The ecological and evolutionary dynamics of inselbergs.

Islands are fundamental model systems in ecology, biogeography, and evolutionary biology. However, terrestrial islands, unlike their aquatic counterparts, have received comparatively less attention. Among these land islands, inselbergs (i.e. isolated rock outcrops with diverse lithologies and a modest topographical prominence) stand out as iconic examples distributed worldwide across global biomes. Due to their durable lithology, inselbergs change slowly, persisting for tens of millions of years. In this review, we propose a biological definition for inselbergs that captures three fundamental characteristics of inselbergs from the perspective of biota. These are old age, isolation and the presence of unique microhabitats that are rare or absent in the surrounding matrix, fostering distinct communities often with unique and endemic biota. We synthesise the state of the art and formulate a set of testable hypotheses to deepen our understanding of the origins and maintenance of diversity on inselbergs, which are increasingly exposed to anthropogenic threats. By offering different habitats compared to the surrounding habitat matrix (e.g. moist microhabitats in dryland landscapes and xeric environments in humid tropical landscapes), inselbergs may allow specific lineages to thrive beyond their typical geographical limits. Particularly in drylands and degraded landscapes, inselbergs may not just provide different habitats but also act as ecological refuges or evolutionary refugia by providing a wider range of potential microhabitats than the surrounding matrix, enhancing resilience and promoting regional biodiversity. The central role of the matrix ensures that the ecological and evolutionary dynamics of inselbergs differ from those of true islands such as oceanic islands. Given that inselberg biota coexist within a terrestrial matrix, interactions between inselberg and matrix populations impact each other significantly. Over evolutionary timescales, matrix species may contract to inselberg refugia, preserving lineages while cycles of isolation and reconnection may drive speciation via a species pump. Although inselberg biodiversity has been studied predominantly from an island biogeography perspective, we argue that depending on the spatial scale, habitat specificity and mobility of the organisms considered, a range of different theories and paradigms can help explain the biogeography and local distribution patterns of different taxonomic and functional groups of inselberg species.

Island biogeography theory and the habitat heterogeneity jointly explain global patterns of Rhododendron diversity.

Mountain biodiversity is of great importance to biogeography and ecology. However, it is unclear what ecological and evolutionary processes best explain the generation and maintenance of its high levels of species diversity. In this study, we determined which of six common hypotheses (e.g., climate hypotheses, habitat heterogeneity hypothesis and island biogeography theory) best explain global patterns of species diversity in Rhododendron. We found that Rhododendron diversity patterns were most strongly explained by proxies of island biogeography theory (i.e., mountain area) and habitat heterogeneity (i.e., elevation range). When we examined other relationships important to island biogeography theory, we found that the planimetric area and the volume of mountains were positively correlated with the Rhododendron diversity, whereas the 'mountains-to-mainland' distance was negatively correlated with Rhododendron diversity and shared species. Our findings demonstrate that Rhododendron diversity can be explained by island biogeography theory and habitat heterogeneity, and mountains can be regarded as islands which supported island biogeography theory.

Synergistic effects of Pleistocene geological and climatic events on complex phylogeographic history of widespread sympatric species of Megaloptera in East Asia.

Unraveling the phylogeographic histories of species remains a key endeavor for comprehending the evolutionary processes contributing to the rich biodiversity and high endemism found in East Asia. In this study, we explored the phylogeographic patterns and demographic histories of three endemic fishfly and dobsonfly species ( Neochauliodes formosanus, Protohermes costalis, and Neoneuromus orientalis) belonging to the holometabolan order Megaloptera. These species, which share a broad and largely overlapping distribution, were analyzed using comprehensive mitogenomic data. Our findings revealed a consistent influence of vicariance on the population isolation of Neoc. formosanus and P. costalis between Hainan, Taiwan, and the East Asian mainland during the early Pleistocene, potentially hindering subsequent colonization of the later diverged Neon. orientalis to these islands. Additionally, we unveiled the dual function of the major mountain ranges in East Asia, serving both as barriers and conduits, in shaping the population structure of all three species. Notably, we demonstrated that these co-distributed species originated from Southwest, Southern, and eastern Central China, respectively, then subsequently migrated along multi-directional routes, leading to their sympatric distribution on the East Asian mainland. Furthermore, our results highlighted the significance of Pleistocene land bridges along the eastern coast of East Asia in facilitating the dispersal of mountain-dwelling insects with low dispersal ability. Overall, this study provides novel insight into the synergistic impact of Pleistocene geological and climatic events in shaping the diversity and distribution of aquatic insects in East Asia.

Landscape, Human Disturbance, and Climate Factors Drive the Species Richness of Alien Invasive Plants on Subtropical Islands.

Invasive alien plants (IAPs) pose a significant threat to island biodiversity and severely impact ecosystems. Understanding the species-area relationship and environmental determinants of growth forms for IAP species on subtropical islands is crucial for establishing an IAP's early warning mechanism, enhancing island ecological management, and protecting the ecosystems of Fujian and other subtropical islands. The study identified significant species-area relationships for IAPs and different life-form plants (trees, shrubs, and herbs), with slopes of 0.27, 0.16, 0.15, and 0.24, respectively. The small island effect does not apply to all species. Isolation has little effect on species richness, and the IAPs on Fujian islands do not conform to the isolation effect in island biogeography. Landscape factors are the main determinants of IAPs and different life-form species richness, with area, shape index, and perimeter-area ratio being the three primary landscape factors. These environmental factors are closely related to habitat heterogeneity. Besides landscape factors, different life forms respond differently to environmental factors. Climate drives the species richness distribution of shrubs and herbs, while trees are mainly influenced by human activities. Overall, landscape, human disturbance, and climate jointly drive the distribution of IAPs, with landscape factors being the most significant.

Diversity of Ladybird Beetles (Coleoptera: Coccinellidae) in Tenerife and La Gomera (Canary Islands): The Role of Size and Other Island Characteristics.

This paper provides new data on the ladybird beetles (Coccinellidae) from two islands in the Canary archipelago: Tenerife, the largest island, and La Gomera, the second smallest. As they clearly differ in size but are similar in location and geological age, they are a suitable model for testing the species-area relationship. Our study shows that, in line with this main assumption of the theory of island biogeography, clearly more species occur on a large island (Tenerife) than on a small one (La Gomera). The field surveys documented the occurrence of 35 ladybird species on Tenerife (including 5 not previously reported from this island) and of 20 species on La Gomera (2 species new to the island). Coelopterus sp. collected on Tenerife (a single female that could not be identified to species) is the first record of this genus for the whole Canary Islands. Taking our data and previously published records into account, 47 species of Coccinellidae are known to occur on Tenerife and 26 species on La Gomera. Tenerife has by far the richest ladybird fauna of all the Canary Islands (the next in line, Gran Canaria, has 41 recorded species), but it also has the highest number of non-native ladybird species. All of the ten non-native species recorded in the Canary Islands are found on Tenerife, and for most of them, Tenerife was the island of their first appearance in the archipelago. This island, much more distant from the mainland than the other relatively large islands (Fuerteventura, Lanzarote), appears to be the main recipient of ladybirds immigrating to the Canary Islands. Tenerife can play this role probably because of its great habitat diversity and altitude variation, as well as intensive tourism and trade-related transport.

Diversity in Irish and British avifauna assemblages: What can variation in diversity profiles reveal about the forces that drive assemblage composition and structure?

Ireland and Britain are two islands located at Europe's westernmost edge, both of which act as the final breeding outposts for many bird species within their European ranges. Despite their similar geographic locations and geological histories, Ireland and Britain host different breeding avifauna assemblages. Diversity profiles, which can serve as more robust alternatives to classic diversity indices, were employed in this study to explore disparities in the two islands' breeding avifauna assemblages. Variations in assemblages were explored, along with their potential drivers, through analyses at three levels: island-scale breeding bird assemblage compositions, island-scale diversity profiles considering 49 common breeding species, and habitat-specific diversity profiles considering assemblages in east/central Irish farmland and East Anglian farmland. Analysis of the two islands' breeding avifauna assemblages revealed that the Irish assemblage is a complete subset of the British assemblage. Analyses of Irish and British assemblages at both an island scale and a habitat scale revealed patterns linking land use to trends within the two islands' avifauna assemblages. Irish assemblages contained greater proportions of insectivorous farmland species by abundance, while British assemblages contained greater proportions of seed-eating farmland species; both trends appeared to be related to structural differences in agricultural land use on the two islands. The British and East Anglian assemblages exhibited higher diversity across all analyses, which appeared to be driven by the assemblages' higher relative abundances of species that were most genetically distinct. This study highlights the ability of diversity profiles to impart more information than classic diversity indices by incorporating species similarity data.

Waves of Colonization and Gene Flow in a Great Speciator.

Secondary contact between previously allopatric lineages offers a test of reproductive isolating mechanisms that may have accrued in isolation. Such instances of contact can produce stable hybrid zones-where reproductive isolation can further develop via reinforcement or phenotypic displacement-or result in the lineages merging. Ongoing secondary contact is most visible in continental systems, where steady input from parental taxa can occur readily. In oceanic island systems, however, secondary contact between closely related species of birds is relatively rare. When observed on sufficiently small islands, relative to population size, secondary contact likely represents a recent phenomenon. Here, we examine the dynamics of a group of birds whose apparent widespread hybridization influenced Ernst Mayr's foundational work on allopatric speciation: the whistlers of Fiji (Aves: Pachycephala). We demonstrate two clear instances of secondary contact within the Fijian archipelago, one resulting in a hybrid zone on a larger island, and the other resulting in a wholly admixed population on a smaller, adjacent island. We leveraged low genome-wide divergence in the hybrid zone to pinpoint a single genomic region associated with observed phenotypic differences. We use genomic data to present a new hypothesis that emphasizes rapid plumage evolution and post-divergence gene flow.

Genomics of ecological adaptation in Canary Island Descurainia (Brassicaceae) and comparisons with other Brassicaceae.

Oceanic archipelagos provide striking examples of lineages that have radiated over pronounced ecological gradients. Accompanying this diversification, lineages have evolved adaptations allowing survival in extreme environments. Here, we investigate the genomic basis of ecological adaptation in Canary Island Descurainia (Brassicaceae), an island relative of Arabidopsis. The seven endemic species have diversified in situ along an elevational and ecological gradient, from low-elevation scrub to high-elevation sub-alpine desert. We first generated a reference genome for Descurainia millefolia, phylogenetic analysis of which placed it as sister to D. sophioides. Ninety-six gene families were found to be specific to D. millefolia and a further 1087 and 1469 gene families have expanded or contracted in size, respectively, along the D. millefolia branch. We then employed genome re-sequencing to sample 14 genomes across the seven species of Canary Island Descurainia and an outgroup. Phylogenomic analyses were consistent with previous reconstructions of Canary Island Descurainia in resolving low- and high-elevation clades. Using the branch-site dN/dS method, we detected positive selection for 275 genes on the branch separating the low- and high-elevation species and these positively selected genes (PSGs) were significantly enriched for functions related to reproduction and stress tolerance. Comparing PSGs to those in analyses of adaptation to elevation and/or latitude in other Brassicaceae, we found little evidence of widespread convergence and gene reuse, except for two examples, one of which was a significant overlap between Descurainia and Draba nivalis, a species restricted to high latitudes. The study of Canary Island Descurainia suggests that the transition to high-elevation environments such as that found in the high mountains of the Canary Islands involves selection on genes related to reproduction and stress tolerance but that repeated evolution across different lineages that have evolved into similar habitats is limited, indicating substantially different molecular trajectories to adaptation.

Chromosome-Level Reference Genome of the Ponza Grayling (Hipparchia sbordonii), an Italian Endemic and Endangered Butterfly.

Islands are crucial evolutionary hotspots, providing unique opportunities for differentiation of novel biodiversity and long-term segregation of endemic species. Islands are also fragile ecosystems, where biodiversity is more exposed to environmental and anthropogenic pressures than on continents. The Ponza grayling, Hipparchia sbordonii, is an endemic butterfly species that is currently found only in two tiny islands of the Pontine archipelago, off the coast of Italy, occupying an area smaller than 10 km2. It has been classified as Endangered (IUCN) because of the extremely limited area of occurrence, population fragmentation, and the recent demographic decline. Thanks to a combination of different assemblers of long and short genomic reads, bulk transcriptome RNAseq, and synteny analysis with phylogenetically close butterflies, we produced a highly contiguous, chromosome-scale annotated reference genome for the Ponza grayling, including 28 autosomes and the Z sexual chromosomes. The final assembly spanned 388.61 Gb with a contig N50 of 14.5 Mb and a BUSCO completeness score of 98.5%. Synteny analysis using four other butterfly species revealed high collinearity with Hipparchia semele and highlighted 10 intrachromosomal inversions longer than 10 kb, of which two appeared on the lineage leading to H. sbordonii. Our results show that a chromosome-scale reference genome is attainable also when chromatin conformation data may be impractical or present specific technical challenges. The high-quality genomic resource for H. sbordonii opens up new opportunities for the accurate assessment of genetic diversity and genetic load and for the investigations of the genomic novelties characterizing the evolutionary path of this endemic island species.

Amphibian survey of Ko Pha-gnan in Surat Thani Province, Thailand.

Insular amphibian species are often overlooked, rendering them vulnerable to habitat encroachment and other anthropogenic threats. The aim of this study was to compile a comprehensive list of amphibian species on Ko Pha-ngan in Surat Thani Province, Thailand. Data were collected via transect surveys and drift line fence traps in three different habitat types from February 2021 to September 2023. Our efforts detected 12 unique amphibian species in each of the three habitat types. The highest number of detections was observed in the Ko Pha-ngan-Than Sadet National Park protected areas. The common tree frog (Polypedatesleucomystax) and the common Asian toad (Duttaphrynusmelanostictus) were the two most abundantly found species on the island, whereas the Koh Tao caecilian (Ichthyophiskohtaoensis) and the newly described false Doria's fanged frog (Limnonectespseudodoriae) where the least commonly found species. In addition, Microhylaheymonsi and Fejervaryalimnocharis tadpoles were observed developing in high-salinity water bodies. Many species have shown a high tolerance in human-dominated landscapes. This study sheds light on the need for additional monitoring to better understand the dynamics of endemic species in addition to the impact tourism-driven development and habitat destruction has on a species with an insularly finite habitat.

Proximity among protected area networks promotes functional connectivity for wintering waterfowl.

The equilibrium theorem provided a fundamental framework for understanding species' distributions and movement in fragmented ecosystems. Wetland-dependent avian species are model organisms to test insular predictions within protected area networks because their mobility allows surveillance of isolated patches without landscape barriers. We hypothesized size and isolation would influence functional connectivity of sanctuaries by GPS-marked wintering mallards (Anas platyrhynchos) within a mesocosm protected sanctuary area network. We evaluated functional connectivity and sanctuary use, measured by movements between sanctuaries, using a multistate modeling framework. Proximity drove connectivity, underscoring that patch isolation-not size-influenced connectivity, even for an avian species with no ascertainable landscape resistance or barriers. We also found that sanctuary use increased overwintering survival by reducing harvest mortality. Our test of equilibrium theory predictions demonstrated that isolation of protected sanctuary areas supersedes their size in determining functional connectivity for mallards and access to these areas may have direct fitness consequences. Our findings could refine land acquisition, restoration, and management practices with equal or greater emphasis on adjacency in protected area network design, especially for wetland-dependent migratory gamebirds.

Landscape configuration can flip species-area relationships in dynamic meta-food-webs.

Spatial and trophic processes profoundly influence biodiversity, yet ecological theories often treat them independently. The theory of island biogeography and related theories on metacommunities predict higher species richness with increasing area across islands or habitat patches. In contrast, food-web theory explores the effects of traits and network structure on coexistence within local communities. Exploring the mechanisms by which landscape configurations interact with food-web dynamics in shaping metacommunities is important for our understanding of biodiversity. Here, we use a meta-food-web model to explore the role of landscape configuration in determining species richness and show that when habitat patches are interconnected by dispersal, more species can persist on smaller islands than predicted by classical theory. When patch sizes are spatially aggregated, this effect flattens the slope of the species-area relationship. Surprisingly, when landscapes have random patch-size distributions, the slope of the species-area relationships can even flip and become negative. This could be explained by higher biomass densities of lower trophic levels that then support species occupying higher trophic levels, which only persist on small and well-connected patches. This highlights the importance of simultaneously considering landscape configuration and local food-web dynamics to understand drivers of species-area relationships in metacommunities.This article is part of the theme issue 'Diversity-dependence of dispersal: interspecific interactions determine spatial dynamics'.

Half of global islands have reached critical area thresholds for undergoing rapid increases in biological invasions.

Biological invasions are among the threats to global biodiversity and social sustainability, especially on islands. Identifying the threshold of area at which non-native species begin to increase abruptly is crucial for early prevention strategies. The small-island effect (SIE) was proposed to quantify the nonlinear relationship between native species richness and area but has not yet been applied to non-native species and thus to predict the key breakpoints at which established non-native species start to increase rapidly. Based on an extensive global dataset, including 769 species of non-native birds, mammals, amphibians and reptiles established on 4277 islands across 54 archipelagos, we detected a high prevalence of SIEs across 66.7% of archipelagos. Approximately 50% of islands have reached the threshold area and thus may be undergoing a rapid increase in biological invasions. SIEs were more likely to occur in those archipelagos with more non-native species introduction events, more established historical non-native species, lower habitat diversity and larger archipelago area range. Our findings may have important implications not only for targeted surveillance of biological invasions on global islands but also for predicting the responses of both non-native and native species to ongoing habitat fragmentation under sustained land-use modification and climate change.

eDNA Metabarcoding Reveals the Species-Area Relationship of Amphibians on the Zhoushan Archipelago.

The species-area relationship is important for understanding species diversity patterns at spatial scales, but few studies have examined the relationship using environmental DNA (eDNA) techniques. We investigated amphibian diversity on 21 islands of the Zhoushan Archipelago and nearby mainland areas in China using the combination of eDNA metabarcoding and the traditional line transect method (TLTM) and identified the species-area relationship for amphibians on the islands. The mean detection probability of eDNA is 0.54, while the mean detection probability of TLTM is 0.24. The eDNA metabarcoding detected eight amphibian species on the islands and nine species in the mainland areas, compared with seven species on the islands and nine species in the mainland areas that were identified by TLTM. Amphibian richness on the islands increased with island area and habitat diversity. The species-area relationship for amphibians in the archipelago was formulated as the power function (S = 0.47A0.21) or exponential function (S = 2.59 + 2.41 (logA)). Our results suggested that eDNA metabarcoding is more sensitive for the detection of amphibian species. The combined use of eDNA metabarcoding and the traditional line transect method may optimize the survey results for amphibians.

Genomic SNPs resolve the phylogeny of an ancient amphibian island radiation from the Seychelles.

Unusually for oceanic islands, the granitic Seychelles host multiple lineages of endemic amphibians. This includes an ancient (likely ca. 60 million years) radiation of eight caecilian species, most of which occur on multiple islands.These caecilians have a complicated taxonomic history and their phylogenetic inter-species relationships have been difficult to resolve. Double-digest RAD sequencing (ddRADseq) has been applied extensively to phylogeography and increasingly to phylogenetics but its utility for resolving ancient divergences is less well established. To address this, we applied ddRADseq to generate a genome-wide SNP panel for phylogenomic analyses of the Seychelles caecilians, whose phylogeny has so far not been satisfactorily resolved with traditional DNA markers. Based on 129,154 SNPs, we resolved deep and shallow splits, with strong support. Our findings demonstrate the capability of genome-wide SNPs for evolutionary inference at multiple taxonomic levels and support the recently proposed synonymy of Grandisonia Taylor, 1968 with Hypogeophis Peters, 1879. We revealed three clades of Hypogeophis (large-, medium- and short-bodied) and identify a single origin of the diminutive, stocky-bodied and pointy-snouted phenotype.

Island biogeography theory provides a plausible explanation for why larger vertebrates and taller humans have more diverse gut microbiomes.

Prior work has shown a positive scaling relationship between vertebrate body size, human height, and gut microbiome alpha diversity. This observation mirrors commonly observed species area relationships (SARs) in many other ecosystems. Here, we expand these observations to several large datasets, showing that this size-diversity scaling relationship is independent of relevant covariates, like diet, body mass index, age, sex, bowel movement frequency, antibiotic usage, and cardiometabolic health markers. Island biogeography theory (IBT), which predicts that larger islands tend to harbor greater species diversity through neutral demographic processes, provides a simple mechanism for positive SARs. Using a gut-adapted IBT model, we demonstrated that increasing the length of a flow-through ecosystem led to increased species diversity, closely matching our empirical observations. We delve into the possible clinical implications of these SARs in the American Gut cohort. Consistent with prior observations that lower alpha diversity is a risk factor for Clostridioides difficile infection (CDI), we found that individuals who reported a history of CDI were shorter than those who did not and that this relationship was mediated by alpha diversity. We observed that vegetable consumption had a much stronger association with CDI history, which was also partially mediated by alpha diversity. In summary, we find that the positive scaling observed between body size and gut alpha diversity can be plausibly explained by a gut-adapted IBT model, may be related to CDI risk, and vegetable intake appears to independently mitigate this risk, although additional work is needed to validate the potential disease risk implications.

In defence of the entity of Macaronesia as a biogeographical region.

Since its coinage ca. 1850 AD by Philip Barker Webb, the biogeographical region of Macaronesia, consisting of the North Atlantic volcanic archipelagos of the Azores, Madeira with the tiny Selvagens, the Canaries and Cabo Verde, and for some authors different continental coastal strips, has been under dispute. Herein, after a brief introduction on the terminology and purpose of regionalism, we recover the origins of the Macaronesia name, concept and geographical adscription, as well as its biogeographical implications and how different authors have positioned themselves, using distinct terrestrial or marine floristic and/or faunistic taxa distributions and relationships for accepting or rejecting the existence of this biogeographical region. Four main issues related to Macaronesia are thoroughly discussed: (i) its independence from the Mediterranean phytogeographical region; (ii) discrepancies according to different taxa analysed; (iii) its geographical limits and the role of the continental enclave(s), and, (iv) the validity of the phytogeographical region level. We conclude that Macaronesia has its own identity and a sound phytogeographical foundation, and that this is mainly based on three different floristic components that are shared by the Macaronesian core (Madeira and the Canaries) and the outermost archipelagos (Azores and Cabo Verde). These floristic components are: (i) the Palaeotropical-Tethyan Geoflora, formerly much more widely distributed in Europe and North Africa and currently restricted to the three northern archipelagos (the Azores, Madeira and the Canaries); (ii) the African Rand Flora, still extant in the coastal margins of Africa and Arabia, and present in the southern archipelagos (Madeira, the Canaries and Cabo Verde), and (iii) the Macaronesian neoendemic floristic component, represented in all the archipelagos, a result of allopatric diversification promoted by isolation of Mediterranean ancestors that manage to colonize Central Macaronesia and, from there, the outer archipelagos. Finally, a differentiating floristic component recently colonized the different archipelagos from the nearest continental coast, providing them with different biogeographic flavours.

A niche-based theory of island biogeography.

The equilibrium theory of island biogeography (ETIB) is a widely applied dynamic theory proposed in the 1960s to explain why islands have coherent differences in species richness. The development of the ETIB was temporarily challenged in the 1970s by the alternative static theory of ecological impoverishment (TEI). The TEI suggests that the number of species on an island is determined by its number of habitats or niches but, with no clear evidence relating species richness to the number of niches however, the TEI has been almost dismissed as a theory in favour of the original ETIB. Here, we show that the number of climatic niches on islands is an important predictor of the species richness of plants, herpetofauna and land birds. We therefore propose a model called the niche-based theory of island biogeography (NTIB), based on the MacroEcological Theory on the Arrangement of Life (METAL), which successfully integrates the number of niches sensu Hutchinson into ETIB. To account for greater species turnover at the beginning of colonisation, we include higher initial extinction rates. When we test our NTIB for resident land birds in the Krakatau Islands, it reveals a good correspondence with observed species richness, immigration and extinction rates. Provided the environmental regime remains unchanged, we estimate that the current species richness at equilibrium is ~45 species (range between 38.39 and 61.51). Our NTIB provides better prediction because it counts for changes in species richness with latitude, which is not considered in any theory of island biogeography.

Population structure in Mixornis tit-babblers across Sunda Shelf matches interfluvia of paleo-rivers.

Rivers constitute an important biogeographic divide in vast areas of tropical rainforest, such as the Amazon and Congo Basins. Southeast Asia's rainforests are currently fragmented across islands divided by sea, which has long obscured their extensive history of terrestrial connectivity as part of a vast (but now submerged) subcontinent - Sundaland - during most of the Quaternary. The role of paleo-rivers in determining population structure in Sundaic rainforests at a time when these forests were connected remains little understood. We examined the coloration of museum skins and used the genomic DNA of museum samples and freshly-collected blood tissue of a pair of Sundaic songbird species, the pin-striped and bold-striped tit-babblers (Mixornis gularis and M. bornensis, respectively), to assess the genetic affinity of populations on small Sundaic islands that have largely been ignored by modern research. Our genomic and morphological results place the populations from the Anambas and Natuna Islands firmly within M. gularis from the Malay Peninsula in western Sundaland, even though some of these islands are geographically much closer to Borneo, where M. bornensis resides. Our results reveal genetic structure consistent with the course of Sundaic paleo-rivers and the location of the interfluvia they formed, and add to a small but growing body of evidence that rivers would have been of equal biogeographic importance in Sundaland's former connected forest landscape as they are in Amazonia and the Congo Basin today.

The importance of migratory drop-off for island colonization in birds.

Seasonal migration is an underappreciated driver of animal diversification. Changes in migratory behaviour may favour the establishment of sedentary founder populations and promote speciation if there is sufficient reproductive isolation between sedentary and migratory populations. From a systematic literature review, we here quantify the role of migratory drop-off-the loss of migratory behaviour-in promoting speciation in birds on islands. We identify at least 157 independent colonization events likely initiated by migratory species that led to speciation, including 44 cases among recently extinct species. By comparing, for all islands, the proportion of island endemic species that derived from migratory drop-off with the proportion of migratory species among potential colonizers, we showed that seasonal migration has a larger effect on island endemic richness than direct dispersal. We also found that the role of migration in island colonization increases with the geographic isolation of islands. Furthermore, the success of speciation events depends in part on species biogeographic and ecological factors, here positively associated with greater range size and larger flock sizes. These results highlight the importance of shifts in migratory behaviour in the speciation process and calls for greater consideration of migratory drop-off in the biogeographic distribution of birds.