Nested areas of endemism analysis

Aim  To develop and evaluate a nested clade approach to aid in the determination of areas of endemism (AoE) in biogeographical studies.
Methods  We adapted the nested clade analysis (NCA) to studies of areas of endemism. For this purpose we adapted several of the programs currently in use. Two data sets were examined using this approach – one involving Sciobius in southern Africa and the other involving terrestrial mammals in Mexico.
Results  Nested clade analysis as applied to areas of endemism produced results similar to those of previous analyses of Sciobus in southern Africa. An analysis of terrestrial mammals in Mexico supports the designation of some biogeographical provinces as areas of endemism while suggesting that other provinces may comprise composite distributions that should be subdivided.
Main conclusions  The nested clade analysis approach utilized primarily in genetic analysis of phylogeographical patterns in population biology studies can be adapted to understanding AoE in the realm of biogeography. This approach offers a statistical paradigm to evaluate AoE suggested by parsimony analysis of endemicity (PAE) trees.     

Areas and algorithms: evaluating numerical approaches for the delimitation of areas of endemism in the Canary Islands archipelago

Aim Areas of endemism are the fundamental units of cladistic biogeographical analysis but there is no consensus on the most appropriate method for their delimitation. In this paper, the relative performance of a number of algorithmic approaches for the delimitation of areas of endemism is investigated within the context of the Canary Islands flora, and areas of endemism within the Canary Islands archipelago are defined. Location The Canary Islands. Methods A data matrix comprising the distributions of 609 endemic spermatophyte taxa (c. 90% of the endemic flora) scored on a 10 × 10 km UTM grid was analysed using: (1) UPGMA (unweighted pair group method with arithmetic mean) clustering of Jaccard and Kulczynski similarity coefficient matrices, (2) parsimony analysis of endemicity (PAE), and (3) the program ndm (eNDeMism). The performance of each method was then determined by the extent to which the resulting areas of endemism met three criteria: (1) possession of two or more strict endemic taxa, (2) diagnosability, and (3) geographical contiguity. Results Each of the four methods resulted in substantially different sets of areas. ndm analysis resolved 17 areas of endemism consistent with all three criteria, and collectively these accounted for 59% of all cells. In the hierarchical analyses none of the methods recovered more than eight areas of endemism, and the total coverage of cells ranged from 13% to 33% when the results were confined to intra‐island areas of endemism. Main conclusions ndm outperforms hierarchical clustering methods in terms of both the number of intra‐island areas of endemism delimited that meet the three evaluation criteria and the total coverage of those areas. ndm may also be considered preferable because it is non‐hierarchical, incorporates spatial information into the delimitation of areas, and permits overlap between areas of endemism where there is evidence to support it. The results support the use of ndm as the most appropriate method currently available for the delimitation of areas of endemism. The areas of endemism identified by the ndm analysis are discussed.     

Areas of endemism for passerine birds in the Atlantic forest, South America

Aim To use the method of parsimony analysis of endemism to identify areas of endemism for passerine birds in the Atlantic Forest, South America, and to compare the locations of these areas with areas previously identified for birds as well as other taxa. Location The Atlantic Forest, eastern South America. Methods We analysed a matrix composed of the presence (1) or absence (0) of 140 endemic species in 24 quadrats of 1 × 1 degree distributed along the Atlantic Forest to find the most parsimonious area cladogram. Results Fourteen most parsimonious cladograms were found and then summarized in a single consensus tree. Four areas of endemism were identified: Pernambuco, Central Bahia, Coastal Bahia, and Serra do Mar. Main conclusions Avian areas of endemism in the Atlantic Forest have significant generality, as they are highly nonrandom and congruent with those of other groups of organisms. A first hypothesis about the historical relationships among the four areas of avian endemism in the Atlantic Forest is delineated. There is a basal dichotomy among areas of endemism in the Atlantic Forest, with Pernambuco forming a northern cluster and Coastal Bahia, Central Bahia and Serra do Mar comprising a southern cluster. Within the southern cluster, Central Bahia and Serra do Mar are more closely related to each other than to Coastal Bahia.     

Areas of endemism of the Patagonian steppe: an approach based on insect distributional patterns using endemicity analysis

Aims  To delimit areas of endemism in the Patagonian steppe using endemicity analysis (EA), which evaluates areas of endemism by means of an endemicity index, and to compare the resulting endemic areas with those proposed for the Patagonian steppe by previous authors.
Location  The Patagonian steppe, a region of South America found approximately below parallel 36° S to the east of the Andes Mountains.
Methods  Distributional data for 159 species of insects collected in the Patagonian steppe, and consisting of 1317 georeferenced samples were used to identify areas of endemism. A data grid of presence and absence (with cells of 1° × 1°) was constructed. Initially, two different types of EA were performed, seeking areas defined by 'four or more' species. A first analysis was performed without taking into consideration those quadrats where no species had been recorded (empty quadrats), which in many cases meant a discontinuous distribution. The second analysis was performed assuming a continuous distribution for each species. A third analysis, assuming continuous distributions, was performed using 'three or more' as the number of species necessary for an area to be identified as an endemic area.
Results  In the first two analyses, EA recognized the same five areas of endemism: western Patagonia, south-western Payunia, northern Suabandean, southern Subandean and Austral Patagonia. The results of the third analysis allowed the identification of three more areas of endemism: northern Payunia, Chubutian and Santacrucian.
Main conclusions  We identified five areas of endemism for the Patagonian steppe, some of which have been defined in previous contributions. These areas are: Western Patagonia, Payunia and Subandean Patagonia (which can be divided into septentrional and meridional), Central Patagonia (Chubutence and Santacrucense) and Austral Patagonia.     

An improved method for the identification of areas of endemism using species co-occurrences

Aim To develop a simple method that (1) combines the notions of biotic elements (groups of taxa with ranges significantly more similar to each other than to the ranges of other taxa) and of areas of endemism (AoE, areas of non‐random distributional congruence among taxa), and (2) overcomes the constraints of a previously suggested null model‐based method that cannot deal with disjunctions and is strictly grid‐dependent. Location We used test data sets from southern Africa and Crete. Methods First, we used a null‐model approach to detect pairs of species that have a significant degree of co‐occurrence, in order to determine biotic elements. Subsequently, we used a parsimony analysis of endemicity to delineate candidate AoE, and multivariate analysis to define groups of biotic elements on the basis of species interactions (co‐occurrence, mutual exclusion, neutral) using only the species detected in the previous step. We applied this method to the well known data set for Sciobius in southern Africa, as well as to endemic invertebrates of Crete (Greece), in order to evaluate its performance. Results Our results are very similar to those of previous analyses, and produce meaningful delineation of AoE and biotic elements in both data sets. The method is flexible regarding null models and significance levels, and eliminates noise in the data. Main conclusions We offer a simple method that provides reasonable identification of both biotic elements and AoE, produces good‐fit statistics, reduces uninformative or junk output, and reduces computational time.     

Panbiogeography of New Caledonia: a response to Heads (2008)

The definition of areas of endemism is one of the most important steps for historical biogeography. Here I review the dataset used by Heads (Journal of Biogeography, 2008, 35 , 2153–2175) for his panbiogeographical analysis of the New Caledonian biota. I highlight that some of the distribution data appear dubious (some localities should have been included while some others should have been deleted) when compared with current databases. In addition, some conclusions are not supported by the data themselves.     

The other face of Lyell: historical biogeography in his Principles of geology

Although some excellent articles about Lyell's work have been published, they do not explicitly deal with Lyell's biogeographical conceptions. The purpose of this paper is to analyse Lyell's biogeographical model in terms of its own internal structure. Lyell tried to explain the distribution of organisms by appealing to a real cause (climate). However, he was aware that environmental conditions were clearly insufficient to explain the existence of biogeographical regions. Lyell's adherence to ecological determinism generated strong tensions within his biogeographical model. He shifted from granting a secondary weight to dispersal to assigning it a major role. By doing so, Lyell was led into an evident contradiction. A permanent tension in Lyell's ideas was generated by the prevalent explanatory pattern of his time. The explanatory model based on laws did not produce satisfactory results in biology because it did not deal with historical processes. We may conclude that the knowledge of organic distribution interested Lyell as long as it could be explained by the uniformitarian principles of his geological system. The importance of the second volume of the Principles of geology lies in its ample and systematic argumentation about the geographical distribution of organisms. Lyell established, independently from any theory about organic change, the first version of dispersalist biogeography.     

Areas of endemism and patterns of diversity for aphids of the Qinghai-Tibetan Plateau and the Himalayas

Aim  The study aimed to identify areas of endemism for aphids in the Qinghai-Tibetan Plateau and the Himalayas (QTPH), and to test congruence between patterns of endemism and patterns of overall species richness identified in a previous study.
Location  The QTPH.
Methods  A distribution data base of 326 endemic aphids in the QTPH was compiled. The study area was divided into a grid of 2°× 2° operative geographical units. Parsimony analysis of endemicity (PAE) was used to identify areas of endemism, and the diversity patterns of endemic species were then mapped using GIS.
Results  We identified 326 endemic species belonging to 138 genera within Adelgidae and 14 subfamilies of Aphididae. Five areas of endemism were found using PAE analysis: the eastern Himalayas, the western Himalayas, north-western Yunnan, southern Tibet and the eastern QTPH. Maps of patterns of endemism identified four major centres for endemic aphids, namely the western Himalayas, the eastern Himalayas (or Sikkim-Assam Himalayas), north-western Hengduan Mountains and the mountains of southern Gansu Province, and three minor centres, southern Tibet, south-eastern Tibet and the eastern Qinghai Province in the north-eastern QTPH.
Main conclusions  Our study identifies major centres of aphid endemism. Furthermore, there is a noticeable congruence between patterns of endemism and patterns of species richness. The patterns of endemism were most likely influenced by the recent uplift of the QTPH.     

Several Pleistocene refugia detected in the high alpine plant Phyteuma globulariifolium sternb & hoppe (Campanulaceae) in the European Alps

Phyteuma globulariifolium is a high alpine plant species growing in the European Alps and the Pyrenees. In order to elucidate its glacial history, 325 individuals from 69 populations were analysed using the amplified fragment length polymorphism (AFLP) technique. A strongly hierarchical phylogeographical pattern was detected: Two major east-west vicariant groups can be separated along a gap in the distributional area. A further subdivision into at least four populational groups is in congruence with presumed peripheral glacial refugia. There is no indication for survival on unglaciated mountain tops (nunataks) in the interior of the Pleistocene ice shield covering the Alps. Our results favour glacial survival in peripheral, unglaciated or not fully glaciated areas. Populations of P. globulariifolium in the Pyrenees are the result of relatively recent long-distance dispersal. Within the Alps, there is strong differentiation among groups of populations, whereas within them the differentiation is weak. This suggests high levels of gene-flow over short to middle distances.     

Post‐glacial migration lag restricts range filling of plants in the European Alps

Aim To evaluate the effect of post‐glacial migration lags on the current distribution of Alpine plants and the factors responsible for possible range‐filling differences among species. Location Austrian Alps. Methods We used species distribution models to predict environmentally suitable sites for 183 Alpine plants at a fine spatial resolution (100 × 100 m²). We overlaid these predictions with independent mapping data (3′× 5′) and calculated the extent to which species fill their potential ranges at this coarser grain based on several different approaches. Moreover, we correlated range‐filling estimates with the magnitude of improvement of distribution models when using the distance to putative glacial refugia as an additional independent variable. Finally, we compared species‐specific range‐filling estimates with traits related to dispersal capacity and competitive ability of these species as well as with characteristics of their habitats. Results Even under a conservative approach, incomplete range filling appears common, with 46% and 31% of the species studied occurring in less than 75% and 50% of their predicted suitable ranges, respectively. Proximity to glacial refugia generally accounts for a lower percentage of the deviance in species distribution data (0–20%, mean 4%) than environmental variables (9–57%, mean 27%). However, its importance correlates closely and negatively with the calculated range‐filling estimates. Range filling significantly increases with the dispersal capacity of a species' propagules and the breadth of its altitudinal niche. Calcicolous species have lower range filling than silicicolous plants and substrate generalists. Conclusions Our results suggest that the current ranges of many Alpine plants are still shaped by delayed Holocene recolonization of suitable sites. Hence, long‐term migration lags also affect plant distribution in mountainous areas, at least on regional scales. These findings question whether high mountain floras will be able to track the expected rapid, climate change driven shifts in habitat.     

Floristic composition and endemism pattern of vascular plants in Ethiopia and Eritrea

Traditional attempts to delineate floristic regions are typically based on the qualitative analysis of species distribution, often ignoring the phylogenetic relationships among their taxa. Ethiopia and Eritrea are in the Horn of Africa, known as one of the world's biodiversity hotspots. We quantitatively classified the flora of Ethiopia and Eritrea into meaningful geographical units by analyzing the taxonomic and phylogenetic β‐diversity at genera, total species, and endemic species levels at a scale of 0.5° × 0.5° grid cells. Hierarchical clustering was used to quantitatively delimitate the flora and analysis of similarities was used to test the significant difference between the derived groups in taxonomic composition and phylogenetic relatedness. In total, two floristic subprovinces, five floristic districts, and 13 floristic subdistricts, as well as three centers of species endemism associated with three floristic subdistricts were identified. Our results also showed that the species diversity, endemism, and turnover of the highlands in Ethiopia and Eritrea were much higher than the lowlands, indicating that the floristic differences are closely related to the topography of the East African Rift. In this study, we provided a scientific framework for the composition and relationships of the floristic units in the Horn of Africa, and similarly provided a scientific basis for better conservation of the diversity in this region.     

Dragging into the open: the polythetic nature of areas of endemism

Areas of endemism represent territories (no matter the size) of non-random overlap in the geographic distribution of two or more taxa, reflecting a common spatial history of these taxa. The common spatial history is a result of different processes that connect areas of endemism to evolutionary theory. Numerous and diverse definitions of areas of endemism have been proposed. All of them have used as the conceptual foundation of the definition a certain degree of non-random congruence of geographic distribution amongst at least two taxa. ‘Certain degree’ means that geographic congruence does not demand complete agreement on the boundaries of those taxa's distributions at all possible scales of mapping. The words ‘certain degree’ mask the polythetic nature of areas of endemism. The polythetic characterization of areas of endemism implies that each locality of the study area has a large number of a set of species. Each species of this set is present in many of those localities and, generally, none of those species is present in every locality of the area. The converse will be a monothetic nature of areas of endemism where a taxon or group of taxa is present in all the localities of the study area. We propose here that the expansion of the definition of areas of endemism, including their polythetic characterization, will improve understanding of large biogeographic areas such as realms, regions, provinces, and districts, and will increase the scientific content (e.g., predictive capability and explanatory power) of areas of endemism.     

高黎贡山南段种子植物区系的特有现象

高黎贡山南段是全球生物多样性研究热点地区之一,迄今计有野生种子植物192科878属2 807种.这一区域不仅植物的种类丰富,而且特有现象较为显著.其中东亚特有科8个、中国特有属17个、中国特有种1 085个、云南特有种305个、高黎贡山南段特有种82个.特殊地质历史和生态环境,使得高黎贡山南段既保留了许多古老特有成分,又孕育了许多新生特有成分.     

Northern Hemisphere origin,transoceanic dispersal,and diversification of Ranunculeae DC. (Ranunculaceae) in the Cenozoic

Aim The role of dispersal versus vicariance for plant distribution patterns has long been disputed. We study the temporal and spatial diversification of Ranunculeae, an almost cosmopolitan tribe comprising 19 genera, to understand the processes that have resulted in the present inter‐continental disjunctions. Location All continents (except Antarctica). Methods Based on phylogenetic analyses of nuclear and chloroplast DNA sequences for 18 genera and 89 species, we develop a temporal–spatial framework for the reconstruction of the biogeographical history of Ranunculeae. To estimate divergence dates, Bayesian uncorrelated rates analyses and four calibration points derived from geological, fossil and external molecular information were applied. Parsimony‐based methods for dispersal–vicariance analysis (diva and Mesquite ) and a maximum likelihood‐based method (Lagrange ) were used for reconstructing ancestral areas. Six areas corresponding to continents were delimited. Results The reconstruction of ancestral areas is congruent in the diva and maximum likelihood‐based analyses for most nodes, but Mesquite reveals equivocal results at deep nodes. Our study suggests a Northern Hemisphere origin for the Ranunculeae in the Eocene and a weakly supported vicariance event between North America and Eurasia. The Eurasian clade diversified between the early Oligocene and the late Miocene, with at least three independent migrations to the Southern Hemisphere. The North American clade diversified in the Miocene and dispersed later to Eurasia, South America and Africa. Main conclusions Ranunculeae diversified between the late Eocene and the late Miocene. During this time period, the main oceanic barriers already existed between continents and thus dispersal is the most likely explanation for the current distribution of the tribe. In the Southern Hemisphere, a vicariance model related to the break‐up of Gondwana is clearly rejected. Dispersals between continents could have occurred via migration over land bridges, such as the Bering Land Bridge, or via long‐distance dispersal.     

Molecular evidence for glacial refugia of mountain plants in the European Alps

Many mountain ranges have been strongly glaciated during the Quaternary ice ages, and the locations of glacial refugia of mountain plants have been debated for a long time. A series of detailed molecular studies, investigating intraspecific genetic variation of mountain plants in the European Alps, now allows for a first synopsis. A comparison of the phylogeographic patterns with geological and palaeoenvironmental data demonstrates that glacial refugia were located along the southwestern, southern, eastern and northern border of the Alps. Additional glacial refugia were present in central Alpine areas, where high-elevation plants survived the last glaciation on ice-free mountain tops. The observed intraspecific phylogeographies suggest general patterns of glacial survival, which conform to well-known centres of Alpine species diversity and endemism. This implies that evolutionary or biogeographic processes induced by climatic fluctuations act on gene and species diversity in a similar way.