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1.
Summary We examined the potential effects of geography on the distribution and speciation of skinks on tropical Pacific archipelagos. The entire tropical Pacific skink fauna was divided into continental (found also in continental areas), Pacific (endemic to the study area but found within more than one archipelago) and endemic (found within only one archipelago) species categories. The number and proportion of skinks within each species category were determined for each of the 27 archipelagos in the study area. Nine geographic variables reflecting archipelago size, isolation and elevation were estimated for each archipelago. Principal components analysis was used to reduce the nine variables to three uncorrelated composite variables that were interpreted as representing archipelago size, isolation and elevation. Numbers and proportions of skinks in each category within an archipelago were related to the composite geographic variables using multiple linear regression analysis. Archipelago size and isolation were important predictors of both skink diversity and endemism. Results were then compared to diversity and endemism of birds within the study area. Skinks showed an archipelago-wide level of endemism similar to that of birds. On an archipelago by archipelago basis, however, large differences between birds and skinks were evident. In particular, the New Caledonia skink fauna was much more endemic than that of birds. The bird faunas of Hawaii and the Marquesas were nearly completely endemic, while no endemic skinks occurred in these two archipelagos. These differences presumably reflect the relative dispersal powers of skinks and birds and, consequently, rates of colonization and speciation. Differences may also be due partly to morphological conservatism among isolated skink populations and the occurrence of cryptic species that have not yet been identified as separate species. The discovery of such cryptic species, however, is unlikely to increase the endemic skink fauna of Hawaii and other distant archipelagos to a level commensurate with that of birds. Differences in endemism between skinks and birds may also be due to unknown local ecological interactions.  相似文献   

2.
Biogeography of mammals on tropical Pacific islands   总被引:2,自引:0,他引:2  
Aim We examine the influence of geography on species richness and endemism of mammals on tropical Pacific archipelagos to determine the importance of intra‐ and inter‐archipelago speciation in promoting local and regional species richness. Location Thirty tropical Pacific archipelagos. Methods A distributional list of mammals on 30 archipelagos was compiled, and values for 10 geographical variables were estimated for each archipelago. Mammal species were placed in three different categories (continental, Pacific and endemic) based on their distribution. The total number of species and numbers of species within each category were related to the geographical variables using Poisson regression analysis. Results Species richness was related positively to variables describing land area, numbers of large islands and elevation; and negatively to variables describing isolation. Levels of endemism did not differ between volant and non‐volant species, but differed between mega‐ and microchiropterans. Main conclusions Variation in species richness of mammals in the tropical Pacific region can be accounted for by a combination of intra‐archipelago speciation within archipelagos composed of large islands, and inter‐archipelago speciation, particularly among more isolated archipelagos. Mammals were less widely distributed throughout the study area than previously found for butterflies, skinks or birds. However, the level of endemism was similar to that of skinks and birds on the same archipelagos, and was higher than that of butterflies.  相似文献   

3.
Summary I analysed avifaunal data from 30 archipelagos and isolated islands in the tropical Pacific Ocean to examine the effects of geography on endemism. I divided the total bird species list (pelagic and migrant species excluded) for each island group into continental (also found outside of the study area), Pacific (found only within the study area but within more than one archipelago), and endemic (found only within a single archipelago) species and estimated ten variables related to the geography of each archipelago. I used multiple linear regression analysis to relate numbers and proportions of species in each category to the geographical variables. Total land area of an archipelago was the most improtant variable in explaining variation in the number of species in each category, with elevation and isolation also being important. The relationships between the proportions of species in each category and the geographical variables underscore the importance of isolation and the number of large islands in promoting endemism, presumably by allowing both inter- and intra-archipelagal speciation to proceed.  相似文献   

4.
One island group suggested to be an exception to the species–area Relationship is the Vanuatu Archipelago, a group of 13 large and 80 small islands in the southwest Pacific Ocean. To test the hypothesis that the lizard fauna of the Vanuatu Archipelago does not meet the predictions of the species-area relationship, and thus is depauperate, we compare diversity among several island groups in the southwest Pacific: Fiji, the Loyalty Islands, New Caledonia, Samoa, the Solomon Islands, Tonga, and Vanuatu. We found that the lizard diversity of Vanuatu meets the pattern of diversity predicted by the species-area relationship. The Solomon Islands, the largest and least isolated oceanic archipelago considered, has the greatest species diversity and endemism of the oceanic islands. Inclusion or exclusion of island groups based on factors such as geologic history or faunal source affects the strength of the relationship between diversity, area, and history of emergence, and influences perceptions of diversity within individual archipelagos. In addition to island size, factors such evolutionary time scale, speciation, and archipelago complexity influence species richness on islands.  相似文献   

5.
Islands are particularly suited to testing hypotheses about the ecological and evolutionary mechanisms underpinning community assembly. Yet the complex spatial arrangements of real island systems have received little attention from both empirical studies and theoretical models. Here, we investigate the extent to which the spatial structure of archipelagos affects species diversity and endemism. We start by proposing a new spatially structured neutral model that explicitly considers archipelago structure, and then investigate its predictions under a diversity of scenarios. Our results suggest that considering the spatial structure of archipelagos is crucial to understanding their diversity and endemism, with structured island systems acting both as “museums” and “cradles” of biodiversity. These dynamics of diversification may change the traditionally expected pattern of decrease in species richness with distance from the mainland, even potentially leading to increasing patterns for taxa with high speciation rates in archipelagos off species‐poor continental areas. Our results also predict that, within spatially structured archipelagos, metapopulation dynamics and evolutionary processes can generate higher diversity on islands more centrally placed than at the periphery. We derive from our results a set of theoretical predictions, potentially testable with empirical data.  相似文献   

6.
Aim To establish the extent to which archipelagos follow the same species–area relationship as their constituent islands and to explore the factors that may explain departures from the relationship. Location Thirty‐eight archipelagos distributed worldwide. Methods We used ninety‐seven published datasets to create island species–area relationships (ISARs) using the Arrhenius logarithmic form of the power model. Observed and predicted species richness of an archipelago and of each of its islands were used to calculate two indices that determined whether the archipelago followed the ISAR. Archipelagic residuals (ArcRes) were calculated as the residual of the prediction provided by the ISAR using the total area of the archipelago, standardized by the total richness observed in the archipelago. We also tested whether any characteristic of the archipelago (geological origin and isolation) and/or taxon accounts for whether an archipelago fits into the ISAR or not. Finally, we explored the relationship between ArcRes and two metrics of nestedness. Results The archipelago was close to the ISAR of its constituent islands in most of the cases analysed. Exceptions arose for archipelagos where (i) the slopes of the ISAR are low, (ii) observed species richness is higher than expected by the ISAR and/or (iii) distance to the mainland is small. The archipelago's geological origin was also important; a higher percentage of oceanic archipelagos fit into their ISAR than continental ones. ArcRes indicated that the ISAR underpredicts archipelagic richness in the least isolated archipelagos. Different types of taxon showed no differences in ArcRes. Nestedness and ArcRes appear to be related, although the form of the relationship varies between metrics. Main conclusions Archipelagos, as a rule, follow the same ISAR as their constituent islands. Therefore, they can be used as distinct units themselves in large‐scale biogeographical and macroecological studies. Departure from the ISAR can be used as a crude indicator of richness‐ordered nestedness, responsive to factors such as isolation, environmental heterogeneity, number and age of islands.  相似文献   

7.
Distributions of danaine butterfly species and associated mimetic patterns were compared among fifteen archipelagos of the tropical Pacific Ocean, and within five major archipelagos (the Bismarcks, Fiji, East and West Solomon Islands, and Vanuatu). Using both simple and stepwise linear regression analysis, variation in the total number of danaine species and number of mimetic patterns was assessed with respect to island size, isolation and elevation. Relative to interarchipelago distributions, the distribution of danaine species and number of mimetic patterns on islands within archipelagos exhibited less dependence upon interisland distance and island area. Geographical features influencing the number of mimetic patterns were similar to those of danaines as a whole. Analysis of residuals from stepwise linear regression suggested that factors influencing danaine distributions were different from those for non-danaine butterflies. This result is consistent with the hypothesis of enhancement of danaine species establishment through Müllerian mimicry, although other factors such as host plant availability and similar habitat use may also be important.  相似文献   

8.
Aim We use molecular‐based phylogenetic methods and ancestral area reconstructions to examine the systematic relationships and biogeographical history of the Indo‐Pacific passerine bird family Pachycephalidae (whistlers). Analysed within an explicit spatiotemporal framework, we elucidate distinct patterns of diversification across the Melanesian and Indonesian archipelagos and explore whether these results may be explained by regional palaeogeological events. We further assess the significance of upstream colonization and its role in species accumulation within the region. Location The Indo‐Pacific region, with an emphasis on the archipelagos on either side of the Australo‐Papuan continent. Methods We used three nuclear and two mitochondrial markers to construct a molecular phylogenetic hypothesis of the Pachycephalidae by analysing 35 of the 49 species known to belong to the family. The programs diva and Mr Bayes were used to reconstruct ancestral area relationships and to examine biogeographical relationships across the family, and beast was implemented to assess the timing of dispersal events. Results We constructed a molecular phylogenetic hypothesis for the Pachycephalidae and estimated divergence times and ancestral area relationships. Different colonization patterns are apparent for the Pachycephalidae in the Indonesian and the Melanesian archipelagos. The Indonesian archipelago was colonized numerous times, whereas one or two colonizations of the Melanesian archipelagos account for the entire diversity of that region. After initial colonization of the Melanesian archipelagos some whistler species recolonized Australia and may have commenced a second round of colonization into Melanesia. Main conclusions The contrasting dispersal patterns of whistlers in archipelagos on either side of the Australo‐Papuan continent are congruent with the arrangement and history of islands in each of the regions and demonstrate that knowledge of palaeogeography is important for an understanding of evolutionary patterns in archipelagos. We also highlight that recolonization of continents from islands may be more common than has previously been assumed.  相似文献   

9.
Island biogeography has greatly contributed to our understanding of the processes determining species' distributions. Previous research has focused on the effects of island geography (i.e., island area, elevation, and isolation) and current climate as drivers of island species richness and endemism. Here, we evaluate the potential additional effects of historical climate on breeding land bird richness and endemism in Wallacea and the West Indies. Furthermore, on the basis of species distributions, we identify island biogeographical network roles and examine their association with geography, current and historical climate, and bird richness/endemism. We found that island geography, especially island area but also isolation and elevation, largely explained the variation in island species richness and endemism. Current and historical climate only added marginally to our understanding of the distribution of species on islands, and this was idiosyncratic to each archipelago. In the West Indies, endemic richness was slightly reduced on islands with historically unstable climates; weak support for the opposite was found in Wallacea. In both archipelagos, large islands with many endemics and situated far from other large islands had high importance for the linkage within modules, indicating that these islands potentially act as speciation pumps and source islands for surrounding smaller islands within the module and, thus, define the biogeographical modules. Large islands situated far from the mainland and/or with a high number of nonendemics acted as links between modules. Additionally, in Wallacea, but not in the West Indies, climatically unstable islands tended to interlink biogeographical modules. The weak and idiosyncratic effect of historical climate on island richness, endemism, and network roles indicates that historical climate had little effects on extinction‐immigration dynamics. This is in contrast to the strong effect of historical climate observed on the mainland, possibly because surrounding oceans buffer against strong climate oscillations and because geography is a strong determinant of island richness, endemism and network roles.  相似文献   

10.
Cluster analyses, Parsimony Analysis of Endemicity, species–area relationships, Mantel tests, partial Mantel tests and levels of endemism were used to investigate the influence of current geography and paleogeography on the composition of the fauna of the Tuscan Islands, a small archipelago in the Mediterranean. The composition of terrestrial mollusc and tenebrionid beetle faunas was prevalently influenced by Pleistocene conditions. The composition of butterfly (and possibly reptile) island fauna was influenced by both present (Holocene) and Pleistocene factors. Finally, chrysids were influenced only by present factors. On the whole, both present and historical factors concurred to determine present distributional patterns, but their relative influence varied according to the mobility of the group considered. Results provided here give some insights into the treatment of endemics to elucidate historical relationships among areas and add further momentum to a general model of biotic dynamics in landbridge archipelagos.  相似文献   

11.
Question: Can we recognize areas of high endemism and high endemic richness, using data from collections, and what are the ecological variables that best explain these areas? Location: Peninsula of Baja California, Mexico. Methods: We analysed the distribution of 723 endemic vascular plants species along the peninsula of Baja California and neighbouring islands distributed in 218 cartographic cells 15’ x 20’ in size. By means of a residual analysis, we identified areas of significantly high endemic species richness, and we calculated the degree of endemicity (or rarity) in each cell by giving to each species a weight factor inversely proportional to the land area it covers. Results: Nine regions of high‐endemicity and/or high endemic species richness were found. Discussion and conclusions: The analyses of rarity and endemic species richness showed two contrasting scenarios: High endemicity values in oceanic and sky islands accounts for a high number of species with a restricted distribution, promoted most likely by genetic isolation and high environmental heterogeneity. High endemic richness along the peninsular coast is related to ecotonal transition along vegetation types. After correcting for collection effort (i.e. the number of specimens collected within a cell), we found the phytogeographic region and altitudinal heterogeneity to be the variables that best predicted endemic richness. Both high endemism and high endemic richness have distinct geographic patterns within our study region. The nine endemic regions provide elements for priority definitions in future conservation programs.  相似文献   

12.
Satoyama, the traditional agricultural landscape in Japan, has drawn much attention from the viewpoint of biodiversity conservation. It is composed of diverse vegetation types, including secondary forests, paddy fields and cultivated fields in a narrow area (ca. 1 km2). To clarify the characteristics of butterfly assemblage and the relative contribution of each vegetation type to butterfly diversity and endemism in satoyama, we conducted a line-transect survey along a 1.1-km route with five sections (two forest interior, one interior-edge, one edge-openland and one openland). Life history features (voltinism, host plant range and host plant type) and endemism of butterflies were discussed in relation to their abundance and vegetation association. Fifty-one species and 856 individuals were recorded. Species richness was highest in the two sections with edges, 32 and 36 species, respectively. Analysis of vegetation association of each butterfly species showed that 16 species were forest interior species, 24 forest edge species and 11 openland species, indicating that vegetation diversity enhanced butterfly diversity. However, the point is that forest interior and edge species contained many specialists sensitive to human impact (univoltine and/or oligophagous species) and many species within temperate East Asia including all endemic species. In contrast, most openland species were generalists (multivoltine and/or polyphagous species) feeding on herbs/grasses with the widest geographic range. Since secondary forests kept more butterfly diversity and specialists than openlands, and had all endemic species, they must be maintained without over fragmentation in order to avoid loss of specialists and endemics in satoyama.  相似文献   

13.
Studies of fragmented habitat, such as island archipelagos, provide insights into the microevolutionary processes that drive early stages of diversification. Here, we examined genetic variation and gene flow among populations of the widespread buff‐banded rail Gallirallus philippensis in Oceania to understand the factors that promote speciation associated within this bird lineage. We analysed mtDNA Control Region sequences and six microsatellite loci from a total of 152 individuals of buff‐banded rail on islands and continental areas. We used a phylogeographic model‐testing approach and a structured spatial design ranging from within to among archipelagoes in the south Pacific. Buff‐banded rail populations in the Philippines archipelago and nearby Palau and Wallacea had high genetic diversity while those in geographically distant Australia showed lower variation. Other archipelagos sampled were found to have less genetic diversity and included haplotypes closely related to Wallacea (Bismarck, Vanuatu, New Caledonia) or Australia (New Zealand, Samoa, Fiji, Cocos Islands). Nucleotide diversity and allele frequency declined with degree of geographic isolation but haplotype diversity remained more even. However, both nucleotide and haplotype diversities were positively correlated with land area. Microsatellite data for a subset of locations showed moderate to high genetic differentiation and significant pairwise FST despite a relatively high migration rate. Our results are mostly consistent with a model of abrupt genetic changes due to founder events with multiple dispersals into Australia from Wallacea and Bismarck. Australia has probably been the source of birds for islands in the Pacific. This is shown by decreasing genetic diversity and growing genetic differentiation when distances separating populations increased from Australia. A history of range expansion and divergent natural selection may help explain the existence of numerous sympatric Gallirallus island endemics.  相似文献   

14.
We examine the effects of ecological opportunity and geographic area on rates of species accumulation and morphological evolution following archipelago colonization in day geckos (genus Phelsuma) in the Indian Ocean. Using a newly generated molecular phylogeny for the genus, we present evidence that these geckos likely originated on Madagascar, whereas colonization of three archipelagos in the Indian Ocean, the Seychelles, Mascarene, and Comoros Islands has produced three independent monophyletic radiations. We find that rates of species accumulation are not elevated following colonization but are roughly equivalent on all three isolated archipelagos and on the larger island of Madagascar. However, rates of species accumulation have slowed through time on Madagascar. Rates of morphological evolution are higher in both the Mascarene and Seychelles archipelagos compared to rates on Madagascar. This negative relationship between rate of morphological evolution and island area suggests that ecological opportunity is an important factor in diversification of day gecko species.  相似文献   

15.
The spatial patterns in the distributions of vertebrates in the rainforests of the wet tropics biogeographic region of north-eastern Australia were examined to form hypotheses on the processes that have shaped vertebrate assemblages and patterns of species richness and regional endemism. These rainforests occur in a relatively narrow and discontinuous strip along the coast of north-eastern Australia. We found that the number of regionally endemic species and the proportion of regional endemics present in each subregion are both strongly related to the geographic shape of subregional patches of rainforest, independent of rainforest area, within Australian tropical rainforests. Shape has a more significant influence on regional endemism than area, and area has a stronger influence on species richness. These patterns were congruent for all terrestrial vertebrate classes manuals, birds, reptiles and frogst, and for the four groups combined. Our results suggest that the combination of current rainforest area and shape are an index of the relative susceptibility of each area of rainforest to historical contractions, with the implication that historical habitat fluctuations, coupled with subsequent localized extinctions species sifting; have been extremely important processes in determining current patterns of endemism in Australia''s wet tropical rainforests. This hypothesis is supported by the highly nested structure of the subregional distribution patterns.  相似文献   

16.
Aim To evaluate the role of island isolation in explaining the distribution of vascular plant species in a dense freshwater archipelago, specifically comparing conventional measures of island isolation with landscape measures of island isolation. Location Data were collected from 35 islands within Massasauga Provincial Park on the eastern shores of the Georgian Bay, Ontario, Canada. Methods Sampled islands were located using stratified random selection based on location and size variation. The number of species was recorded along stratified random transects. Island isolation variables included distance to the mainland, distance to the nearest island, largest gap in a stepping‐stone sequence, distance to the closest upwind point of land, and a landscape measure of island isolation. The landscape measure of isolation was quantified as the percentage of the land area within 100, 250, 500, 1000, 1500 and 2000 m of each island’s perimeter. The isolation variables were calculated within a geographical information system (GIS). Dependent variables in the regression analyses included species richness, the logarithm of species richness and residuals of the species–area relationship. Independent variables included island isolation variables and their logarithmic transformations. Results Isolation plays a role, albeit small, in explaining species richness in the study area. In the regression analyses, the landscape measure of isolation provided a better fit than conventional measures of island isolation. Islands with less land than water within a 250‐m buffer were more effectively isolated and had fewer species present than islands surrounded by a greater proportion of water. Main conclusions Consistent with the species–isolation relationship, fewer species were present on more isolated islands within the Massasauga study area, as elucidated using a series of island buffers in a GIS. Applying a landscape measure of isolation to similar dense, freshwater archipelagos may elucidate species–isolation patterns not evident through conventional, straight‐line distance measurements of island isolation. The low value of the regression coefficients as well as the isolation history and high density of the Massasauga islands suggests caution in extending the results, especially to dissimilar archipelagos.  相似文献   

17.
Aim We used a phylogenetic framework to examine island colonization and predictions pertaining to differentiation within Macaronesian Tarphius (Insecta, Coleoptera, Zopheridae), and explain the paucity of endemics in the Azores compared with other Macaronesian archipelagos. Specifically, we test whether low diversity in the Azores could be due to recent colonization (phylogenetic lineage youth), cryptic speciation (distinct phylogenetic entities within species) or the young geological age of the archipelago. Location Macaronesian archipelagos (Azores, Madeira and the Canary Islands), northern Portugal and Morocco. Methods Phylogenetic analyses of mitochondrial and nuclear genes of Tarphius beetles of the Azores, other Macaronesian islands and neighbouring continental areas were used to investigate the origin of island biodiversity and to compare patterns of colonization and differentiation. A comparative nucleotide substitution rate test was used to select the appropriate substitution rate to infer clade divergence times. Results Madeiran and Canarian Tarphius species were found to be more closely related to each other, while Azorean taxa grouped separately. Azorean taxa showed concordance between species and phylogenetic clades, except for species that occur on multiple islands, which segregated by island of origin. Divergence time estimates revealed that Azorean Tarphius are an old group and that the most recent intra‐island speciation event on Santa Maria, the oldest island, occurred between 3.7 and 6.1 Ma. Main conclusions Our phylogenetic approach provides new evidence to understand the impoverishment of Azorean endemics: (1) Tarphius have had a long evolutionary history within the Azores, which does not support the hypothesis of fewer radiation events due to recent colonization; (2) the current taxonomy of Azorean Tarphius does not reflect common ancestry and cryptic speciation is responsible for the underestimation of endemics; (3) intra‐island differentiation in the Azores was found only in the oldest island, supporting the idea that young geological age of the archipelago limits the number of endemics; and (4) the lack of evidence for recent intra‐island diversification in Santa Maria could also explain the paucity of Azorean endemics. Phylogenetic reconstructions of other species‐rich taxa that occur on multiple Macaronesian archipelagos will reveal whether our conclusions are taxon specific, or of a more general nature.  相似文献   

18.
The general dynamic model of oceanic island biogeography describes the evolution of species diversity properties, including species richness (SR), through time. We investigate the hypothesis that SR in organisms with high dispersal capacities is better predicted by island area and elevation (as a surrogate of habitat diversity) than by time elapsed since island emergence and geographic isolation. Linear mixed effect models (LMMs) subjected to information theoretic model selection were employed to describe moss and liverwort SR patterns from 67 oceanic islands across 12 archipelagos. Random effects, which are used to modulate model parameters to take differences among archipelagos into account, included only a random intercept in the best‐fit model for liverworts and in one of the two best‐fit models for mosses. In this case, the other coefficients are constant across archipelagos, and we interpret the intercept as a measure of the intrinsic carrying capacity of islands within each archipelago, independently of their size, age, elevation and geographic isolation. The contribution of area and elevation to the models was substantially higher than that of time, with the least contribution made by measures of geographic isolation. This reinforces the idea that oceanic barriers are not a major impediment for migration in bryophytes and, together with the almost complete absence of in situ insular diversification, explains the comparatively limited importance of time in the models. We hence suggest that time per se has little independent role in explaining bryophyte SR and principally features as a variable accounting for the changing area and topographic complexity during the life‐cycle of oceanic islands. Simple area models reflecting habitat availability and diversity might hence prevail over more complex temporal models reflecting in‐situ speciation and dispersal (time, geographic connectivity) in explaining patterns of biodiversity for exceptionally mobile organisms.  相似文献   

19.
20.
Aim We compare the influence of contemporary geography and historical influences on butterfly diversity for islands in the Aegean archipelago. Location The Aegean archipelago (Greece) and two islands (Cyprus and Megisti) in the Levantine Sea. Methods Thirty‐one islands were examined. Data are taken from own surveys (Coutsis and Olivier) and from the literature. Stepwise multiple regression is used to determine relationships between species richness, frequency, rarity and endemicity against potential geographical predictors. Stepwise logit regression is used to determine geographical predictors of species incidence on islands. Inter‐island and inter‐species associations have been examined using multivariate ordination and clustering techniques. Results The Aegean butterfly fauna is characterized by decreasing diversity and rarity, and increasing homogeneity, from the periphery to the present geographical centre of the archipelago (Cyclades). Diversity and rarity are shown to relate closely to species richness, and species richness, in turn, is largely explained by contemporary geography, particularly the degree of isolation from the nearest mainland sources of Greece or Turkey, and island dimensions. Islands towards the centre of the archipelago are characterized by a group of mobile species (n ≥ 20 species) with extensive ranges across Europe; species that would have recolonized Santorini (Thira) following the VI6 eruption there c. 1630 bc . Endemic components, indicative of autochthonous evolutionary events, are few (5% of species are endemic) compared to known sedentary organisms (molluscs and isopods), but exceed those for more mobile animals (i.e. birds); their distribution is mainly confined to large isolated islands along the Aegean arc (i.e. Kriti) and in the Dodecanese group. Main conclusions Contemporary geography, i.e. processes currently operating in ecological time, dominates butterfly diversity gradients (species richness, frequency, rarity and incidence) in the archipelago. Two reasons are suggested to account for the lack of endemism and the pattern of decreasing diversity into the Cyclades. First, relict butterfly elements may have become extinct on all but a few larger islands, particularly from environmental changes since the Neolithic (fire and overgrazing). Second, colonization from the continental landmasses is ongoing with more mobile species transferring even to the most isolated islands.  相似文献   

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