycline Resistance Profile in Darwin Finches in the Galapagos Islands

Antimicrobial Resistance (AMR), which is the ability of microorganisms to withstand attack by antimicrobial drugs, has a worldwide impact. Thus, it is important to identify resistance mechanisms circulating in the environment. Among antimicrobials widely used in human and veterinary health, tetracycline is ideal due to its safety and has been categorized by the WHO as a critically important antimicrobial. Based on the fact that wild animals are bioindicators of environmental contamination and that wild birds are considered sentinels of AMR in the ecosystem, in the present study, we studied AMR patterns in Darwin finches (Geospiza spp.) in the Galapagos Islands, Ecuador. To this end, a total of 384 cloacal swabs from Darwin finches were collected from three zones of Santa Cruz Island (an urban, an agricultural, and a protected zone). Phenotypic antibiotic resistance was analyzed by the Kirby-Bauer disk diffusion method in Escherichia coli (n = 136) and Enterococcus spp. (n = 332) isolates. PCR was performed for the detection of the tetA gene in E. coli strains and of the tetM gene in Enterococcus isolates. Resistance for at least one of nine and one of eight antimicrobials was observed in 62.5% (85/136) and 28.6% (95/332) of E. coli and Enterococcus isolates, respectively. A high percentage of phenotypic resistance to tetracycline was identified for both bacteria. The tetA and tetM genes were identified in 83.7% (36/43) of E. coli strains and 75.4% (49/65) of Enterococcus spp. isolates, respectively. The highest percentages of AMR were observed in the agricultural zone. We also found the presence of multi-drug-resistant strains. These results show that Darwin finches might be proposed as sentinels of AMR in the Galapagos Islands.

The Role of Photobionts as Drivers of Diversification in an Island Radiation of Lichen-Forming Fungi

Speciation in oceanic islands has attracted the interest of scientists since the 19th century. One of the most striking evolutionary phenomena that can be studied in islands is adaptive radiation, that is, when a lineage gives rise to different species by means of ecological speciation. Some of the best-known examples of adaptive radiation are charismatic organisms like the Darwin finches of the Galapagos and the cichlid fishes of the great African lakes. In these and many other examples, a segregation of the trophic niche has been shown to be an important diversification driver. Radiations are known in other groups of organisms, such as lichen-forming fungi. However, very few studies have investigated their adaptive nature, and none have focused on the trophic niche. In this study, we explore the role of the trophic niche in a putative radiation of endemic species from the Macaronesian Region, the Ramalina decipiens group. The photobiont diversity was studied by Illumina MiSeq sequencing of the ITS2 region of 197 specimens spanning the phylogenetic breadth and geographic range of the group. A total of 66 amplicon sequence variants belonging to the four main clades of the algal genus Trebouxia were found. Approximately half of the examined thalli showed algal coexistence, but in most of them, a single main photobiont amounted to more than 90% of the reads. However, there were no significant differences in photobiont identity and in the abundance of ITS2 reads across the species of the group. We conclude that a segregation of the trophic niche has not occurred in the R. decipiens radiation.

Darwin's small and medium ground finches might have taste preferences, but not for human foods.

Urbanization is rapidly changing ecological niches. On the inhabited Galapagos Islands, Darwin's finches consume human-introduced foods preferentially; however, it remains unclear why. Here, we presented pastry with flavour profiles typical of human foods (oily, salty and sweet) to small ground finches (Geospiza fuliginosa) and medium ground finches (Geospiza fortis) to test if latent taste preferences might drive the selection of human foods. If human food flavours were consumed more than a neutral or bitter control only at sites with human foods, then we predicted tastes were acquired after urbanization; however, if no site differences were found then this would indicate latent taste preferences. Contrary to both predictions, we found little evidence that human food flavours were preferred compared with control flavours at any site. Instead, finches showed a weak aversion to oily foods, but only at remote (no human foods present) sites. This was further supported by behavioural responses, with beak-wiping occurring more often at remote sites after finches tasted flavours associated with human foods. Our results suggest, therefore, that while Darwin's finches regularly exposed to human foods might have acquired a tolerance to human food flavours, latent taste preferences are unlikely to have played a major role in their dietary response to increased urbanization.

Female in-nest attendance predicts the number of ectoparasites in Darwin's finch species.

Selection should act on parental care and favour parental investment decisions that optimize the number of offspring produced. Such predictions have been robustly tested in predation risk contexts, but less is known about alternative functions of parental care under conditions of parasitism. The avian vampire fly (Philornis downsi) is a myasis-causing ectoparasite accidentally introduced to the Galápagos Islands, and one of the major mortality causes in Darwin's finch nests. With an 11-year dataset spanning 21 years, we examine the relationship between parental care behaviours and number of fly larvae and pupae in Darwin's finch nests. We do so across three host species (Camarhynchus parvulus, C. pauper, Geospiza fuliginosa) and one hybrid Camarhynchus group. Nests with longer female brooding duration (minutes per hour spent sitting on hatchlings to provide warmth) had fewer parasites, and this effect depended on male food delivery to chicks. Neither male age nor number of nest provisioning visits were directly associated with number of parasites. While the causal mechanisms remain unknown, we provide the first empirical study showing that female brooding duration is negatively related to the number of ectoparasites in nests. We predict selection for coordinated host male and female behaviour to reduce gaps in nest attendance, especially under conditions of novel and introduced ectoparasites.

Evaluating Volatile Plant Compounds of Psidium galapageium (Myrtales: Myrtaceae) as Repellents Against Invasive Parasitic Diptera in the Galapagos Islands.

Plant-based repellents represent a safe, economic, and viable alternative to managing invasive insects that threaten native fauna. Observations of self-medication in animals can provide important cues to the medicinal properties of plants. A recent study in the Galapagos Islands found that Darwin's finches apply the leaves of Psidium galapageium (Hooker 1847) to their feathers, extracts of which were repellent to mosquitoes and the parasitic fly Philornis downsi (Dodge & Aitkens 1968; Diptera: Muscidae). Introduced mosquitoes are suspected vectors of avian pathogens in the Galapagos Islands, whereas the larvae of P. downsi are blood-feeders, causing significant declines of the endemic avifauna. In this study, we investigated the volatile compounds found in P. galapageium, testing each against a model organism, the mosquito Anopheles arabiensis (Patton 1905; Diptera: Culicidae), with the aim of singling out the most effective compound for repelling dipterans. Examinations of an ethanolic extract of P. galapageium, its essential oil and each of their respective fractions, revealed a mixture of monoterpenes and sesquiterpenes, the latter consisting mainly of guaiol, trans-nerolidol, and β-eudesmol. Of these, trans-nerolidol was identified as the most effective repellent to mosquitoes. This was subsequently tested at four different concentrations against P. downsi, but we did not find a repellence response. A tendency to avoid the compound was observed, albeit significance was not achieved in any case. The lack of repellence suggests that flies may respond to a combination of the volatile compounds found in P. galapageium, rather than to a single compound.

Geometry and dynamics link form, function, and evolution of finch beaks

Darwin's finches are a classic example of adaptive radiation, exemplified by their adaptive and functional beak morphologies. To quantify their form, we carry out a morphometric analysis of the three-dimensional beak shapes of all of Darwin's finches and find that they can be fit by a transverse parabolic shape with a curvature that increases linearly from the base toward the tip of the beak. The morphological variation of beak orientation, aspect ratios, and curvatures allows us to quantify beak function in terms of the elementary theory of machines, consistent with the dietary variations across finches. Finally, to explain the origin of the evolutionary morphometry and the developmental morphogenesis of the finch beak, we propose an experimentally motivated growth law at the cellular level that simplifies to a variant of curvature-driven flow at the tissue level and captures the range of observed beak shapes in terms of a simple morphospace. Altogether, our study illuminates how a minimal combination of geometry and dynamics allows for functional form to develop and evolve.

Urbanization alters interactions between Darwin's finches and Tribulus cistoides on the Galápagos Islands.

Emerging evidence suggests that humans shape the ecology and evolution of species interactions. Islands are particularly susceptible to anthropogenic disturbance due to the fragility of their ecosystems; however, we know little about the susceptibility of species interactions to urbanization on islands. To address this gap, we studied how the earliest stages of urban development affect interactions between Darwin's finches and its key food resource, Tribulus cistoides, in three towns on the Galápagos Islands. We measured variation in mericarp predation rates, mericarp morphology, and finch community composition using population surveys, experimental manipulations, and finch observations conducted in habitats within and outside of each town. We found that both seed and mericarp removal rates were higher in towns than natural habitats. We also found that selection on mericarp size and defense differed between habitats in the survey and experimental populations and that towns supported smaller and less diverse finch communities than natural habitats. Together, our results suggest that even moderate levels of urbanization can alter ecological interactions between Darwin's finches and T. cistoides, leading to modified natural selection on T. cistoides populations. Our study demonstrates that trophic interactions on islands may be susceptible to the anthropogenic disturbance associated with urbanization. Despite containing the highest diversity in the world, studies of urbanization are lacking from the tropics. Our study identified signatures of urbanization on species interactions in a tropical island ecosystem and suggests that changes to the ecology of species interactions has the potential to alter evolution in urban environments.

A multispecies BCO2 beak color polymorphism in the Darwin’s finch radiation

Carotenoid-based polymorphisms are widespread in populations of birds, fish, and reptiles,1 but generally little is known about the factors affecting their maintenance in populations.2 We report a combined field and molecular-genetic investigation of a nestling beak color polymorphism in Darwin's finches. Beaks are pink or yellow, and yellow is recessive.3 Here we show that the polymorphism arose in the Galápagos half a million years ago through a mutation associated with regulatory change in the BCO2 gene and is shared by 14 descendant species. The polymorphism is probably a balanced polymorphism, maintained by ecological selection associated with survival and diet. In cactus finches, the frequency of the yellow genotype is correlated with cactus fruit abundance and greater hatching success and may be altered by introgressive hybridization. Polymorphisms that are hidden as adults, as here, may be far more common than is currently recognized, and contribute to diversification in ways that are yet to be discovered.

Vocal learning in animals and humans.

Vocal learning in animals and humans.

Morphological ghosts of introgression in Darwin’s finch populations

Many species of plants, animals, and microorganisms exchange genes well after the point of evolutionary divergence at which taxonomists recognize them as species. Genomes contain signatures of past gene exchange and, in some cases, they reveal a legacy of lineages that no longer exist. But genomic data are not available for many organisms, and particularly problematic for reconstructing and interpreting evolutionary history are communities that have been depleted by extinctions. For these, morphology may substitute for genes, as exemplified by the history of Darwin's finches on the Galápagos islands of Floreana and San Cristóbal. Darwin and companions collected seven specimens of a uniquely large form of Geospiza magnirostris in 1835. The populations became extinct in the next few decades, partly due to destruction of Opuntia cactus by introduced goats, whereas Geospiza fortis has persisted to the present. We used measurements of large samples of G. fortis collected for museums in the period 1891 to 1906 to test for unusually large variances and skewed distributions of beak and body size resulting from introgression. We found strong evidence of hybridization on Floreana but not on San Cristóbal. The skew is in the direction of the absent G. magnirostris We estimate introgression influenced 6% of the frequency distribution that was eroded by selection after G. magnirostris became extinct on these islands. The genetic residuum of an extinct species in an extant one has implications for its future evolution, as well as for a conservation program of reintroductions in extinction-depleted communities.

Darwin’s finches habitually anoint their feathers with leaves of the endemic tree Psidium galapageium during the non‐breeding season

AbstractBirds host a wide range of ectoparasites and have developed behavioural strategies to combat them, such as preening, dust bathing and water bathing. In addition, a wide range of avian taxa anoint their feathers with insects or plants that have pharmaceutical properties, though most observations on anointing are anecdotal. Darwin's finches preen with leaves of an endemic tree (Psidium galapageium) and a previous laboratory study has shown that this plant has compounds that repel both mosquitoes and the invasive parasitic fly Philornis downsi, whose larvae suck blood from nestlings and incubating females and cause high nestling mortality. In the current study, we tested the hypothesis that preening with P. galapageium leaves serves to repel these parasites with an indirect approach. Mosquitoes and P. downsi affect their hosts mainly during the bird breeding season and P. downsi only affects breeding females, but not adult males. To test our hypothesis, we gathered quantitative data on leaf‐preening behaviour in Darwin's finches during their breeding and non‐breeding season and also investigated the influence of time of day and humidity, as humid conditions facilitate the release of volatile organic compounds. Contrary to our predictions, anointing occurred significantly more often during the non‐breeding season when mosquito and P. downsi numbers are lower. Four Darwin's finch species anointed their feathers habitually, and during the non‐breeding season, 56% of all preening events were with leaves. We found no effect of sex, but preening with leaves occurred predominately in the morning when leaves were wet. Our study is the first to provide quantitative data on anointing behaviour in birds and the high percentage of preening with leaves in the non‐breeding season suggests that the behaviour has an adaptive value. However, further studies are needed to test whether it reduces the negative impact of parasites other than mosquitoes and P. downsi.

Behavioural response to songs between genetically diverged allopatric populations of Darwin's small tree finch in the Galápagos.

Empirical data that identify contemporary mechanisms of divergence shed light on how species could multiply. In this study, we measured population genetic structure, song syllable diversity and response to simulated intruder song in Darwin's small tree finch (Camarhynchus parvulus) on Santa Cruz and Floreana Islands, Galápagos archipelago. Our aim was to test whether the magnitude of contemporary behavioural response in resident birds was consistent with patterns of genetic or cultural differences between populations. We analysed genetic structure and the occurrence of song syllable types, and experimentally measured the response of resident birds to intruder bird song from different geographical origin (i.e., island) or syllable type. We discovered a weak signal of population genetic structure between Santa Cruz and Floreana Islands. Although some song syllables occurred on both islands, others were unique to each island; Santa Cruz Island males used more unique syllables than Floreana Island males. Both Santa Cruz and Floreana resident males discriminated their response towards a simulated intruder song based on the geographical origin of the intruder song, but not on the syllable type sung by the intruder. We conclude that the populations are diverging in genetic and cultural traits and identified a signal of contemporary behavioural response that could maintain divergence upon secondary contact.

Urban living influences the nesting success of Darwin's finches in the Galápagos Islands.

Urbanization is expanding worldwide with major consequences for organisms. Anthropogenic factors can reduce the fitness of animals but may have benefits, such as consistent human food availability. Understanding anthropogenic trade‐offs is critical in environments with variable levels of natural food availability, such as the Galápagos Islands, an area of rapid urbanization. For example, during dry years, the reproductive success of bird species, such as Darwin's finches, is low because reduced precipitation impacts food availability. Urban areas provide supplemental human food to finches, which could improve their reproductive success during years with low natural food availability. However, urban finches might face trade‐offs, such as the incorporation of anthropogenic debris (e.g., string, plastic) into their nests, which may increase mortality. In our study, we determined the effect of urbanization on the nesting success of small ground finches (Geospiza fuliginosa; a species of Darwin's finch) during a dry year on San Cristóbal Island. We quantified nest building, egg laying and hatching, and fledging in an urban and nonurban area and characterized the anthropogenic debris in nests. We also documented mortalities including nest trash‐related deaths and whether anthropogenic materials directly led to entanglement‐ or ingestion‐related nest mortalities. Overall, urban finches built more nests, laid more eggs, and produced more fledglings than nonurban finches. However, every nest in the urban area contained anthropogenic material, which resulted in 18% nestling mortality while nonurban nests had no anthropogenic debris. Our study showed that urban living has trade‐offs: urban birds have overall higher nesting success during a dry year than nonurban birds, but urban birds can suffer mortality from anthropogenic‐related nest‐materials. These results suggest that despite potential costs, finches benefit overall from urban living and urbanization may buffer the effects of limited resource availability in the Galápagos Islands.

Sex identification in embryos and adults of Darwin's finches.

Darwin's finches are an iconic example of adaptive radiation and evolution under natural selection. Comparative genetic studies using embryos of Darwin's finches have shed light on the possible evolutionary processes underlying the speciation of this clade. Molecular identification of the sex of embryonic samples is important for such studies, where this information often cannot be inferred otherwise. We tested a fast and simple chicken embryo protocol to extract DNA from Darwin's finch embryos. In addition, we applied minor modifications to two of the previously reported PCR primer sets for CHD1, a gene used for sexing adult passerine birds. The sex of all 29 tested embryos of six species of Darwin's finches was determined successfully by PCR, using both primer sets. Next to embryos, hatchlings and fledglings are also impossible to distinguish visually. This extends to juveniles of sexually dimorphic species which are yet to moult in adult-like plumage and beak colouration. Furthermore, four species of Darwin's finches are monomorphic, males and females looking alike. Therefore, sex assessment in the field can be a source of error, especially with respect to juveniles and mature monomorphic birds outside of the mating season. We caught 567 juveniles and adults belonging to six species of Darwin's finches and only 44% had unambiguous sex-specific morphology. We sexed 363 birds by PCR: individuals sexed based on marginal sex specific morphological traits; and birds which were impossible to classify in the field. PCR revealed that for birds with marginal sex specific traits, sexing in the field produced a 13% error rate. This demonstrates that PCR based sexing can improve field studies on Darwin's finches, especially when individuals with unclear sex-related morphology are involved. The protocols used here provide an easy and reliable way to sex Darwin's finches throughout ontogeny, from embryos to adults.

The flippingt‐ratio test: Phylogenetically informed assessment of the Pareto theory for phenotypic evolution

AbstractAn organism cannot be fully optimized for all tasks that are needed for its survival because of the existence of intrinsic trade‐offs among those tasks. It was recently proposed that an economics‐originated theory, the Pareto theory, is a general theory explaining the rules of phenotypic evolution under such trade‐offs. To date, many phenotype datasets have been argued to fit the Pareto theory based on a statistical method named thet‐ratio test.Here, we show that this test suffers a serious defect in that general phenotype data can be wrongly concluded to fit the Pareto theory with a very high false‐positive rate and that the claim that the Pareto theory is a general theory should definitely be considered with caution. This is because thet‐ratio test assumes that all phenotypic traits are independent of one another, but this assumption does not hold true—different traits of organisms have usually been affected by the same phylogenetic history and are thus typically not independent.We developed the flippingt‐ratio test to accurately test the Pareto theory by considering phylogenetic background as well as artefacts that can be induced during dimensionality reduction. Using this improved method, we confirm that the phenotype data analysed in previous studies, including the well‐known Darwin's ground finch dataset, no longer support the Pareto theory.We hope that the flippingt‐ratio test will contribute to examining which phenotype datasets truly fit the Pareto theory and understanding how diverse phenotypes evolve in natural ecosystems.

Structural variation and phylogenetic relationship of Geospiza magnirostris based on mitochondrial control region

The control region of mitochondrial genome is the main non-coding region. There is no details about Geospiza magnirostris’s control region, who is the only species of Geospiza in Darwin finches whose mitochondrial genome has been sequenced. In this study, the control region mitochondrial genome of G. magnirostris was analyzed. Compare research for 15 Passerine species, belonging to 4 families (Darwin ground finch, Thraupidae, Icteridae, Emberizidae), was also been done. The result showed the length was 1202 bp and it could been divided into three parts, domainI, domainII and domainIII. Genetic distances showed average divergence of different family ranged from 0.36% (Darwin ground finch) to 9.20% (Emberizidae), indicated that the Darwin ground finch may be a very conservative single-source species. Some specific sequence fragments such as conservative sequence box, boxes C, D, F and E, HSP/LSP, CSB-1, TAS-1 were also found in the control region of G. magnirostris, while B-box, CSB-2 and CSB-3 were not found in it. The F-box was detected in all selected 15 species and was 29 bp in length with no variant sites. In the E-box, 20 of the 21 nucleotide positions were identical as conserved among the selected species. There are two variant sites in the D-box, and five variant sites in the C-box. Basing the phylogenetic tree, it can be clearly seen that the relationship of G. magnirostris and family Thraupidae is the closest in all selected family. Through this study, the structure characteristics of G. magnirostris control region were explored and understood. This study provided more basic information for in-depth research of G. magnirostris. It’s also reminded us that we should pay more attention to explore the characteristics of the control region of Darwin’s finches.

Ecological and morphological determinants of evolutionary diversification in Darwin's finches and their relatives.

Darwin's finches are a classic example of adaptive radiation, a process by which multiple ecologically distinct species rapidly evolve from a single ancestor. Such evolutionary diversification is typically explained by adaptation to new ecological opportunities. However, the ecological diversification of Darwin's finches following their dispersal to Galápagos was not matched on the same archipelago by other lineages of colonizing land birds, which diversified very little in terms of both species number and morphology. To better understand the causes underlying the extraordinary variation in Darwin's finches, we analyze the evolutionary dynamics of speciation and trait diversification in Thraupidae, including Coerebinae (Darwin's finches and relatives) and, their closely related clade, Sporophilinae. For all traits, we observe an early pulse of speciation and morphological diversification followed by prolonged periods of slower steady‐state rates of change. The primary exception is the apparent recent increase in diversification rate in Darwin's finches coupled with highly variable beak morphology, a potential key factor explaining this adaptive radiation. Our observations illustrate how the exploitation of ecological opportunity by contrasting means can produce clades with similarly high diversification rate yet strikingly different degrees of ecological and morphological differentiation.

The Radiation of Darwin’s Giant Daisies in the Galápagos Islands

Evolutionary radiations on oceanic islands have fascinated biologists since Darwin's exploration of the Galápagos archipelago [1, 2]. Island radiations can provide key insights for understanding rapid speciation, including evolutionary patterns and the processes behind them. However, lack of resolution of species relationships has historically hindered their investigation, particularly in the plant kingdom [3-5]. Here, we report a time-calibrated phylogenomic analysis based on genotyping-by-sequencing data [6] of the 15 species of Scalesia (Darwin's giant daisies), an iconic and understudied plant radiation endemic to the Galápagos Islands and considered the plant counterpart to Darwin's finches [1, 7-9]. Results support a Pliocene to early Pleistocene divergence between Scalesia and the closest South American relatives, and rapid diversification of extant Scalesia species from a common ancestor dated to the Middle Pleistocene. Major evolutionary patterns in Scalesia include the following: (1) lack of compliance with the "progression rule" hypothesis, in which earlier diverging lineages are expected to occupy older islands; (2) a predominance of within-island speciation over between-island speciation; and (3) repeated convergent evolution of potentially adaptive traits and habitat preferences on different islands during the course of diversification. Massive sequencing provided the essential framework for investigating evolutionary and ecological processes in the complex natural laboratory of the Galápagos, thereby advancing our understanding of island plant radiations.