Modeling Species Distributions from Heterogeneous Data for the Biogeographic Regionalization of the European Bryophyte Flora

The definition of biogeographic regions provides a fundamental framework for a range of basic and applied questions in biogeography, evolutionary biology, systematics and conservation. Previous research suggested that environmental forcing results in highly congruent regionalization patterns across taxa, but that the size and number of regions depends on the dispersal ability of the taxa considered. We produced a biogeographic regionalization of European bryophytes and hypothesized that (1) regions defined for bryophytes would differ from those defined for other taxa due to the highly specific eco-physiology of the group and (2) their high dispersal ability would result in the resolution of few, large regions. Species distributions were recorded using 10,000 km² MGRS pixels. Because of the lack of data across large portions of the area, species distribution models employing macroclimatic variables as predictors were used to determine the potential composition of empty pixels. K-means clustering analyses of the pixels based on their potential species composition were employed to define biogeographic regions. The optimal number of regions was determined by v-fold cross-validation and Moran’s I statistic. The spatial congruence of the regions identified from their potential bryophyte assemblages with large-scale vegetation patterns is at odds with our primary hypothesis. This reinforces the notion that post-glacial migration patterns might have been much more similar in bryophytes and vascular plants than previously thought. The substantially lower optimal number of clusters and the absence of nested patterns within the main biogeographic regions, as compared to identical analyses in vascular plants, support our second hypothesis. The modelling approach implemented here is, however, based on many assumptions that are discussed but can only be tested when additional data on species distributions become available, highlighting the substantial importance of developing integrated mapping projects for all taxa in key biogeographically areas of Europe, and the Mediterranean peninsulas in particular.     

Concepts of classic and modern biogeography: Contribution of Russian entomologists

The main stages and ways of development of insect biogeography in Russia are discussed. Three general research programs (studies of ranges, zones, and regions) are described. The role of the regional approach in biogeography is characterized. Some problems related to studies of insect ranges are evaluated, and the need for creating practical range maps is emphasized. The relationships between the insect distribution patterns and the life zones are discussed. The specific features of zonation in insect biogeography are noted. Special attention is paid to establishing the zones with unified population distribution patterns within a species range. The role of geozoological investigations and opportunities of modern information technologies are characterized relative to evaluation of general patterns of insect assemblage distribution. The problems of regionalization in insect biogeography are discussed. Three main approaches (intuitive, a priori, and a posteriori) are determined. Some advantages of using population and assemblage distribution patterns in regionalization are emphasized. The role and possibilities of insect biogeography studies for in the field of reconstruction of past and long-term forecasting are characterized.     

Biogeographic structure of the northeastern Pacific rocky intertidal: the role of upwelling and dispersal to drive patterns

Identifying the environmental conditions that drive biogeographic structure remains a major challenge of biogeography, evolutionary ecology and increasingly, conservation biology. Here, we use multivariate classification trees to assess the biogeographic structure of northeast Pacific (～ 26–58°N) rocky intertidal species (406 species of algae and invertebrates) from 102 field sites. Random forest analyses are used to assess the importance of 29 environmental variables, encompassing a broad range of potential drivers, to predict biogeographic structure. Analyses are repeated for species with different larval dispersal capabilities and by broad taxonomic categories (invertebrates and algae). Results show that overall biogeographic structure is in general agreement with classic classification schemes, but patterns are variable among species with different larval dispersal capabilities. Random forest models show a very high fit (pseudo r² > 0.94) and indicate that biogeographic structure can be predicted by a relatively modest subset of variables. Upwelling related variables are the best overall predictors of biogeographic structure (nutrient concentrations, sea‐surface temperature, upwelling/downwelling seasonal switch index), but the relative importance of predictors is geographically variable and top predictors are dependent on the type of larval dispersal. Upwelling related variables are more important to predict biogeographic structure for invertebrates with lower‐medium dispersal capabilities and algae, whereas species with high larval dispersal (planktotrophic) are better predicted by a different subset of variables (i.e. salinity, precipitation seasonality). Our results lend support to the influence of coastal upwelling in structuring biogeographic patterns and highlight the potential for climate change‐induced alterations of upwelling regimes to profoundly affect biodiversity at biogeographic scales.     

INTRASPECIFIC PHYLOGEOGRAPHY ACROSS THE POINT CONCEPTION BIOGEOGRAPHIC BOUNDARY

Recent studies of intraspecific phylogeography have suggested that the geographic location of genetic discontinuities, or phylogeographic breaks, may frequently coincide with biogeographic boundaries. The concordance is hypothesized to reflect similarity in the processes governing species boundaries and intraspecific lineage boundaries. This concordance has not, however, been widely tested. In the case of the Point Conception biogeographic boundary between the Oregonian and Californian marine biotas, only the supralittoral copepod Tigriopus californicus has been found to have a coincident phylogeographic break. Here I show that the apparent relationship between this break and Point Conception was, in fact, an artifact of insufficient geographic sampling. Mitochondrial DNA analyses of T. californicus populations between Morro Bay and San Diego reveal at least five equally deep phylogeographic breaks in the region (where only one biogeographic boundary is recognized). Limited nuclear DNA sequence data and allozyme data also support the occurrence of multiple genetic discontinuities along this geographic range. Lack of one-to-one correspondence between intraspecific phylogeography and biogeographic boundaries indicates that the processes affecting the genetic differentiation of populations of T. californicus differ from those responsible for determining species distributional limits at the Point Conception biogeographic boundary. A review of genetic data from other species also fails to provide evidence for concordance of biogeography and intraspecific phylogeography across Point Conception. I suggest that the concordance of phylogeography with biogeography will only be pronounced where the biogeographic boundary separates biotas that are phylogenetically related. The numerous cases of interspecific hybrid zones in the region of Cape Canaveral, for example, indicate that many sister-species pairs occur across this biogeographic boundary. Such hybrid zones are not common at Point Conception, and there appears to be no cases of intraspecific phylogeographic breaks associated with this well-recognized biogeographic boundary.     

Marine shelf habitat: biogeography and evolution

We synthesize the evolutionary implications of recent advances in the fields of phylogeography, biogeography and palaeogeography for shallow‐water marine species, focusing on marine speciation and the relationships among the biogeographic regions and provinces of the world. A recent revision of biogeographic provinces has resulted in the recognition of several new provinces and a re‐evaluation of provincial relationships. These changes, and the information that led to them, make possible a clarification of distributional dynamics and evolutionary consequences. Most of the new conclusions pertain to biodiversity hotspots in the tropical Atlantic, tropical Indo‐West Pacific, cold‐temperate North Pacific, and the cold Southern Ocean. The emphasis is on the fish fauna, although comparative information on invertebrates is utilized when possible. Although marine biogeographic provinces are characterized by endemism and thus demonstrate evolutionary innovation, dominant species appear to arise within smaller centres of high species diversity and maximum interspecies competition. Species continually disperse from such centres of origin and are readily accommodated in less diverse areas. Thus, the diversity centres increase or maintain species diversity within their areas of influence, and are part of a global system responsible for the maintenance of biodiversity over much of the marine world.     

The niche, biogeography and species interactions

In this paper, I review the relevance of the niche to biogeography, and what biogeography may tell us about the niche. The niche is defined as the combination of abiotic and biotic conditions where a species can persist. I argue that most biogeographic patterns are created by niche differences over space, and that even ‘geographic barriers’ must have an ecological basis. However, we know little about specific ecological factors underlying most biogeographic patterns. Some evidence supports the importance of abiotic factors, whereas few examples exist of large-scale patterns created by biotic interactions. I also show how incorporating biogeography may offer new perspectives on resource-related niches and species interactions. Several examples demonstrate that even after a major evolutionary radiation within a region, the region can still be invaded by ecologically similar species from another clade, countering the long-standing idea that communities and regions are generally ‘saturated’ with species. I also describe the somewhat paradoxical situation where competition seems to limit trait evolution in a group, but does not prevent co-occurrence of species with similar values for that trait (called here the ‘competition–divergence–co-occurrence conundrum’). In general, the interface of biogeography and ecology could be a major area for research in both fields.     

Diversity and geographic distribution of ciliates (Protista: Ciliophora)

About 4,500 free-living ciliate morphospecies have been described, applying an average synonymy rate of 20%. We estimate that 83–89% of the ciliate diversity is still undescribed, using the following probabilities: detailed habitat studies suggest that the described number of morphospecies must be doubled: 4,500 → 9,000; this figure has to be increased by about 50% due to species with similar interphase morphology but different resting cysts: 9,000 → 13,500; the genetic and molecular data suggest that this value must be doubled or trebled: 13,500 → 27,000 to 40,000 free-living, biological ciliate species. The knowledge on geographic distribution of ciliates heavily depends on flagship species and statistical analyses because reliable faunistic studies are rare and molecular data are still in its infancy. We present a list of 52 ciliate flagship species as a testable hypothesis, i.e., the hypothesis of restricted distribution of certain ciliate species must be refused when a considerable number of them is found in all or most biogeographic regions. Flagship species and statistical analyses consistently show Gondwanan and Laurasian ciliate communities, suggesting that the split of Pangaea deeply influenced ciliate distribution and rare species play a key role in geographic differentiation. However, there is also substantial evidence for continental, regional, and local endemism of free-living ciliates. The molecular studies usually show a high level of genetic diversity underlying ciliate morphospecies, suggesting that morphologic and molecular evolution may be decoupled in many ciliate species. Molecular studies on ciliate biogeography are at variance, possibly because most are still focusing on single molecular markers. In sum, the data indicate that ciliate biogeography is similar to that of plants and animals, but with an increased proportion of cosmopolites, favouring the moderate endemicity model. Special Issue: Protist diversity and geographic distribution. Guest editor: W. Foissner.     

内蒙古苏崩地点的古新世软食中兽(哺乳纲,中兽目):古新世-始新世界线附近“东方伊甸园”扩散模式的新证据(英文)

记述了发现于内蒙古苏崩晚古新世格沙头期的中兽类软食中兽Hapalodectes属的一个新种。这是软食中兽在中国古新世地层中的首次发现,也是亚洲格沙头期的第二种软食中兽。已有的系统学和生物地层学证据支持软食中兽属和软食中兽科(Hapalodectidae)亚洲起源的观点。软食中兽显然是在古新世-始新世极热事件(PETM)期间通过白令陆桥扩散到北美大陆的,因而符合"东方伊甸园"学说中的生物地理格局。软食中兽有限的(即非欧洲的)地理分布使得我们可以重建该属生物地理学历史。如同软食中兽一样的"东方伊甸园"式的扩散模式,可以看作是大的环境变化事件导致多个支系产生相似的系统学和生物地理学分布格局的生物地理扩散机制。严格地检查了所谓的在古新世-始新世界线上或其附近的与"东方伊甸园"模式相矛盾的大陆间哺乳动物扩散事例,结果发现这些例子都是不可靠的。"东方伊甸园"生物地理学说充分解释了PETM时期哺乳动物群更替以及劳亚古陆哺乳动物地理分布格局的成因。     

Defining and Dividing the Greater Caribbean: Insights from the Biogeography of Shorefishes

The Greater Caribbean biogeographic region is the high-diversity heart of the Tropical West Atlantic, one of four global centers of tropical marine biodiversity. The traditional view of the Greater Caribbean is that it is limited to the Caribbean, West Indies, southwest Gulf of Mexico and tip of Florida, and that, due to its faunal homogeneity, lacks major provincial subdivisions. In this scenario the northern 2/3 of the Gulf of Mexico and southeastern USA represent a separate temperate, “Carolinian” biogeographic region. We completed a comprehensive re-assessment of the biogeography of the Greater Caribbean by comparing the distributions of 1,559 shorefish species within 45 sections of shelf waters of the Greater Caribbean and adjacent areas. This analysis shows that that the Greater Caribbean occupies a much larger area than usually thought, extending south to at least Guyana, and north to encompass the entire Carolinian area. Rather than being homogenous, the Greater Caribbean is divided into three major provinces, each with a distinctive, primarily tropical fauna: (1) a central, tropical province comprising the West Indies, Bermuda and Central America; (2) a southern, upwelling-affected province spanning the entire continental shelf of northern South America; and (iii) a northern, subtropical province that includes all of the Gulf of Mexico, Florida and southeastern USA. This three-province pattern holds for both reef- and soft bottom fishes, indicating a general response by demersal fishes to major variation in provincial shelf environments. Such environmental differences include latitudinal variation in sea temperature, availability of major habitats (coral reefs, soft bottom shorelines, and mangroves), and nutrient additions from upwelling areas and large rivers. The three-province arrangement of the Greater Caribbean broadly resembles and has a similar environmental basis to the provincial arrangement of its sister biogeographic region, the Tropical Eastern Pacific.     

Biogeography and conservation in Southeast Asia: how 2.7 million years of repeated environmental fluctuations affect today’s patterns and the future of the remaining refugial-phase biodiversity

Understanding the historical biogeography of this global biodiversity hotspot is as important to long-term conservation goals as ecology and evolution are to understanding current patterns and processes. Today’s geography is, however, misleading and typical of only ~2% of the last million years; >90% of that time the region’s land area was 1.5–2.0 times larger as mean sea levels were 62 m below today’s, climates were cooler, and extensive forests and savanna covered the emerged Sunda plains. The region’s land area varied two-fold as sea levels fluctuated up to ±50 m with each of ~50 Pleistocene glacial cycles, and forests expanded and contracted with oscillations in land area and seasonality. This dynamic geographic history is relevant to the development of biogeographic regionalism and shows that it is today’s forests that are refugial, not those of the Last Glacial Maximum. This history affects how species will adapt or shift their ranges in response to global warming and further decreases in land area (submergence of low-lying coastal areas) during the 21st century. The alternative is mass species extinction. The biota is also threatened by the continued destruction of forest, destruction of Mekong River flood-pulse based ecosystems, and continued human population growth. Human biogeography will become more important in conservation planning as tens of millions of people who depend on protected area forests, riverine ecosystems, and coastal habitats become environmental refugees. Conservation scientists need to become more involved in regional ecological education, environmental stewardship, and ecosystem-based adaptation to sustain as much as possible of this rich biota and the ecological services it provides.     

Extinctions of montane mammals reconsidered: putting a global-warming scenario on ice

McDonald and Brown developed a model based on island biogeographic principles to predict the magnitude and composition of small mammal extinctions from isolated boreal habitats atop mountains of the Great Basin following global warming. The model predicts that three of 14 boreal mammals will go extinct regionally and that four of 19 mountain ranges will lose upwards of 50% of their present faunas. Here, we re-examine the model on the basis of its underlying assumptions, on the statistical and biogeographic protocols used, and on its predictive power. A key assumption, that populations of these small mammals are isolated by absolute barriers to dispersal, is challenged by published field observations and by extensive trapping records. Statistical procedures used to construct the model are questionable and the model itself yields imprecise estimates. The biogeographic principle used to identify extinction-prone species, nested subsets of species, makes predictions that are at odds with available autecological information. The demonstration of a nested pattern of species occurrences does not provide definitive evidence in resolving SLOSS-or whether a single large island or several small islands of equivalent total area will contain more species. We conclude that the model is not a reliable method for forecasting species extinctions following global warming. The final resolution of the biogeography of montane mammals (and predictive models of extinction) in the Great Basin must await a full and accurate accounting of past and present species distributions.     

中国鸟类生物地理学研究回顾与展望

本文回顾总结了中国鸟类生物地理学的产生、发展、重要成果和未来的发展趋势.主要论述了以下五个方面:1.中国动物地理学简史及鸟类生物地理学的萌芽;2.鸟类动物地理区划的发展,并重点论述了岛屿鸟类区系调查与生物地理格局、中国台湾岛和海南岛与大陆鸟类区系的联系以及鸟类特有化现象;3.鸟类多样性的格局、形成与维持,并重点归纳了几...     

Panbiogeography and conservation science in New Zealand

Abstract

New Zealand’s official conservation science framework, the Protected Natural Areas Programme (PNA), is evaluated in relation to current biogeographic/systematic methods and principles with reference to philosophy, taxonomy, information content, and historical geological/biological relations. The PNA Programme and its parent International Union for Conservation of Nature and Natural Resources (IUCN) framework is based on a phenetic system of classification that does not reflect process (phylogenetic) characters of ecological systems. The PNA philosophy of conservation science is shown to be essentialist, rooted in traditional medieval Western metaphysics, and out of step with current developments in biogeography, systematics, and ecology. Panbiogeography is an appropriate global perspective for developing a conservation science because it meets the requirements of homology, monophyly, increased information content, and empirical testability whereas the PNA programme does not. Establishment of a Panbiogeographic Track Atlas is proposed as a suitable conservation framework for historical ecology and biogeography. The atlas could provide an empirical natural resource inventory to identify priority areas for conservation at an economically acceptable cost compared to the PNA Programme. Application of cladistic techniques to ecological and biogeographic patterns in relation to the Atlas can provide aphylogenetically sound hierarchical classification for conservation science. The potential benefits of the panbiogeographic approach for conservation education and tourism are discussed.     

Tectonic blocks and molecular clocks

Evolutionary timescales have mainly used fossils for calibrating molecular clocks, though fossils only really provide minimum clade age constraints. In their place, phylogenetic trees can be calibrated by precisely dated geological events that have shaped biogeography. However, tectonic episodes are protracted, their role in vicariance is rarely justified, the biogeography of living clades and their antecedents may differ, and the impact of such events is contingent on ecology. Biogeographic calibrations are no panacea for the shortcomings of fossil calibrations, but their associated uncertainties can be accommodated. We provide examples of how biogeographic calibrations based on geological data can be established for the fragmentation of the Pangaean supercontinent: (i) for the uplift of the Isthmus of Panama, (ii) the separation of New Zealand from Gondwana, and (iii) for the opening of the Atlantic Ocean. Biogeographic and fossil calibrations are complementary, not competing, approaches to constraining molecular clock analyses, providing alternative constraints on the age of clades that are vital to avoiding circularity in investigating the role of biogeographic mechanisms in shaping modern biodiversity.This article is part of the themed issue ‘Dating species divergences using rocks and clocks’.     

Reconstruction of adaptive radiation in insular habitats using integrated biogeographic information

Aim To develop an analytical method for accurately reconstructing the biogeographic events associated with adaptive radiation in a system of insular habitats. Location An idealized, two-dimensional model system of insular habitats is based on the altitudinally stratified vegetation zones on the island-like mountains of eastern Africa. Methods The model system is treated as a two-dimensional array of insular habitats, and adaptive radiation is treated as a ‘space-filling’ process according to six premises based on uniformitarian principles. Previous approaches to this class of problems have used (1) Hennig's progression rule, (2) optimization of biogeographic character states, and (3) reasoned argumentation with an intuitive synthesis of information. The strengths and limitations of these previous approaches are evaluated. Results A closed analytical method is presented that accurately reconstructs biogeographic events. This methodological approach integrates atomized information back up to the appropriate level of biological organization and has general applicability to phylogenetic analysis. Main conclusions In a truly evolutionary approach to phylogenetic systematics and biogeography, the initial analysis of the pattern of descent should be complemented with a subsequent analysis of the pattern of modification. The method presented here offers one approach to analysing the pattern of modification, which in this case constitutes biogeographic movement.     

Inference of biogeographic history by formally integrating distinct lines of evidence: genetic,environmental niche and fossil

A primary focus of historical biogeography is to understand changes in species ranges, abundance and genetic connectivity, and changes in community composition. Traditionally, biogeographic inference has relied on distinct lines of evidence, including DNA sequences, fossils and hindcasted ecological niche models. In this review we propose that the development of integrative modeling approaches that leverage multiple distinct data types from diverse disciplines has the potential to revolutionize the field of biogeography. Although each data type contains information on a distinct aspect of species’ biogeographic histories, few studies formally integrate multiple types in analysis. For example, post hoc congruence among analyses based on different data types (e.g. fossils and genetics) is commonly assumed to indicate likely biogeographic histories. Unfortunately, analyses of different data often reach discordant conclusions. Thus, fundamental and unresolved debates continue regarding speed and timing of postglacial migration, location and size of glacial refugia, and degree of long distance dispersal. Formal statistical integration can help address these issues. More specifically, formal integration can leverage all available evidence, account for inherent biases associated with different data types, and quantify data and process uncertainty. Novel, quantitative integration of data and models across fields is now possible due to recent advances in cyberinfrastructure, spatial modeling, online and aggregated ecological databases, data processing and quantitative methods. Our purpose is to make the case for and give examples of rigorous integration of genetic, fossil and environmental/occurrence data for inferring biogeographic history. In particular, we 1) review the need for such a framework; 2) explain common data types and approaches used to infer biogeographic history (and the challenges with each); 3) review state‐of‐the‐art examples of data integration in biogeography; 4) lay out a series of novel, suggested improvements on current methods; and 5) provide an outlook on technical feasibility and future opportunities.     

Psephellus turcicus sp. nov. (Asteraceae), a new chasmophyte species from central Anatolia, Turkey

A new species, Psephellus turcicus A.Duran & Hamzaoğlu (Asteraceae) in sect. Psephelloidei (Boiss.) Wagenitz & Hellwig is described and illustrated from Anatolia, Turkey. The species grows on limestone crevices in Kazankaya Canyon (A5 Yozgat) in central Anatolia. A neo-endemic confined to Kazankaya Canyon, it is closely related to Psephellus psephelloides (Freny & Sint.) Wagenitz. Diagnostic morphological characters from closely similar taxa are discussed, and arranged in a key of Turkish similar Psephellus Cass. The ecology, biogeography and conservation status of the species are also presented. Achene surface morphology of P. turcicus and P. psephelloides are examined by SEM. The geographical distribution of the new species and other related species is mapped.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 495–500.     

Toad radiation reveals into-India dispersal as a source of endemism in the Western Ghats-Sri Lanka biodiversity hotspot

Background  

High taxonomic level endemism in the Western Ghats-Sri Lanka biodiversity hotspot has been typically attributed to the subcontinent's geological history of long-term isolation. Subsequent out of – and into India dispersal of species after accretion to the Eurasian mainland is therefore often seen as a biogeographic factor that 'diluted' the composition of previously isolated Indian biota. However, few molecular studies have focussed on into-India dispersal as a possible source of endemism on the subcontinent. Using c. 6000 base pairs of mitochondrial and nuclear DNA, we investigated the evolutionary history and biogeography of true toads (Bufonidae), a group that colonized the Indian Subcontinent after the Indo-Asia collision.     

The temporal dimension of marine speciation

Speciation is a process that occurs over time and, as such, can only be fully understood in an explicitly temporal context. Here we discuss three major consequences of speciation’s extended duration. First, the dynamism of environmental change indicates that nascent species may experience repeated changes in population size, genetic diversity, and geographic distribution during their evolution. The present characteristics of species therefore represents a static snapshot of a single time point in a species’ highly dynamic history, and impedes inferences about the strength of selection or the geography of speciation. Second, the process of speciation is open ended—ecological divergence may evolve in the space of a few generations while the fixation of genetic differences and traits that limit outcrossing may require thousands to millions of years to occur. As a result, speciation is only fully recognized long after it occurs, and short-lived species are difficult to discern. Third, the extinction of species or of clades provides a simple, under-appreciated, mechanism for the genetic, biogeographic, and behavioral ‘gaps’ between extant species. Extinction also leads to the systematic underestimation of the frequency of speciation and the overestimation of the duration of species formation. Hence, it is no surprise that a full understanding of speciation has been difficult to achieve. The modern synthesis—which united genetics, development, ecology, biogeography, and paleontology—greatly advanced the study of evolution. Here we argue that a similarly synthetic approach must be taken to further our understanding of the origin of species.     

Psephellus yusufeliensis sp. nov. from north-east Anatolia, Turkey

Psephellus yusufeliensis O. Tugay & Uysal sp. nov. (Asteraceae) from Yusufeli (B7 Artvin-Turkey) is described and illustrated. It is closely related to P. aucheriana (DC) Wagenitz & F. H. Hellw. Diagnostic morphological characters are provided and the relationship to other Psephellus species in Turkey is discussed. The geographical distribution of the new species and other related species are mapped. Notes are also presented on the ecology, biogeography and conservation status of the new species.