Island Syndrome Research Articles

Convergent evolution toward a slow pace of life predisposes insular endotherms to anthropogenic extinctions.

Island vertebrates have evolved a number of morphological, physiological, and life history characteristics that set them apart from their mainland relatives. However, to date, the evolution of metabolism and its impact on the vulnerability to extinction of insular vertebrates remains poorly understood. This study used metabolic data from 2813 species of tetrapod vertebrates, including 695 ectothermic and 2118 endothermic species, to reveal that island mammals and birds evolved convergent metabolic strategies toward a slow pace of life. Insularity was associated with shifts toward slower metabolic rates and greater generation lengths in endotherms, while insularity just drove the evolution of longer generation lengths in ectotherms. Notably, a slow pace of life has exacerbated the extinction of insular endemic species in the face of anthropogenic threats. These findings have important implications for understanding physiological adaptations associated with the island syndrome and formulating conservation strategies across taxonomic groups with different metabolic modes.

Genome-wide relaxation of selection and the evolution of the island syndrome in Orkney voles.

Island populations often experience different ecological and demographic conditions than their counterparts on the continent, resulting in divergent evolutionary forces affecting their genomes. Random genetic drift and selection both may leave their imprints on island populations, although the relative impact depends strongly on the specific conditions. Here we address their contributions to the island syndrome in a rodent with an unusually clear history of isolation. Common voles (Microtus arvalis) were introduced by humans on the Orkney archipelago north of Scotland >5000 years ago and rapidly evolved to exceptionally large size. Our analyses show that the genomes of Orkney voles were dominated by genetic drift, with extremely low diversity, variable Tajima's D, and very high divergence from continental conspecifics. Increased d N/d S ratios over a wide range of genes in Orkney voles indicated genome-wide relaxation of purifying selection. We found evidence of hard sweeps on key genes of the lipid metabolism pathway only in continental voles. The marked increase of body size in Orkney-a typical phenomenon of the island syndrome-may thus be associated to the relaxation of positive selection on genes related to this pathway. On the other hand, a hard sweep on immune genes of Orkney voles likely reflects the divergent ecological conditions and possibly the history of human introduction. The long-term isolated Orkney voles show that adaptive changes may still impact the evolutionary trajectories of such populations despite the pervasive consequences of genetic drift at the genome level.

Behavioral Island Syndrome and Its Ecological Drivers in the Mednyi Island Arctic Fox Population

Behavioral Island Syndrome and Its Ecological Drivers in the Mednyi Island Arctic Fox Population

Prenatal and neonatal phenotype of Larsen of La Réunion Island syndrome (B4GALT7-linkeropathy)

Larsen of La Réunion Island syndrome (LRS) is an autosomal recessive condition associated with multiple large joint dislocations, clubfeet, severe dwarfism, and distinctive facial features. LRS is caused by a recurrent homozygous variant in B4GALT7 gene with a founder effect in La Réunion population. Proteoglycans (PG) that are a major component of the extracellular matrix, are composed of a core protein connected to a glycosaminoglycans side chain via a tetrasaccharide linker region. B4GALT7 encodes galactosyltransferase I, one of the enzymes involved in the biosynthesis of the linker region. Conditions caused by pathogenic biallelic variants in genes implicated in the synthesis of the tetrasaccharide linker of PG are known as linkeropathies.Prenatal features are rarely described in this group of chondrodysplasias. We present a series of 12 unpublished patients having LRS and describe the perinatal phenotype. All the patients had a prenatal growth restriction with brevity of limbs. The other features revealed by ultrasounds were increased nuchal translucency at 10–12 weeks of gestation (50 %), feet abnormalities (clubfeet or metatarsus varus) (25 %), dislocation affecting at least one large joint (elbow, knee, wrist) (25 %). Bilateral bowing of femora was noted for two fetuses. Fibular hypertrophy was noted for one fetus. Prenatal helical computed tomography (CT) performed in three pregnancies showed additional data such as bowing of the forearm bones, proximal radio-ulnar synostosis, or dislocation of large joints. Prenatal sonographic and helical CT findings led to the prenatal diagnosis of LRS in four patients. We confirm that the neonatal clinical picture of LRS has an important overlap with that reported in patients with B4GALT7 deficiency outside La Réunion Island and other linkeropathies. The core of the phenotypic spectrum combines low birth height, micromelia, hypermobility, dislocation of at least one large joint, facial features with prominent eyes, microstomia, depressed nasal bridge, and midface hypoplasia. Other clinical features include clubfeet (33%), bifid thumb in one patient, and cardiac abnormalities in two patients. Radiological findings include radio-ulnar synostosis (75%), metaphyseal flaring, precocious carpal ossification, and a Swedish key appearance of the proximal femora. Finally, we also report radiological features rarely described in B4GALT7-linkeropathies, including bowing of the femora and fibular hypertrophy. Our results confirm the phenotypic continuum of LRS within linkeropathies with some additional findings, including a high frequency of clubfeet usually described in B3GALT6-linkeropathies, the presence of congenital heart diseases usually described in B3GAT3-linkeropathies, and a high frequency of metaphyseal flaring usually reported in B3GALT6 or XITLT1-linkeropathies.This is the first study that describes the perinatal phenotype in a cohort of patients with LRS. This study can help improve the prenatal diagnosis of the linkeropathies and add this group of conditions to the differential diagnosis of chondrodysplasias with multiple dislocations.In view of the founder effect for LRS in La Réunion Island, this disease should be suspected in fetuses with growth restriction and micromelia. Thus in case of LOH which include B4GALT7 identified in SNP-array, we recommend performing a targeted Sanger sequencing for the recurrent mutation c.808C > T; p. (Arg270Cys).

Evidence for Reductions in Physical and Chemical Plant Defense Traits in Island Flora.

Reduced defense against large herbivores has been suggested to be part of the "island syndrome" in plants. However, empirical evidence for this pattern is mixed. In this paper, we present two studies that compare putative physical and chemical defense traits from plants on the California Channel Islands and nearby mainland based on sampling of both field and common garden plants. In the first study, we focus on five pairs of woody shrubs from three island and three mainland locations and find evidence for increased leaf area, decreased marginal leaf spines, and decreased concentrations of cyanogenic glycosides in island plants. We observed similar increases in leaf area and decreases in defense traits when comparing island and mainland genotypes grown together in botanic gardens, suggesting that trait differences are not solely driven by abiotic differences between island and mainland sites. In the second study, we conducted a common garden experiment with a perennial herb-Stachys bullata (Lamiaceae)-collected from two island and four mainland locations. Compared to their mainland relatives, island genotypes show highly reduced glandular trichomes and a nearly 100-fold reduction in mono- and sesquiterpene compounds from leaf surfaces. Island genotypes also had significantly higher specific leaf area, somewhat lower rates of gas exchange, and greater aboveground biomass than mainland genotypes across two years of study, potentially reflecting a broader shift in growth habit. Together, our results provide evidence for reduced expression of putative defense traits in island plants, though these results may reflect adaptation to both biotic (i.e., the historical absence of large herbivores) and climatic conditions on islands.

Birds that breed exclusively on islands have smaller clutches

Abstract The “island syndrome” refers to similarity in the biology of island organisms, but its generality is questionable, as the scope of species and traits examined are often limited. Here, I show that birds breeding exclusively on islands (breeding island endemics) evolved smaller clutches, using a dataset of 4,530 bird species. Using an inclusive definition of a breeding island endemic, which also encompasses migratory species and seabirds, I examine the evolution of clutch sizes in island breeding species using phylogenetic generalized linear models. Across disparate phylogenetic hypotheses, and after accounting for biological and geographical co-variables, I show that breeding island endemic landbirds (470 species) evolved smaller clutch sizes than continental breeding species (3,818 species). I show that the evolution of clutch size follows the expectations of the island syndrome, as among breeding island endemic landbirds there is a positive relationship between clutch size and breeding range area. Finally, I reinforce the view that the island syndrome is a general pattern in birds, spanning diverse phylogenetic and ecological groups, by showing that in a seabird-only dataset (242 species), breeding island endemic seabirds show evolution of smaller clutch sizes. In a model of the full dataset of both landbirds and seabirds (4,530 species) there was no evidence of an interaction of being a seabird with breeding island endemicity, showing that seabirds and landbirds respond in the same way. This study, using more than 40% of all bird species, provides the first evidence of a general evolutionary response in a life history trait, clearly showing the island syndrome as a general evolutionary tendency associated with island environments.

Evolutionary Plasticity in Insular Lizard, Adapting over Reproduction, Metabolism, and Color Variation.

The Italian wall lizard (Podarcis siculus) living on islets exhibits a melanic skin coloration and a suite of adaptive traits lacking in nearby mainland populations. On islets, the unpredictable environmental conditions and highly fluctuating population densities are believed to have produced reversed island syndrome (RIS). Several physiological, behavioral, and life-history changes based on the RIS could result from positive selection on increased activity of melanocortins. We hypothesize that phenotypes on islets are the product of a plastic variation depending on the regulation of specific genes. Focusing on control systems that determine the insular-adapted phenotype, we demonstrated that reproductive markers, involved in the hypothalamus-hypophysis-gonadal axis, and metabolism markers, flags for hypophysis-melanocortin receptors, are all up-regulated in island lizards under the RIS. This behavior, combined with the observed limited variation in the mitochondrial genome, agrees with the hypothesis that plasticity enables populations to persist in novel environmental conditions and that over time, natural selection will "fine-tune" the population to the environment by modifying the phenotype under selection. We believe that analysis of the transcriptome and the single gene expression, such that all the variations observed in the island populations, can be useful to shed light on evolutionary plasticity as a process affecting animals' populations in general.

A test of island plant syndromes using resource‐use traits

AbstractDespite representing a fraction of the global terrestrial surface area, oceanic islands are disproportionately diverse in species, resulting from high rates of endemicity. Island plants are thought to share a unique phenotype—referred to as an island syndrome—which is thought to be driven by convergent evolution in response to selection by shared abiotic and biotic factors. One aspect of the island plant syndrome that has received relatively little research focus is that island plants are expected to have converged on conservative resource use associated with slow growth rates and weak competitive abilities. Here we tested whether native, woody Hawaiian plant species are phenotypically distinct—with more resource‐conservative leaf traits—compared to a globally distributed sample of continental species. Using an archipelago‐wide trait data set, we detected that on average, native Hawaiian species had lower leaf nutrient concentrations overall, and lower nutrient concentrations at high leaf mass per area, but no other phenotypic differences compared with continental plants. There was also considerable overlap in the trait spaces of native Hawaiian species and continental species. Our findings indicate that an island plant syndrome for leaf traits is not present in the Hawaiian flora, and that island species can demonstrate extensive variation in their resource‐use strategies, on a scale that is comparable with that of continental species worldwide.

The island syndrome in birds

AbstractThe island syndrome is a widespread biological phenomenon that describes a suite of morphological, behavioural, demographic and life‐history changes associated with island dwelling. These similar evolutionary responses among disparate groups of animals and plants represent a remarkable case of convergent evolution. Among animals, birds are a highly suitable group to study the island syndrome; they are a comparatively data‐rich taxon, are frequent island colonisers, and sometimes display extreme adaptations such as the loss of flight. However, the avian island syndrome literature is fragmented, and multiple components are rarely considered together even though many are inextricably linked. We reviewed multi‐species comparative studies, single‐species or population‐level studies and anecdotal accounts, to summarise and assess the support for individual components of the island syndrome for birds, and to identify suites of traits that should be considered together. The weight of evidence for island syndrome patterns in morphology is substantial, but is more partial or even anecdotal for various aspects of behaviour, life history and physiology. Full validation of the island syndrome in birds will require the less‐studied components to be treated in a comparative framework, and for covarying components to be examined in an integrated way. An improved description of the scope of the syndrome will pave the way to understanding its drivers.

Towards a holistic perspective on the development of island syndrome by examining its occurrence patterns in insular plants

Towards a holistic perspective on the development of island syndrome by examining its occurrence patterns in insular plants

Insular giant leporid matured later than predicted by scaling

The island syndrome describes morphological, behavioral, and life history traits that evolve in parallel in endemic insular organisms. A basic axiom of the island syndrome is that insular endemics slow down their pace of life. Although this is already confirmed for insular dwarfs, a slow life history in giants may not be adaptive, but merely a consequence of increasing body size. We tested this question in the fossil insular giant leporid Nuralagus rex. Using bone histology, we constructed both a continental extant taxon model derived from experimentally fluorochrome-labeled Lepus europaeus to calibrate life history events, and a growth model for the insular taxon. N.rex grew extremely slowly and delayed maturity well beyond predictions from continental phylogenetically corrected scaling models. Our results support the life history axiom of the island syndrome as generality for insular mammals, regardless of whether they have evolved into dwarfs or giants.

Insularity determines nestling sex ratio variation in Egyptian vulture populations.

Variation in offspring sex ratio, particularly in birds, has been frequently studied over the last century, although seldom using long-term monitoring data. In raptors, the cost of raising males and females is not equal, and several variables have been found to have significant effects on sex ratio, including food availability, parental age, and hatching order. Sex ratio differences between island populations and their mainland counterparts have been poorly documented, despite broad scientific literature on the island syndrome reporting substantial differences in population demography and ecology. Here, we assessed individual and environmental factors potentially affecting the secondary sex ratio of the long-lived Egyptian vulture Neophron percnopterus. We used data collected from Spanish mainland and island populations over a ca. 30-year period (1995-2021) to assess the effects of insularity, parental age, breeding phenology, brood size, hatching order, type of breeding unit (pairs vs. trios), and spatial and temporal variability on offspring sex ratio. No sex bias was found at the population level, but two opposite trends were observed between mainland and island populations consistent with the island syndrome. Offspring sex ratio was nonsignificantly female-biased in mainland Spain (0.47, n = 1112) but significantly male-biased in the Canary Islands (0.55, n = 499), where a male-biased mortality among immatures could be compensating for offspring biases and maintaining a paired adult sex ratio. Temporal and spatial variation in food availability might also have some influence on sex ratio, although the difficulties in quantifying them preclude us from determining the magnitude of such influence. This study shows that insularity influences the offspring sex ratio of the Egyptian vulture through several processes that can affect island and mainland populations differentially. Our research contributes to improving our understanding of sex allocation theory by investigating whether sex ratio deviations from parity are possible as a response to changing environments comprised by multiple and complexly interrelated factors.

Exploring island syndromes: Variable matrix permeability in Phalaenopsis pulcherrima (Orchidaceae), a specialist lithophyte of tropical Asian inselbergs.

Plants confined to island-like habitats are hypothesised to possess a suite of functional traits that promote on-spot persistence and recruitment, but this may come at the cost of broad-based colonising potential. Ecological functions that define this island syndrome are expected to generate a characteristic genetic signature. Here we examine genetic structuring in the orchid Phalaenopsis pulcherrima, a specialist lithophyte of tropical Asian inselbergs, both at the scale of individual outcrops and across much of its range in Indochina and on Hainan Island, to infer patterns of gene flow in the context of an exploration of island syndrome traits. We sampled 323 individuals occurring in 20 populations on 15 widely scattered inselbergs, and quantified genetic diversity, isolation-by-distance and genetic structuring using 14 microsatellite markers. To incorporate a temporal dimension, we inferred historical demography and estimated direction of gene flow using Bayesian approaches. We uncovered high genotypic diversity, high heterozygosity and low rates of inbreeding, as well as strong evidence for the occurrence of two genetic clusters, one comprising the populations of Hainan Island and the other those of mainland Indochina. Connectivity was greater within, rather than between the two clusters, with the former unequivocally supported as ancestral. Despite a strong capacity for on-spot persistence conferred by clonality, incomplete self-sterility and an ability to utilize multiple magnet species for pollination, our data reveal that P. pulcherrima also possesses traits that promote landscape-scale gene flow, including deceptive pollination and wind-borne seed dispersal, generating an ecological profile that neither fully conforms to, nor fully contradicts, a putative island syndrome. A terrestrial matrix is shown to be significantly more permeable than open water, with the direction of historic gene flow indicating that island populations can serve as refugia for postglacial colonisation of continental landmasses by effective dispersers.

Behavioral island syndrome and its ecological drivers in the Mednyi Island Arctic fox

Reptiles, birds, and mammals inhabiting oceanic islands tend to change many ecological, behavioral, and genetic characteristics. These changes are referred to as the “island syndrome”. The behavioral components of these changes have been little studied so far. Based on a 40-year study of the biology of the Arctic fox (Vulpes lagopus) on the Mednyi Island, we show that the island Arctic foxes changed many behavioral characteristics compared to mainland foxes. They use smaller home ranges, travel shorter daily distances, and their dispersal distance from the natal territory is much smaller. An increase in the family size of the island Arctic foxes, together with a decrease in litter size, leads to an increase in parental and kin investment and increased cooperation between family members. At the same time, the island foxes, to a large extent, lost their fear of humans. These features are consistent with those found in other island populations. On the other hand, in contrast to other island populations, Mednyi Arctic foxes have increased territoriality, apparently in response to an increased risk of infanticide. Furthermore, Mednyi Arctic foxes expose sex-biased dispersal and maternal inheritance of home range, which are absent in the mainland foxes. The sex-biased dispersal can be considered a mechanism for avoiding inbreeding. Comparison with island fox (Urocyon littoralis) populations in the Channel Islands (Southern California) suggests that these traits have evolved under the influence of specific environmental drivers of the Mednyi Island: patchy and highly predictable resource distribution. The particular resource distribution led to the increased importance of another driver, social landscape, and a new behavioral metatrait of the Mednyi fox – increased conservatism in spatial, reproductive, and foraging behavior.

The ‘island syndrome’ is an alternative state

AbstractAimIn the half‐century since publication of the Theory of Island Biology, ecologists have come to recognize the importance of predation as a decisive determinant of alternate states in many ecosystems. Island species are notorious for their vulnerability to introduced predators, yet the strength of island predator regimes has not been fully incorporated into our understanding of the forces that structure island consumer communities.LocationThe Greater and Lesser Antilles.TaxonBirds andAnolislizards.MethodsField surveys of sclerophyll and rainforest sites on islands ranging in size from 3.5 km2Terre‐de‐Haut to 76,000 km2Hispaniola.ResultsEvidence gathered in the 1970s and 1980s shows that Antillean anoles live at higher densities on fewer resources, grow more slowly, reproduce later and live longer than mainland counterparts in conformity with the ‘island syndrome’. Data from this period show that Antillean bird communities display density overcompensation, community saturation, size‐structured foraging guilds, low species diversity and low species packing, all traits consistent with the island syndrome and a regime of low predation and intense competition. Mainland species and communities display none of these features.Main conclusionsI propose that the island syndrome is an alternative state that distinguishes low‐predation island communities from high‐predation mainland counterparts. It follows that strong mainland predation regimes tend to prevent island species from colonizing. Conversely, invasion‐resistant, size‐structured island communities, despite low species diversity, prevent mainland species from colonizing islands. These predictions are experimentally testable withAnolislizards and, if confirmed, could set island biogeography on a new course.

Evolution of immune genes in island birds: reduction in population sizes can explain island syndrome

Shared ecological conditions encountered by species that colonize islands often lead to the evolution of convergent phenotypes, commonly referred to as “island syndrome”. Reduced immune functions have been previously proposed to be part of this syndrome, as a consequence of the reduced diversity of pathogens on island ecosystems. According to this hypothesis, immune genes are expected to exhibit genomic signatures of relaxed selection pressure in island species. In this study, we used comparative genomic methods to study immune genes in island species (N = 20) and their mainland relatives (N = 14). We gathered public data as well as generated new data on innate (TLR: Toll-Like Receptors, BD: Beta Defensins) and acquired immune genes (MHC: Major Histocompatibility Complex classes I and II), but also on hundreds of genes with various immune functions. As a control, we used a set of 97 genes, not known to be involved in immune functions based on the literature, to account for the increased drift effects of the lower effective population sizes in island species. We used synonymous and non-synonymous variants to estimate the selection pressure acting on immune genes. We found that BDs and TLRs have higher ratios of non-synonymous over synonymous polymorphisms (Pn/Ps) than randomly selected control genes, suggesting that they evolve under a different selection regime. However, simulations show that this is unlikely to be explained by ongoing positive selection or balancing selection. For the MHC genes, which evolve under balancing selection, we used simulations to estimate the impact of population size variation. We found a significant effect of drift on immune genes of island species leading to a reduction in genetic diversity and efficacy of selection. However, the intensity of relaxed selection was not significantly different from control genes, except for MHC class II genes. These genes exhibit a significantly higher level of non-synonymous loss of polymorphism than expected assuming only drift and evolution under frequency dependent selection, possibly due to a reduction of extracellular parasite communities on islands. Overall, our results showed that demographic effects lead to a decrease in the immune functions of island species, but the relaxed selection that is expected to be caused by a reduced parasite pressure may only occur in some categories of immune genes.

Epigenetics and island-mainland divergence in an insectivorous small mammal.

Geographically isolated populations, specifically island-mainland counterparts, tend to exhibit phenotypic variation in many species. The so-called island syndrome occurs when different environmental pressures lead to insular divergence from mainland populations. This phenomenon can be seen in an island population of Nova Scotia masked shrews (Sorex cinereus), which have developed a specialized feeding habit and digestive enzyme compared to their mainland counterparts. Epigenetic modifications, such as DNA methylation (DNAm), can impact phenotypes by altering gene expression without changing the DNA sequence. Here, we used a de novo masked shrew genome assembly and a mammalian methylation array profiling 37 thousand conserved CpGs to investigate morphological and DNA methylation patterns between island and mainland populations. Island shrews were morphologically and epigenetically different than their mainland counterparts, exhibiting a smaller body size. A gene ontology enrichment analyses of differentially methylated CpGs implicated developmental and digestive system related pathways. Based on our shrew epigenetic clock, island shrews might also be aging faster than their mainland counterparts. This study provides novel insight on phenotypic and epigenetic divergence in island-mainland mammal populations and suggests an underlying role of methylation in island-mainland divergence.

Insularity and early domestication: anthropogenic ecosystems as habitat islands

Over the past decade research into early domestication has been transformed by the genomics revolution and increased archaeological investigation. Despite clarification of the timing, locations and genetic processes, most scholars still envision evolutionary responses to human innovations, such as sickle harvesting, tilling, selection for docility or directed breeding. Stepping away from anthropocentric models, evolutionary parallels in the wild can provide case studies for understanding what ecological pressures drove the evolution of the first domestication traits. I contrast evolutionary trends seen among plants and animals confined on oceanic islands with the changes seen in the first cultivated crops and animals. I argue that the earliest villages functioned as habitat islands, applying parallel selective pressures as those on oceanic islands. In this view, the collective assemblage of parallel evolving traits that some scholars refer to as either an island syndrome or domestication syndrome results from similar ecological pressures of insularity, notably ecological release.

Parasite island syndromes in the context of nidicolous ectoparasites: Fleas (Insecta: Siphonaptera) in wild passerine birds from Azores Archipelago.

Island syndrome, previously established for isolation process of insular vertebrates' populations, have been adapted to insular parasites communities, termed parasite island syndromes. In this work, were studied for the first time the insular syndromes for nidicolous ectoparasites of the bird species, Turdus merula, Sylvia atricapilla, Fringilla coelebs and Erithacus rubecula from Azores and the mainland Portugal. Flea species were only recorded on Azorean birds, namely Dasypsyllus gallinulae and Ctenocephalides felis felis, known as not host-specific parasites. In the absence of shared flea species between mainland and islands birds, a comparison among our fleas prevalence to Azores Islands and mainland fleas prevalence, recorded to others European studies, showed that Azorean host populations undergo higher prevalence than the mainland one. This result was consistent with parasite island syndromes predictions recorded to ectoparasites, hippoboscid flies and chewing lice, that fleas have higher prevalence on the Azores Islands compared to mainland Portugal. However, our results provide a new perspective to parasite island syndromes assumptions, namely in the context of nidicolous ectoparasites that spend only brief periods on the hosts' body.

Gut microbiota plasticity in insular lizards under reversed island syndrome

Animals living on small islands are more drastically exposed to environmental changes, such as food or water starvation, and rapid temperature shifts. Facing such conditions, and probably thank to adaptive plasticity mechanisms, some animals display a Reversed Island Syndrome (RIS), a suite of traits, including skin pigmentation, voracity, sexual dimorphism, showed differently from mainland relatives. Here, we analyse a so far poorly explored aspect of RIS: the effect of this on the microbiota composition of host Italian wall lizard (Podarcis siculus), strongly influenced by the animal’s lifestyle, and conditioning the same. We compare mainland and island populations, assessing the difference between their microbial communities and their response under unexpected food, experimentally provided. Our observations showed a significant difference in microbiota communities between island and mainland groups, depended mainly from changes in relative abundance of the shared genera (difference due to decrease/increase). Exposure to experimental diet regimes resulted into significative reshaping of bacterial composition of microbiota and a greater variation in body mass only in the island population. Our results could be an evidence that gut microbial community contributes to adaptive plasticity mechanisms of island lizards under RIS to efficiently respond to unexpected changes.