Hidden areas of endemism: Small units in the South-eastern Neotropics

This study aimed to establish if the Lower Río de la Plata Basin (LRPB) wetlands can be considered a biogeographic unit. The species of this area were compiled and segregated according to the habitat, selecting only 87 endemic taxa restricted to the LRPB and linked to wetlands. Distributional data of species obtained from the literature, web databases, biological collections, and field trips were georeferenced. The areas of endemism were established as those areas where the distribution of two or more taxa overlaps in groups of rivers’ sections with geographic continuity and were tested with a cluster analysis. This congruence is due to ecological, geomorphological, and historical factors. Four areas of endemism were found: a broad area that comprises the whole study area (Riverine district), which is divided into three nested smaller areas (Paraguay–Paraná Flooding Valleys, Uruguay Basin, and Paraná Delta subdistricts). Then, we analysed 170 taxa distributions to evaluate the relationship between the study area and the neighbouring biogeographic units. According to the results, the study area belongs to the Paraná biogeographic province. Some areas of endemism are hidden inside broader areas and are hardly detected with the currently used biogeographic grid-methods. We propose to combine the information about ecological requirements of each taxon with its georeferenced records to estimate their areas of distribution as a primary step for searching areas of endemism in intracontinental studies.     

Patterns of endemism in south-eastern Pacific benthic polychaetes of the Chilean coast

Aim In this study we evaluate patterns of endemism for benthic polychaete species along the southeastern Pacific coast of Chile. Our goals were (1) to describe latitudinal gradients of endemism and identify areas of high endemism, (2) to evaluate the effect of biogeographical limits on endemism patterns, and (3) to evaluate indirectly the role played by evolutionary dynamics on patterns of endemism. Location South‐eastern Pacific coast of Chile, ranging from Arica (18° S) to Cape Horn (56° S). Methods We used a list of 178 species of endemic, shallow benthic polychaetes to evaluate patterns of endemism. Parsimony analysis of endemicity (PAE) and the endemism index (EI) were used to evaluate hierarchical relationships of endemism between different latitudinal bands, and to identify areas with high degrees of endemism and differences in endemism. We evaluated the effect of biogeographical limits on endemic polychaete fauna by testing for the existence of geometric constraints (mid‐domain effect). The role of evolutionary dynamics on latitudinal patterns of endemism was evaluated with nestedness analysis (NA) using the temperature index. Results The PAE analysis indicated two large, separate areas of endemism: (1) the northern area between 18° S and 38° S, and (2) the southern area between 39° S and 56° S. The endemism index showed a maximum value (32 species) around 39°–41° S. Species‐richness curves of each 3° band of latitude showed a clear mid‐domain effect (69%), but the two maximum points of species richness at mid‐latitudes (36° S to 38° S and 39° S to 41° S) did not correspond to the mid‐domain peak in species richness, presenting a greater number of species than expected by the mid‐domain effect. The nestedness analysis showed that the number of genera reaches a maximum of 70 at mid‐latitudes (36°–41° S), decreasing towards both the northern and southern areas. The spatial distribution of the entire data set of endemic species showed a nested pattern (T° = 24.5°, P < 0.0001). Main conclusions Our results strongly support the existence of a latitudinal gradient of endemism for benthic polychaete species along the Chilean coast. The shape of this gradient is clearly non‐linear, with a marked peak of endemism occurring at mid‐latitudes (36°–41° S, endemism hotspot), which also corresponds to a peak in species richness. Furthermore, this hotspot is the midpoint separating two distinct areas of endemism to the north and south. We suggest that the observed pattern of endemism for benthic polychaete taxa of the Chilean coast can be explained by a combination of geometric constraints and historical mechanisms, such as the processes that affected the Chilean coast during the Neogene (e.g. ENSO, oxygen minimum zone, glaciations).     

Levels of endemism are not necessarily biased by the co-presence of species with different range sizes: a case study of Vilenkin and Chikatunov's models

Aim  To illustrate problems in the methods proposed by B. Vilenkin and V. Chikatunov to study levels of endemism and species–area relationships.
Location  The study used data on the distribution of tenebrionid beetles (Coleoptera, Tenebrionidae) on the Aegean Islands (Greece).
Methods  A total of 32 islands and 170 taxa (species and subspecies) were included in this study. Levels of endemism were evaluated both as the proportion of endemic taxa, and according to the methods proposed by Vilenkin and Chikatunov, which are based on the number of non-endemic taxa and various relationships with area. A model of the species–area relationship proposed by these authors was also analysed.
Results  The number of endemic taxa was positively correlated with the number of taxa with different distribution types, but this positive correlation did not influence the estimation of the level of endemism. In fact, the commonly used estimate of endemicity as a percentage was strongly correlated with the endemism values calculated according to the method of Vilenkin and Chikatunov. The usual power function fitted the species–area relationship as well as the most complicated method of Vilenkin and Chikatunov.
Main conclusions  As hypothesized by Vilenkin and Chikatunov, the number of endemic taxa was influenced both by the number of taxa of other biogeographical ranks, and by an island's area. However, explanations for the positive relationship between the number of endemic taxa and taxa of different biogeographical ranks are equivocal. Importantly, this relationship did not necessarily influence the level of endemism, which could be expressed adequately by percentages. The method proposed by Vilenkin and Chikatunov to estimate the species–area relationship cannot be clearly justified on theoretical grounds and is of questionable practical utility.     

The biogeography of tropical reef fishes: endemism and provinciality through time

The largest marine biodiversity hotspot straddles the Indian and Pacific Oceans, driven by taxa associated with tropical coral reefs. Centred on the Indo‐Australian Archipelago (IAA), this biodiversity hotspot forms the ‘bullseye’ of a steep gradient in species richness from this centre to the periphery of the vast Indo‐Pacific region. Complex patterns of endemism, wide‐ranging species and assemblage differences have obscured our understanding of the genesis of this biodiversity pattern and its maintenance across two‐thirds of the world's oceans. But time‐calibrated molecular phylogenies coupled with ancestral biogeographic estimates have provided a valuable framework in which to examine the origins of coral reef fish biodiversity across the tropics. Herein, we examine phylogenetic and biogeographic data for coral reef fishes to highlight temporal patterns of marine endemism and tropical provinciality. The ages and distribution of endemic lineages have often been used to identify areas of species creation and demise in the marine tropics and discriminate among multiple hypotheses regarding the origins of biodiversity in the IAA. Despite a general under‐sampling of endemic fishes in phylogenetic studies, the majority of locations today contain a mixture of potential paleo‐ and neo‐endemic fishes, pointing to multiple historical processes involved in the origin and maintenance of the IAA biodiversity hotspot. Increased precision and sampling of geographic ranges for reef fishes has permitted the division of discrete realms, regions and provinces across the tropics. Yet, such metrics are only beginning to integrate phylogenetic relatedness and ancestral biogeography. Here, we integrate phylogenetic diversity with ancestral biogeographic estimation of lineages to show how assemblage structure and tropical provinciality has changed through time.     

Quantity versus quality: Endemism and protected areas in the temperate forest of South America

Abstract Identification of biodiversity hotspots is essential to conservation strategies aimed at minimizing the possibility of losing half of the world's species in the next 50 years. The aims of the present study were: (i) to locate and designate zones of endemism in the temperate forest of South America; and (ii) to compare the distribution of these areas with the distribution of existing protected areas in this habitat type. Endemism areas were determined by using parsimonious analysis of endemism, which identified zones of endemism on the basis of sets of endemic species that were restricted to two or more study areas. We used distribution information for five unrelated taxa (ferns, trees, reptiles, birds and mammals) to provide more reliable results and patterns than would work with only a single taxon or related taxa. The northern part of this region has high endemism for all of the taxa considered in this study. We demonstrate that although the temperate forest of South America has more than 30% of its area under some type of protection, correlation between protected areas and the areas of endemism is remarkably low. In fact, less than 10% of protected areas are situated in areas that have the greatest value for conservation (i.e. high endemism). Under the current strategy, biodiversity within South America's temperate forest is in danger despite the large amount of protected area for this forest type.     

Contribution à la connaissance de la végétation rupicole des hautes vallées de l'ubaye et de l'ubayette. (Alpes cottiennes)

Summary The study of the saxatile vegetation of the alpine zone in the Upper Ubaye Valley shows a very sharp demarcation from a phytogeographical point of view.The saxatile groupings on limestone show septentrional affinities. It is interesting to note that the associations described have at least as much relationship to the Swiss Alp groups as to those of The Maritime Alps in fact closer.In our opinion, the explanation is not to be found in very different climatical or ecological considerations but in the presence of important barrier-mountains which have deprived the Upper Ubaye Valley of alien postglacial immigrants.There endemism is apparent but less important than in the neighbouring zone of The Maritime Alps; here it concerns surviving endemic species proceeding from an old native preglacial or interglacial element.These characters are to be found more in the northern areas of Queyras, Embrunais and Briançonnais, regions which have an incontestable relationship with our district.     

Avian distribution patterns in the Guiana Shield: implications for the delimitation of Amazonian areas of endemism

Aim To develop a methodology for defining the boundaries of the Guianan area of endemism using complementary approaches that include GIS tools, multivariate statistics and analyses of physical barriers in the distribution patterns of an entire endemic avifauna. As a case study, I used the distribution patterns of lowland terra firme forest birds. Location Guiana Shield, northern South America. Methods I identified Guianan endemics using the ornithological literature, subsequently gathering distributional data for these taxa using mainly museum collections and my own fieldwork in the region. I used these distributional data to map the spatial patterns of endemicity in the region and to compare distributions across taxa. I employed community composition data from 34 localities from throughout the Guiana Shield to identify spatial patterns of clustering using an ordination analysis (non‐metric multidimensional scaling), and to recognize the region’s main biogeographical barriers for birds using Monmonier’s algorithm. Results At least 88 avian taxa are restricted to the terra firme forests of the Guianan area of endemism, which is roughly delimited by the Amazon, Negro and Branco rivers. These large rivers, however, are not the only boundaries. I identified seven additional barriers, including medium‐sized rivers, non‐forested areas and mountains, which also contribute to delimiting the area of endemism. Within the endemic avifauna, I identified three distinct distribution patterns. The ordination analysis shows the presence of two distinct avifaunas within the Guiana Shield. Main conclusions Although the proposed boundaries of the Guianan area of endemism are consistent with previously postulated configurations, this study reveals a more complex delimitation than formerly recognized, highlighting the importance of several landscape features besides large rivers, and the existence of three distinct distributional patterns within one endemic avifauna. The Branco/Negro interfluvium, often included within this area of endemism, actually represents a transition zone between two distinct avifaunas. The longstanding view of the Amazon Basin as a mosaic of parapatric areas of endemism delimited by major rivers appears to be an oversimplification, at least for the Guiana Shield. This finding suggests the need for more rigorous approaches to re‐evaluate the traditional boundaries of such areas.     

Haplotype richness in refugial areas: phylogeographical structure of <Emphasis Type="Italic">Saxifraga callosa</Emphasis>

This paper illustrates the phylogeographical structure of Saxifraga callosa in order to describe its genetic richness in refugial areas and to reconstruct its glacial history. S. callosa is a species spread throughout south-east France and Italy with a high distribution in the Maritime Alps. Four chloroplast microsatellite and AFLP markers were analyzed in populations of S. callosa. The size variants of all tested loci amount to 11 different haplotypes. Intrapopulational haplotype variation was found in two of the populations analyzed: on the Mt. Toraggio in the Maritime Alps, and in the Apuan Alps. On the other hand, no intrapopulational variation was found in 25 populations, most of which were sampled from isolated areas. Analysis of the haplotype distribution showed that population subdivision across all populations was high (G _ST = 0.899). Moreover, its genetic structure was studied using AMOVA and STRUCTURE analysis. The study legitimated inferred conclusions about the phylogeographical structure of the species and identified centers of diversity. Considerations concerning genetic structure and divergence among three major clades (Maritime Alps, Apuan Alps and Apennines), the patchy distribution of haplotypes, and the high number of private haplotypes support the proposal that S. callosa survived in some refugia within the Italian Peninsula refugium, and that mainly northern populations of refugia were involved in postglacial recolonization.     

Humans and other animals and the plants they ingest

Primula allionii is endemic to a tiny area of the Maritime Alps and has one of the narrowest distribution ranges in this hotspot of biodiversity. Phylogeographical patterns in P. allionii were studied using plastid DNA markers and dominantly inherited markers (AFLP and ISSR) to verify any admixture between P. allionii and the sympatric P. marginata and to detect the phylogeographical history of the species. Morphometric measurements of flowers and admixture analysis support the hypothesis that hybridization occurs in nature. Species distribution models using two climate models (CCSM and MIROC) suggested a reduction in habitat suitability during cold periods. Phylogeographical analysis suggested an old allopatric divergence during the mid‐Pleistocene transition (about 0.8 Mya) without recolonization/contraction cycles. The Alps watershed does not act as a strong barrier between the two main areas of the distribution range, and moderate gene flow by pollen seems to create the admixture recorded among the stands. According to our results, the persistence of P. allionii throughout the Ice Age appears to be linked to the capacity of the Maritime Alps to provide a wide diversity of microhabitats consistent with the recent biogeographical pattern proposed for the Mediterranean Basin. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 637–653.     

Distribution range and ecological niche of Primula marginata Curtis (Primulaceae)

The distribution range of Primula marginata Curtis (Primulaceae) has never been fully characterized. In the present study, authors did a revision of the distribution range using herbaria material, database records and in situ populations' check-up. P. marginata was confirmed extending from Cottian to Maritime and Ligurian Alps, with few outlier occurrences in the northern Apennines. The localities previously reported from northern Piedmont (Val d'Ossola) were not confirmed. Maximum entropy model (Maxent) was used to simulate the potential distribution of P. marginata under current climate conditions. According to the distribution modelling performed, the species prefers rocky calcareous habitats mainly at high elevations, with abundant precipitation, but low moisture retention at soil level and marked temperature range between winter and summer seasons. The potential distribution area drawn by Maxent seemed to describe P. marginata at its maximum extension, and any future climate changes might cause limitations for the survival of the species.     

Parsimony analysis of endemicity and analysis of endemicity: A fair comparison

Parsimony Analysis of Endemicity (PAE) has been compared with other methods regarding its performance to identify areas of endemism. It is frequently compared with the Analysis of Endemicity (AE), which seems to perform better than PAE to identify these areas. Here I compare PAE and AE considering the sympatric taxa diagnosed as endemic, being as strictly close as possible to sympatry, and using previously published data of Sciobius (Coleoptera: Curculionidae). AE identified more candidate areas of endemism than PAE, but the number of highly restricted endemic taxa to these areas was insufficient to support them as areas of endemism. Considering strictly sympatry (homopatry), PAE performed better than AE; however, both methods may identify areas with some grade of sympatry, but the recognition of which areas constitute real areas of endemism in the strict sense depends on the interpretation of the researcher.     

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.     

Phytogeographic analysis of taxa endemic to the Yucatán Peninsula using geographic information systems, the domain heuristic method and parsimony analysis of endemicity

Abstract. The distribution of plant taxa endemic to the Yucatán Peninsula was studied using Parsimony Analysis of Endemicity (PAE). The known distribution of 162 endemic plant taxa was plotted and the DOMAIN method together with environmental data were used to model the potential distribution for each taxon. The Peninsula was divided into a grid of quarter‐degree cells for the purpose of identifying distribution patterns. A total of 294 cells were analysed using known collection records and potential distribution of endemic taxa data. Two data matrices were constructed, a matrix of known distribution and a matrix of both the known and potential distribution. The two matrices were included in the PAE to identify areas of endemism. The areas determined with the known distribution were restricted and almost half of them remained unresolved, whereas with the potential distribution, approximately 90% of the cells were assigned to any one of the endemicity areas. Four endemism areas were identified: the Yucatán dry zone, Yucatán, El Petén and Belize. The areas of Yucatán and El Petén could be explained by current and Pleistocene climatic conditions and their congruence with other biological groups. Analysis of the potential distribution identified areas with patterns that share current environmental characteristics and a palaeoclimate history. Potential distribution modelling can eliminate uncertainties in biogeographical analysis caused by lack of data distribution and sample variation and produce information about the relationships between areas and taxa as well as the environmental affinities of taxa.     

A spider diversity model for the Caucasus Ecoregion

Precise information on spatial patterns of species richness and endemic species distribution is important for effective species conservation. In the Caucasus Ecoregion such information is virtually non-existent for invertebrate taxa. Using occurrence data from a large database we calculated species distribution models with the GARP algorithm for 471 spider species to visualize the diversity distribution of spider species in this region. Overall species diversity was highest in mountain forests of the North Caucasus, east-central Georgia, the southern slopes of the eastern Great Caucasus and south-east Azerbaijan. A regression tree analysis Chi squared automatic interaction detector method revealed the mean temperature of the driest quarter and precipitation parameters to be the main environmental factors shaping these patterns. Diversity of endemic species was correlated with overall species diversity but hotspots of endemic species (10+ percent of all species) exists in high-mountain areas, suggesting post-glacial speciation events in the high mountains as the main sources of high endemism in Caucasus. Further information on the spatial distribution of species diversity of invertebrate taxa in the Caucasus Ecoregion is needed to improve conservation efforts in this biodiversity hotspot.     

Parsimony analysis of the distribution pattern of Japanese primary freshwater fishes, and its application to the distribution of the bagrid catfishes

The geographic distribution of Japanese primary and some secondary freshwater fishes was analyzed using parsimony analysis of endemicity (PAE). Analysis of 73 taxa (species and intraspecific forms) on the four main islands of Japan (divided into 25 geographic areas), produced 44 most parsimonious cladograms. In all of the latter, a total of eight single and compound areas were recognized as endemic areas in nested relationships. The area cladograms showed Japan as comprising a middle-eastern Hokkaido area plus southern areas, the latter containing mainly a northeastern-Honshu endemic area and more heterogeneous southwestern areas, including four endemic areas (western Kyushu, southeastern Chugoku, middle Kinki and Tokai around Ise Bay) and several peripheral areas. Some patterns, e.g., the distinction in fauna across Fossa Magna, noted by previous studies, were supported by these results. Even though the analysis had some problems (e.g., not all geographic ranges of taxa could be included), it provided evidence for the detection of general distribution patterns, because the relationships or similarities among areas were clearly defined by shared taxa. To demonstrate the historical implications of the analysis, the allopatric distribution of, four bagrid catfishes was reconsidered in the area cladogram. The general pattern implied secondary extinction ofPseudobagrus nudiceps around lse Bay, which was in keeping with the fossil record.     

Dispersal limitations, rather than bottlenecks or habitat specificity, can restrict the distribution of rare and endemic rainforest trees

Despite their narrow distribution, Australian rainforests still contain considerable levels of diversity and include many ancient, but often rare, lineages. Very little is known about the general biology of rainforest species, yet their long-term management depends on a better understanding of the main factors leading to rarity. For instance, are they highly endemic taxa, at the early stages of expansion, nearing the end of a period of decline, or persisting at low numbers over the long term? In this study we combine molecular, environmental, and ecological data to identify the factors responsible for the narrow distribution of a paleoendemic rainforest tree: Elaeocarpus sedentarius (Elaeocarpaceae). Between-population and between-generation comparisons of genetic diversity across all known populations of E. sedentarius show evidence of mutation-drift equilibrium rather than evidence of a recent bottleneck. Similarly, floristic and environmental data negate the hypothesis of rarity as a consequence of highly specialized habitat requirements. Instead, genetic structure and the available ecological data support the hypothesis of dispersal limitation as the main cause of endemism and that the species may have attained genetic equilibrium without realizing its full niche potential. We suggest that these factors are likely to explain narrow endemism in a broader range of taxa.     

Disturbance,seeds, restoration,and the importance of experiments and long‐term observations: the Editors' Award for 2005

Question: Can the geographic patterning of endemic plant species inform reserve selection in a region of high endemism? Location: The Southeastern Coastal Plain of North America, focusing primarily on the imperiled longleaf pine (Pinus palustris P. Miller) ecosystem. Methods: We documented the high level of plant endemism in the region, and characterized the endemic taxa into distributional subregions. Results: A total of 1630 plant taxa are endemic to the Coastal Plain, a large proportion of which are endemic to phytogeographical subregions within the Coastal Plain, with particularly large numbers of narrow endemics occurring in the East Gulf Coastal Plain and Florida Peninsula. Conclusions: This pattern of local endemism presents challenges in conserving the full biota of the region: a reserve system focusing on few and large conservation areas has theoretical benefits for long‐term management and viability, but will fail to capture many local endemics. We propose that the dispersed distribution of endemic species will require a mixture of large core reserves and smaller satellite reserves.