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1.
The isolated archipelagos of Remote Oceania provide useful microcosms for understanding the impacts of initial human colonization. Palaeoecological data from most islands reveal catastrophic transformations, with losses of many species through over-hunting, deforestation and the introduction of novel mammalian predators, the most ubiquitous and devastating being commensal rats. Two case studies from the Austral Islands and New Zealand demonstrate the potential of direct human proxies from palaeoecological archives to detect initial human impacts on islands. We show how pollen from introduced crop plants, and buried seeds with gnaw marks from the introduced Pacific rat (Rattus exulans) provide a reliable means of detecting initial human colonization and highlight the downstream ecological consequences of agriculture and rat introduction on previously uninhabited pristine island ecosystems. Previous studies have relied on indirect signals of human arrival based on charcoal and associated vegetation changes, the causes of which are often more difficult to interpret with certainty.  相似文献   

2.
Speciation on islands, and particularly the divergence of species in situ, has long been debated. Here, we present one of the first, complete assessments of the geographic modes of speciation for the flora of a small oceanic island. Cocos Island (Costa Rica) is pristine; it is located 550 km off the Pacific coast of Central America. It harbors 189 native plant species, 33 of which are endemic. Using phylogenetic data from insular and mainland congeneric species, we show that all of the endemic species are derived from independent colonization events rather than in situ speciation. This is in sharp contrast to the results of a study carried out in a comparable system, Lord Howe Island (Australia), where as much as 8.2% of the plant species were the product of sympatric speciation. Differences in physiography and age between the islands may be responsible for the contrasting patterns of speciation observed. Importantly, comparing phylogenetic assessments of the modes of speciation with taxonomy‐based measures shows that widely used island biogeography approaches overestimate rates of in situ speciation.  相似文献   

3.
Here, we investigate Mid- to Late-Holocene vegetation changes in low-lying coastal areas in Tonga and how changing sea levels and recurrent volcanic eruptions have influenced vegetation dynamics on four islands of the Tongan archipelago (South Pacific). To investigate past vegetation and environmental change at Ngofe Marsh (‘Uta Vava’u), we examined palynomorphs (pollen and spores), charcoal (fire), and sediment characteristics (volcanic activity) from a 6.7-m-long sediment core. Radiocarbon dating indicated the sediments were deposited over the last 7700 years. We integrated the Ngofe Marsh data with similar previously published data from Avai’o’vuna Swamp on Pangaimotu Island, Lotofoa Swamp on Foa Island, and Finemui Swamp on Ha’afeva Island. Plant taxa were categorized as littoral, mangrove, rainforest, successional/ disturbance, and wetland groups, and linear models were used to examine relationships between vegetation, relative sea level change, and volcanic eruptions (tephra). We found that relative sea level change has impacted vegetation on three of the four islands investigated. Volcanic eruptions were not identified as a driver of vegetation change. Rainforest decline does not appear to be driven by sea level changes or volcanic eruptions. From all sites analyzed, vegetation at Finemui Swamp was most sensitive to changes in relative sea level. While vegetation on low-lying Pacific islands is sensitive to changing sea levels, island characteristics, such as area and elevation, are also likely to be important factors that mediate specific island responses to drivers of change.  相似文献   

4.
In the oceans of the tropical and warm-temperate zone (40° N–40° S), only a small number of islands are high enough to show timberline and alpine vegetation. Excluding large islands with a more continental climate, only the following oceanic islands are relevant: Pico (Azores), Madeira, Tenerife, Gran Canaria and La Palma (Canary islands), Fogo (Cape Verde islands), Fernando Poo (Bioko) and Tristan da Cunha in the Atlantic Ocean, Réunion and Grande Comore (Ngazidja) in the Indian Ocean, Yakushima (Japan), Maui and Hawaii (Hawaiian islands), and Mas Afuera (Juan Fernandez islands) in the Pacific Ocean. Timberline and alpine vegetation exist here under a unique combination of a highly oceanic climate and a marked geographic isolation which contrasts with the tropical alpine vegetation in the extended mountains of South America, Africa and Southeast Asia.This review seeks to identify common physiognomic patterns in the high elevation vegetation that exist despite the fact that the islands belong to different floristic regions of the world. Based on the existing literature as well as personal observation, an overview of the elevation, physiognomy and floristics of the forest (and tree) line and the alpine vegetation on 15 island peaks is given.The forest line ecosystems are dominated either by conifers (Canary islands, Yakushima), heath woodland (Azores, Madeira, Réunion, Grande Comore, Fernando Poo) or broad-leaved trees (Hawaiian islands, Juan Fernandez islands, Tristan da Cunha). In the subalpine and alpine belts, dry sclerophyllous scrub occurs on island mountains that are exposed to the trade winds (Canary islands, Cape Verde islands, Hawaiian islands, Réunion, Grande Comore). These peaks are more or less arid above the forest line because a temperature inversion restricts the rise of humid air masses further upslope. In the summit regions of the remaining islands, which are located either in the wet equatorial and monsoonal regions or in the temperate westerly zones without an effective inversion layer, mesic to wet vegetation types (such as grassland, alpine heathland and fern scrub) are found.Compared to mountains at a similar latitude in continental areas, the forest line on the islands is found at 1000 to 2000 m lower elevations. The paper discusses four factors that are thought to contribute to this forest line depression: (1) drought on trade-wind exposed island peaks with stable temperature inversions, (2) the absense of well-adapted high-altitude tree species on isolated islands, (3) immaturity of volcanic soils, and (4) an only small mountain mass effect that influences the vertical temperature gradient.  相似文献   

5.
Islands are bounded ecosystems and serve as excellent laboratories for assessing changes in Biodiversity. Some oceanic islands, such as Madagascar, Bermuda and notably the islands in the Pacific (e.g. Hawaii), are home to unique forms of endemic plants and animals that have evolved in isolation over millions of years. The palaeoecological record indicates that such islands are characterized by waves of extinctions concomitant with colonization by humans. By way of contrast, the biota of the islands of the north Atlantic (Greenland, Iceland and Faroe) do not follow the expected pattern and the few extinctions recorded are very localized. This is not a result of the scale of human impact, which is as great as on other islands, but relates to the virtual lack of endemics. The dearth of endemic forms and the disharmonic nature of these island communities indicates a youthful biota and the operation of severe filters and sweepstakes during colonisation over the last 10,000 years. This paper draws upon an extensive invertebrate fossil record to contrast and examine these spatial and temporal patterns in island Biodiversity.  相似文献   

6.
7.
Humans have introduced many species onto remote oceanic islands. The house mouse (Mus musculus) is a human commensal and has consequently been transported to oceanic islands around the globe as an accidental stowaway. The history of these introductions can tell us not only about the mice themselves but also about the people that transported them. Following a phylogeographic approach, we used mitochondrial D‐loop sequence variation (within an 849‐ to 864‐bp fragment) to study house mouse colonization of the Azores. A total of 239 sequences were obtained from all nine islands, and interpretation was helped by previously published Iberian sequences and 66 newly generated Spanish sequences. A Bayesian analysis revealed presence in the Azores of most of the D‐loop clades previously described in the domesticus subspecies of the house mouse, suggesting a complex colonization history of the archipelago as a whole from multiple geographical origins, but much less heterogeneity (often single colonization?) within islands. The expected historical link with mainland Portugal was reflected in the pattern of D‐loop variation of some of the islands but not all. A more unexpected association with a distant North European source area was also detected in three islands, possibly reflecting human contact with the Azores prior to the 15th century discovery by Portuguese mariners. Widening the scope to colonization of the Macaronesian islands as a whole, human linkages between the Azores, Madeira, the Canaries, Portugal and Spain were revealed through the sharing of mouse sequences between these areas. From these and other data, we suggest mouse studies may help resolve historical uncertainties relating to the ‘Age of Discovery’.  相似文献   

8.
In order to emphasize the importance of vegetation mapping for nature conservation purposes a case study in Terceira island (Azores) is presented, in which the importance of the natural vegetation of the eastern slope of Santa Bárbara volcano (which is part of the Site of Community Importance of Santa Bárbara–Pico Alto) is evaluated through the elaboration of its vegetation map. Fourteen (14) different natural vegetation types were identified: grasslands (1 type), peat bogs (2 types), scrubs (2), forests (5), successional vegetation (3) and vegetation of rocky slopes (1). All communities are protected under the Habitat and Species Directive (EC/92/43) and most of them are endemic to the Azores Islands. This fact, together with the significant number of Azorean endemic taxa (18), Macaronesian endemic taxa (5) and species protected under the Habitat and Species Directive (7), gives this area an important conservation value that justifies future protection actions. Vegetation mapping is an important tool for the characterization, evaluation and implementation of managing plans of natural areas of the Azores islands. The use of a floristic-based classification, supported by multivariate analysis and structural data, is an efficient methodology for the construction of these maps. The data collected comprise an important set of information about the distribution and abundance of natural vegetation types and endemic and rare species. This information was not available until now and is indispensable for the elaboration of management plans of Special Zones for Conservation that will be part of the NATURA 2000 network.  相似文献   

9.
The Azores archipelago (Portugal) is formed by nine islands whose relative positions define them as three geographical groups: Eastern (S. Miguel and Sta. Maria), Central (Terceira, Faial, Pico, Graciosa and S. Jorge) and Western (Flores and Corvo). Using the father's surname of 187,398 individuals living on the nine Azorean Islands, a population analysis based on inter-island relationship and hierarchical organization was conducted. The relation between islands was investigated using summary statistics, analysis of molecular variance (AMOVA) as well as graphical methods. When the values of heteronymy were contrasted with values of gene diversity based on haplogroup frequencies of the Y chromosome, it was possible to verify that Graciosa and Sta. Maria appeared to have the least diverse populations, and that Flores, despite its smaller population size and geographical isolation, has considerably higher levels of diversity. As for inter-island relationships, the difficulty of directly interpreting summary statistics values was evidenced. The AMOVA revealed that only 0.77% of the variation in surnames can be attributed to among-island variation, a value that, although small, can be considered significant. Application of Malécot's model revealed that geographic distance has an important impact in the genetic structure of the archipelago. Monmonier's maximum-difference algorithm demonstrated that the most isolated island of the archipelago appears to be Graciosa, followed by the islands of the Western group and by Sta. Maria. After integrating values of summary statistics with results from AMOVA and graphical methods, a more accurate genetic profile of the Azores, highly supported by genetic data, has emerged.  相似文献   

10.
Oceanic islands are vulnerable ecosystems and their flora has been under pressure since the arrival of the first humans. Human activities and both deliberately and inadvertently introduced biota have had and continue to have a severe impact on island endemic plants. The number of alien plants has increased nearly linearly on many islands, perhaps resulting in extinction‐based saturation of island floras. Here, we provide evidence for such a scenario in Alejandro Selkirk, Robinson Crusoe Islands (Archipelago Juan Fernández, Chile). We compared species richness and species composition of historical vegetation samples from 1917 with recent ones from 2011. Changes in species’ relative occurrence frequency were related to their taxonomic affiliation, dispersal mode, distribution status, and humidity and temperature preferences. While total species richness of vascular plants remained relatively similar, species composition changed significantly. Plants endemic to the Robinson Crusoe Islands declined, exotic species increased substantially within the period of ca. 100 years. Further, the relative occurrence frequency of plants with preferences for very warm and humid climate decreased, while the opposite was found for plants preferring drier and colder environments. Potential drivers responsible for this dramatic shift in the vegetation within only one century might have been the large goat population affecting especially small populations of endemic plants and climatic changes. Taking into account a substantial extinction debt, we expect further shifts in the vegetation of this small oceanic island toward alien plants. This would have significant negative consequences on global biodiversity, considering that island floras contribute substantially to global plant species richness due to their high proportion of endemics.  相似文献   

11.
Territorial songs in island populations of songbirds are often highly divergent from those of neighbouring continental relatives. This is shown for the three goldcrest subspecies (Regulus regulus azoricus, R. r. sanctae-mariae and R. r. inermis) endemic on six islands of the Azorean archipelago. All investigated populations display a high intra- and inter-individual acoustic variation. On each island, up to six different song types have been found; and a single male sings up to three types. In contrast, all northwestern European populations of R. r. regulus and R. r. anglorum share only a single song type. In playback experiments, none of 18 tested dialect songs of Azorean goldcrests evoked notable territorial reaction in German and Czech goldcrest males (ssp. regulus). Two differing dialect groups of the goldcrest can be distinguished on the Azores. Populations of the eastern islands, São Miguel and Santa Maria, share common song types which are not found on the islands of the central and western groups. Dialect repertoires on the westernmost islands, Flores, Faial and Pico, are dominated by a different song type. In the geologically younger western crater of São Miguel, both western and eastern song types coexist. Acoustic similarities to a population from neighbouring Terceira suggest the western part of São Miguel as the origin for the westward expansion of R. regulus on the Azores.  相似文献   

12.
This paper introduces the integration of additive partitioning with species—area relationships to island biogeography in order to address the question “How are the pteridophyte and spermatophyte native and endemic flora of different oceanic archipelagos partitioned across islands?”.Species richness data of all endemic species and all native species of pteridophytes and spermatophytes were obtained for the Azores, Canaries and Cape Verde in the Atlantic Ocean and Galápagos, Hawaii and Marquesas in the Pacific Ocean. Additive partitioning of species diversity was used to quantify how much of the total diversity of an oceanic archipelago flora (γ-diversity) is due to (i) the mean species richness of the flora of each island (α-diversity), (ii) the variability in species richness of the floras across islands (βNestedness) and (iii) the complementarity in species composition of the floras of different islands (βReplacement). The analysis was separately performed for the native and endemic pteridophyte and spermatophyte floras.The diversity partitioning of the six archipelagos showed large differences in how the flora of each archipelago is partitioned among the α, βNestedness and βReplacement components, for pteridophytes and spermatophytes and for all endemic species and all native species. The α-diversity was more important for all native species than for endemic species and more important for pteridophytes than for spermatophytes, with the Azores showing outstanding high values of α-diversity. The βNestedness was higher for pteridophytes than for spermatophytes and higher for endemic species than for all native species in both pteridophytes and spermatophytes. The values of βReplacement suggested that: (i) the spermatophyte native flora is more differentiated across islands than the pteridophyte native flora and (ii) the pteridophyte endemic flora and, especially, the spermatophyte endemic flora are more differentiated across islands than the corresponding native flora. An outstanding value of βReplacement for endemic and all native spermatophytes was found in Hawaii, confirming the biogeographical island differentiation in this archipelago.  相似文献   

13.
Aim Speciation processes on islands are still poorly understood. Previous studies based on the analysis of distribution data from checklists found that the flora of the Azores archipelago differs from other island floras in the exceptionally low number of radiations and the low number of single‐island endemics. The general mechanism(s) responsible for these apparently unique patterns remained unclear. One possible explanation for the distinctiveness of the Azorean endemic flora is the lack of a consistent and critical taxonomic framework for the floras of the Atlantic archipelagos. In this study, molecular variation within a range of Azorean endemic plant lineages was analysed to determine whether inadequacies in the current taxonomy of endemics might be an explanation for the unusual diversity patterns observed in the endemic flora of the Azores. Location Azores archipelago. Method Sixty‐nine populations of eight endemic species or subspecies belonging to five genetic lineages were sampled from all Azorean islands but one. Nuclear and plastid DNA regions were sequenced, and relationships among internal transcribed spacer (ITS) region ribotypes established using statistical parsimony. Results Molecular diversity patterns differ from current taxonomic groupings, with all lineages comprising previously overlooked genetic entities. Main conclusions Recognition as distinct taxa of the genetically distinct entities discovered in this study would drastically change the diversity patterns and make them more similar to those of other Atlantic archipelagos. The results serve to highlight that current knowledge of endemic diversity on oceanic islands may be far from complete, even in relatively well‐known groups such as angiosperms. This limitation is rarely considered in macroecological and evolutionary studies that make use of data from taxonomic checklists to draw inferences about oceanic island biogeographic processes.  相似文献   

14.
The Azores archipelago was selected as a case study since there are few studies on macroalgae introduction in oceanic islands. While at a global scale, around 3 % of macroalgae are considered non-indigenous; in the remote oceanic islands of the Azores, over 6 % of the marine algal flora is non-indigenous. The taxa distribution pattern of non-indigenous species in the Azores is significantly different from the distribution pattern in the globe. The most representative group was Rhodophyta species, being 84 % of the total non-indigenous macroalgae, mainly introduced via maritime traffic. This study highlights the vulnerability of remote islands to the introduction of macroalgae and the need to develop further studies on other archipelagos to understand whether the observed vulnerability is generally characteristic of oceanic islands. The development of local monitoring and mitigation programs and the necessity of regulatory and preventive measures for the maritime traffic vector are strongly suggested.  相似文献   

15.
Islands have long provided material and inspiration for the study of evolution and ecology. The West Indies are complex historically and geographically, providing a rich backdrop for the analysis of colonization, diversification and extinction of species. They are sufficiently isolated to sustain endemic forms and close enough to sources of colonists to develop a dynamic interaction with surrounding continental regions. The Greater Antilles comprise old fragments of continental crust, some very large; the Lesser Antilles are a more recent volcanic island arc, and the low-lying Bahama Islands are scattered on a shallow oceanic platform. Dating of island lineages using molecular methods indicates over-water dispersal of most inhabitants of the West Indies, although direct connections with what is now southern Mexico in the Early Tertiary, and subsequent land bridges or stepping stone islands linking to Central and South America might also have facilitated colonization. Species-area relationships within the West Indies suggest a strong role for endemic radiations and extinction in shaping patterns of diversity. Diversification is promoted by opportunities for allopatric divergence between islands, or within the large islands of the Greater Antilles, with a classic example provided by the Anolis lizards. The timing of colonization events using molecular clocks permits analysis of colonization-extinction dynamics by means of species accumulation curves. These indicate low rates of colonization and extinction for reptiles and amphibians in the Greater Antilles, with estimated average persistence times of lineages in the West Indies exceeding 30Myr. Even though individual island populations of birds might persist an average of 2Myr on larger islands in the Lesser Antilles, recolonization from within the archipelago appears to maintain avian lineages within the island chain indefinitely. Birds of the Lesser Antilles also provide evidence of a mass extinction event within the past million years, emphasizing the time-heterogeneity of historical processes. Geographical dynamics are matched by ecological changes in the distribution of species within islands over time resulting from adaptive radiation and shifts in habitat, often following repeatable patterns. Although extinction is relatively infrequent under natural conditions, changes in island environments as a result of human activities have exterminated many populations and others--especially old, endemic species--remain vulnerable. Conservation efforts are strengthened by recognition of aesthetic, cultural and scientific values of the unique flora and fauna of the West Indies.  相似文献   

16.
Invasive species are a threat for ecosystems worldwide, especially oceanic islands. Predicting the invasive potential of introduced species remains difficult, and only a few studies have found traits correlated to invasiveness. We produced a molecular phylogenetic dataset and an ecological trait database for the entire Azorean flora and find that the phylogenetic nearest neighbour distance (PNND), a measure of evolutionary relatedness, is significantly correlated with invasiveness. We show that introduced plant species are more likely to become invasive in the absence of closely related species in the native flora of the Azores, verifying Darwin's 'naturalization hypothesis'. In addition, we find that some ecological traits (especially life form and seed size) also have predictive power on invasive success in the Azores. Therefore, we suggest a combination of PNND with ecological trait values as a universal predictor of invasiveness that takes into account characteristics of both introduced species and receiving ecosystem.  相似文献   

17.
Origin and evolution of endemic plants of the Bonin (Ogasawara) Islands   总被引:1,自引:0,他引:1  
The Bonin Islands are typical oceanic islands, located at the western part of the North Pacific Ocean and approximately 1,000 km south of mainland Japan. This archipelago consists of about 20 small islands. Although floristic diversity is low due to the small area and limited environmental diversity, the Bonin Islands harbor unique endemic flora as in other well-known oceanic islands. This paper presents a brief summary of the results obtained from recent studies on the endemic flora of the Bonin Islands. The results are reviewed in relation to the four stages of the evolution of endemic flora in the oceanic islands; migration, establishment, enlargement and diversification. The ancestors of the flora originated mostly from tropical and subtropical Southeast Asia or mainland Japan by rare events of long distance dispersal. The proportion of bird-dispersed species is relatively high as for other oceanic islands. Genetic data sets obtained from allozyme variation in some endemic species suggest that migration occurred several million years ago and genetic diversity is correlated with current population size. At the time of establishment, self-compatible plants are expected to have an advantage. However, the percentage of dioecious plants is relatively high. This is partly due to evolutionary changes from hermaphroditic ancestors to dioecy which occurred in two genera in the Bonin Islands. In addition, there are some examples of evolutionary changes from herbaceous ancestors to woody endemics. Adaptive radiation is found in some genera, although the number of congeneric endemic species is less than five. Studies of allozyme variation inPittosporum, Symplocos andCrepidiastrum showed that genetic identity is generally very high between congeneric species in spite of their distinct morphologies. This result suggests that divergence of these species occurred rather recently and distinct morphological differences are based on a limited number of genetic changes.  相似文献   

18.
Madagascar is surrounded by archipelagos varying widely in origin, age and structure. Although small and geologically young, these archipelagos have accumulated disproportionate numbers of unique lineages in comparison to Madagascar, highlighting the role of waif-dispersal and rapid in situ diversification processes in generating endemic biodiversity. We reconstruct the evolutionary and biogeographical history of the genus Psiadia (Asteraceae), a plant genus with near equal numbers of species in Madagascar and surrounding islands. Analyzing patterns and processes of diversification, we explain species accumulation on peripheral islands and aim to offer new insights on the origin and potential causes for diversification in the Madagascar and Indian Ocean Islands biodiversity hotspot. Our results provide support for an African origin of the group, with strong support for non-monophyly. Colonization of the Mascarenes took place by two evolutionary distinct lineages from Madagascar, via two independent dispersal events, each unique for their spatial and temporal properties. Significant shifts in diversification rate followed regional expansion, resulting in co-occurring and phenotypically convergent species on high-elevation volcanic slopes. Like other endemic island lineages, Psiadia have been highly successful in dispersing to and radiating on isolated oceanic islands, typified by high habitat diversity and dynamic ecosystems fuelled by continued geological activity. Results stress the important biogeographical role for Rodrigues in serving as an outlying stepping stone from which regional colonization took place. We discuss how isolated volcanic islands contribute to regional diversity by generating substantial numbers of endemic species on short temporal scales. Factors pertaining to the mode and tempo of archipelago formation and its geographical isolation strongly govern evolutionary pathways available for species diversification, and the potential for successful diversification of dispersed lineages, therefore, appears highly dependent on the timing of arrival, as habitat and resource properties change dramatically over the course of oceanic island evolution.  相似文献   

19.
Mitochondrial DNA sequence data were obtained for eight species of flightless Galapaganus endemic weevils and one winged close relative in order to study their colonization history and modes of diversification in the Galápagos Archipelago. Contrary to most other insular radiations, the phylogeny estimates we recovered for Galapaganus do not follow the progression rule of island biogeography. The penalized likelihood age estimates of colonization of the archipelago exceed the age of the emerged islands and underscore the potential role of now sunken seamounts for the early evolution of Galapaganus . The phylogeny proposes one intra-island origin for Galapaganus endemics, but monophyly tests suggest a larger contribution of in-situ speciation on older islands. Generalist habitat preferences were reconstructed as ancestral while shifts to highland habitats were reconstructed as having evolved independently on different islands. Magnitudes and patterns of diversification rate were found to differ between older and younger islands. Our analyses reveal that the colonization sequence of islands and timing of colonization of Galapaganus could be linked with the geological and volcanic history of the islands in a rather complex scenario. Even though most islands appear to have been colonized soon after their emergence, there are notable deviations from the pattern of sequential colonization expected under the progression rule when considering only the extant emerged islands. Patterns of diversification rate variation on older and younger islands correspond to the volcanic activity or remnants of such activity, while the pattern of independent evolution of restricted habitat preferences in different islands suggests that habitat shifts could also have contributed to species diversity in Galapaganus .  相似文献   

20.
This paper focuses on medicinal plants found on the Caribbean island of Montserrat; however, other plant uses were also documented when found. Two hundred seventy-two taxa from seventy-eight families are listed along with habitat information and ethnobotanical notes when pertinent. Brief treatments of the islands’ vegetation and geographical conditions are also included. Montserrat is a tropical island with a diverse flora and a rich heritage of plant folklore, the latter encompassing traditions passed down from Africans, Caribbean Amerindians, and Europeans. Prior to the recent volcanic eruptions, the author did a taxonomic study of ethnobotanically important plants found on Montserrat. Although the main purpose of this study was to gather ethnobotanical information, a number of plants which are not known to be ethnobotanically important to Montserratians were collected in order to augment our knowledge of the flora of the island. The island’s recent devastating volcanic eruptions (resulting in the deaths of several informants and obliteration of large tracts of vegetation) have made the data collected in this project all the more significant.  相似文献   

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