Species richness,endemism, and conservation of American tree ferns (Cyatheales)

Analyses of richness and endemism of Cyatheales (tree ferns) in tropical America were performed and evidence of a diversity gradient is presented. For this, the occurrence ranges of 239 species were plotted into a 5° × 5° grid-cell map and then analyzed using species richness and endemism indices. Here we show that species richness and endemism are not distributed randomly over the landscape, but do aggregate into defined regions of high diversity in tropical America: the northern Andes, lower Central America, upper Central America and Mexico, the Guyana Highlands, southeastern Brazil, and the Antilles. These distributional patterns are congruent with the geographical distribution of cloud forest, which in turn is determined by topography, high humidity, and persistent cloud immersion. The mountain regions of tropical America, especially the cloud forests, harbour most of the species of American Cyatheales and have high levels of habitat loss and climatic fragility. Conservation policies for Cyatheales are centred on the local use and trade of many tree fern species, but none such policies focus on cloud forest habitat loss. This makes tree ferns a critically endangered group of plants. In the face of the current environmental crisis and global climate change, the presence of Cyatheales in these regions sounds the alarm on their conservation priorities.     

Habitat suitability models to make conservation decisions based on areas of high species richness and endemism

Biodiversity positively relates with the provisioning of ecosystem services and preserving areas with elevated diversity of highly-functional species could help to ensure human well-being. Most studies addressed to make these decisions use maps relying on species occurrences, where sites containing several species are proposed as priority conservation areas. These maps, however, may underestimate species richness because of the incompleteness of occurrence data. To improve this methodology, we propose using habitat suitability models to estimate the potential distribution of species from occurrence data, and later shaping richness maps by overlapping these predicted distribution ranges. We tested this proposal with Mexican oaks because they provide several ecosystem services and habitat suitability models of species were calibrated with MaxEnt. We used linear regressions to compare the outputs of these predictive maps with those of maps based on species occurrences only and, for both mapping methods, we assessed how much surface of sites with elevated richness and endemism of oaks is currently included within nature reserves. Both mapping methods indicated that oak species are concentrated in mountain regions of Mexico, but predictive maps based on habitat suitability models indicated higher oak richness and endemism that maps based on species occurrences only. Our results also indicated that nature reserves cover a small fraction of areas harboring elevated richness and endemism of oaks. These results suggest that estimating richness across extensive geographic regions using habitat suitability models quickly provides accurate information to make conservation decisions for highly-functional species groups.     

Seamounts: centres of endemism or species richness for ophiuroids?

Aim To test the hypotheses that seamounts exhibit high rates of endemism and/or species richness compared to surrounding areas of the continental slope and oceanic ridges. Location The south‐west Pacific Ocean from 19–57° S to 143–171° E. Methods Presence/absence museum data were compiled for seamount and non‐seamount areas at depths between 100 and 1500 m for the Ophiuroidea (brittle‐stars), an abundant and speciose group of benthic invertebrates. Large‐scale biogeographical gradients were examined through multivariate analyses at two spatial scales, at the scale of seamounts (< 1° of latitude/longitude) and regions (5–9°). The robustness of these patterns to spatially inconsistent sampling effort was tested using Monte Carlo‐style simulations. Levels of local endemism and species richness over numbers of samples were compared for seamount and non‐seamount areas using linear regressions. Non‐seamount populations were randomly generated from areas and depth ranges that reflected the typical sampling profile of seamounts. Results Seamount ophiuroid assemblages did not exhibit elevated levels of species richness or narrow‐range endemism compared with non‐seamount areas. Seamounts can exhibit high overall species richness for low numbers of samples, particularly on seamounts supporting a dense coral matrix, but this does not increase with additional sampling at the rates found in non‐seamount areas. There were relatively few identifiable seamount specialists. In general, seamount faunas reflected those found at similar depths in surrounding areas, including the continental slope. Seamount and non‐seamount faunas within the study area exhibited congruent latitudinal and bathymetric species turnover. Main conclusions Seamount faunas were variable for ophiuroid faunal composition, species richness and narrow‐range endemism, reflecting their environmental diversity and complex history. The continental slope was also variable, with some areas being particularly species rich. Broad geomorphological habitat categories such as ‘seamounts’ or ‘continental slope’ may be at the wrong scale to be useful for conservation planning.     

Areas of endemism and distribution patterns for Neotropical Piper species (Piperaceae)

Aim The study aimed to establish areas of endemism and distribution patterns for Neotropical species of the genus Piper in the Neotropical and Andean regions by means of parsimony analysis of endemicity (PAE) and track‐compatibility analysis. Location The study area includes the Neotropical region and the Northern Andean region (Páramo‐Punan subregion). Methods We used distribution information from herbarium specimens and recent monographic revisions for 1152 species of Piper from the Neotropics. First, a PAE was attempted in order to delimit the areas of endemism. Second, we performed a track‐compatibility analysis to establish distribution patterns for Neotropical species of Piper. Terminology for grouping Piper is based on recent phylogenetic analyses. Results The PAE yielded 104 small endemic areas for the genus Piper, 80 of which are in the Caribbean, Amazonian and Paranensis subregions of the Neotropical region, and 24 in the Páramo‐Punan subregion of the Andean region. Track‐compatibility analysis revealed 26 generalized tracks, one in the Páramo‐Punan subregion (Andean region), 19 in the Neotropical region, and six connecting the Andean and Neotropical regions. Both the generalized tracks and endemic areas indicate that distribution of Piper species is restricted to forest areas in the Andes, Amazonia, Chocó, Central America, the Guayana Shield and the Brazilian Atlantic coast. Main conclusions Piper should not be considered an Andean‐centred group as it represents two large species components with distributions centred in the Amazonian and Andean regions. Furthermore, areas of greater species richness and/or endemism are restricted to lowland habitats belonging to the Neotropical region. The distribution patterns of Neotropical species of Piper could be explained by recent events in the Neotropical region, as is the case for the track connecting Chocó and Central America, where most of the species rich groups of the genus are found. Two kinds of event could explain the biogeography of a large part of the Piper taxa with Andean–Amazonian distribution: pre‐Andean and post‐Andean events.     

Global patterns in species richness of pelagic seabirds: the Procellariiformes

Quantitative information on large scale spatial patterns of biodiversity remains poor, especially for pelagic systems. In this paper the regional diversity of procellariiforms is mapped worldwide at the species level. These seabirds do not display a conventional latitudinal gradient of decreasing species richness towards high latitudes, but rather are most speciose between 37° and 59°S in all ocean basins. Based on data for foraging ranges, areas with the highest species richness and the most species with smaller range sizes are all found in the vicinity of New Zealand and its sub-Antarctic islands. In contrast, data for breeding ranges show islands in the southern Indian and Atlantic oceans to have the highest number of breeding species, while these islands and New Zealand have the most species with smaller range sizes. No northern hemisphere regions are amongst the top ten grid cells for foraging and breeding species richness, although Hawaii has the highest species richness of procellariilforms north of the equator. Northern Baja California. Madeira, the Canary islands, and the west coast of South America are all important sites of narrow endemism in the northern hemisphere. High species richness and narrow endemism coincide with areas of significant longline fishing activity in the southern hemisphere. Near-minimum sets based on one and three representations demonstrate that if all procellariiform species are to be retained, large areas of the ocean and almost all breeding sites require conservation.     

Phylogenetic endemism of the orchids of Megamexico reveals complementary areas for conservation

Orchid diversity provides a unique opportunity to further our understanding of biotic and abiotic factors linked to patterns of richness, endemism, and phylogenetic endemism in many regions. However, orchid diversity is consistently threatened by illegal trade and habitat transformation. Here, we identified areas critical for orchid conservation in the biogeographic province of Megamexico. For this purpose, we evaluated orchid endemism, phylogenetic diversity, and phylogenetic endemism within Megamexico and characterized orchid life forms. Our results indicate that the majority of the regions with the highest estimates of endemism and phylogenetic endemism are in southern Mexico and northern Central America, mostly located on the Pacific side of Megamexico. Among the most important orchid lineages, several belong to epiphytic lineages such as Pleurothallidinae, Laeliinae and Oncidiinae. We also found that species from diverse and distantly related lineages converge in montane forests where suitable substrates for epiphytes abound. Furthermore, the southernmost areas of phylogenetic diversity and endemism of Megamexico are in unprotected areas. Thus, we conclude that the most critical areas for orchid conservation in Megamexico are located in southern Mexico and northern Central America. We recommend that these areas should be given priority by the Mexican system of natural protected areas as complementary conservation areas.     

Influences of interpolation of species ranges on elevational species richness gradients

Interpolation of species ranges has been a common approach to compensate for the unevenness or incompleteness in sampling effort in studies of geographic species richness gradients. However, potential biases introduced by this estimation method remain unclear. Here, we presented an explicit examination of the influences of one‐dimensional interpolation on elevational species richness gradients, and discussed potential causes and processes of these influences. We conducted intensive surveys of birds along the elevational gradients of the Ailao Mountains, southwestern China, and compared richness patterns based on interpolation with raw data as well as estimated data from rarefaction and Chao1 non‐parametric estimator; we also compared results of multiple linear regressions and hierarchical partitioning analyses explaining these four measures of richness. Actual evapotranspiration (AET) and the mid‐domain effect (MDE) were highly correlated and separately provided a good potential explanation for the unimodal richness pattern in the Ailao Mountains, with modifying and suppressive effects of other variables such as area. Interpolation consistently and significantly increased the effects of AET/MDE, while it reduced contributions of area and human disturbance. Our results demonstrated that while compensating for biases in sampling effort, interpolation may also spuriously fill genuine distribution gaps, and tend to underestimate the effects of the non‐monotonic or discontinuous influencing factors that are responsible for these gaps, and overestimate the effects of other factors actually suppressed by these factors. These influences were most strong for species with relatively medium elevational ranges. We conclude that at the regional scale, interpolation method is a potential source of bias in identifying and explaining species richness gradients and should be used with careful consideration. It may be advantageous to adopt other robust estimation methods besides interpolation to gain a more accurate assessment of species richness and a more objective understanding of their underlying mechanisms.     

Patterns of speciation are similar across mountainous and lowland regions for a Neotropical plant radiation (Costaceae: Costus)

High species richness and endemism in tropical mountains are recognized as major contributors to the latitudinal diversity gradient. The processes underlying mountain speciation, however, are largely untested. The prevalence of steep ecogeographic gradients and the geographic isolation of populations by topographic features are predicted to promote speciation in mountains. We evaluate these processes in a species-rich Neotropical genus of understory herbs that range from the lowlands to montane forests and have higher species richness in topographically complex regions. We ask whether climatic niche divergence, geographic isolation, and pollination shifts differ between mountain-influenced and lowland Amazonian sister pairs inferred from a 756-gene phylogeny. Neotropical Costus ancestors diverged in Central America during a period of mountain formation in the last 3 million years with later colonization of Amazonia. Although climatic divergence, geographic isolation, and pollination shifts are prevalent in general, these factors do not differ between mountain-influenced and Amazonian sister pairs. Despite higher climatic niche and species diversity in the mountains, speciation modes in Costus appear similar across regions. Thus, greater species richness in tropical mountains may reflect differences in colonization history, diversification rates, or the prevalence of rapidly evolving plant life forms, rather than differences in speciation mode.     

Application of parsimony analysis of endemicity in Amazonian biogeography: an example with primates

The distributions of 51 non-human primate species are used for Parsimony Analysis of Endemicity (PAE) to determine the relationships among 14 interfluvial regions in the Amazon basin, South America. Two most parsimonious cladograms were found. The strict consensus tree of these cladograms suggests an early separation between Lower Amazonia (eastern) and Upper Amazonia (western). The major clusters of interfluvial regions identified in the PAE cladogram are congruent with the areas of endemism delimited for birds. When interfluvial regions are converted into avian areas of endemism, the PAE cladogram is congruent with one of the two general areas cladograms suggested for Amazonia based on phylogenies of several clades of forest birds. Our analysis suggests that PAE can be used as a tool to objectively identify areas of endemism at an intra-continental scale as well as to make historical inferences. However, the value of a PAE cladogram in this latter application should be always evaluated by congruence with area cladograms built upon cladistic biogeography procedures.     

Patterns in species richness and endemism of European freshwater fish

Aim  To analyse the patterns in species richness and endemism of the native European riverine fish fauna, in the light of the Messinian salinity crisis and the Last Glacial Maximum (LGM).
Location  European continent.
Methods  After gathering native fish faunistic lists of 406 hydrographical networks, we defined large biogeographical regions with homogenous fish fauna, based on a hierarchical cluster analysis. Then we analysed and compared the patterns in species richness and endemism among these regions, as well as species–area relationships.
Results  Among the 233 native species present in the data set, the Cyprinidae family was strongly dominant (> 50% of the total number of species). Seven biogeographical regions were defined: Western Peri-Mediterranea, Central Peri-Mediterranea, Eastern Peri-Mediterranea, Ponto-Caspian Europe, Northern Europe, Central Europe and Western Europe. The highest regional species richness was observed for Central Peri-Mediterranea and Ponto-Caspian Europe. The highest endemic richness was found in Central Peri-Mediterranea. Species–area relationships were characterized by high slope values for Peri-Mediterranean Europe and low values for Central and Western Europe.
Main conclusions  The results were in agreement with the 'Lago Mare' hypothesis explaining the specificity of Peri-Mediterranean fish fauna, as well as with the history of recolonization of Central and Western Europe from Ponto-Caspian Europe following the LGM. The results also agreed with the mechanisms of speciation and extinction influencing fish diversity in hydrographical networks. We advise the use of the seven biogeographical regions for further studies, and suggest considering Peri-Mediterranean Europe and Ponto-Caspian Europe as 'biodiversity hotspots' for European riverine fish.     

Analysing botanical collecting effort in Amazonia and correcting for it in species range estimation

Aim The aim of this study is to analyse the distribution pattern of the botanical collecting effort in Amazonia so that it can be accounted for when interpreting phytogeographical patterns such as inferred species ranges. We also develop a mechanistic and transparent method for taking into account the bias in collecting effort when estimating likelihoods of species occurrences. Location Amazonia, Neotropics. Methods We utilized electronic data sets of georeferenced herbarium collections (1,063,530 in total). We plotted collecting localities (68,246 in total) on maps overlaid with 1° and 0.5° square grids, and analysed collecting effort using a geographical information system (GIS). We also drew a map of Thiessen polygons, using collecting localities as polygon centres, to visualize collecting density in a scale‐independent way. We then created a ‘collecting activity landscape’ in which well‐collected areas appear as peaks and poorly studied areas as valleys. We demonstrate how this surface can be utilized when estimating species distributions. Results The data available to us confirm that botanical collecting activity is still severely biased in Amazonia. The uncollected area represents 43% of the total area of Amazonia, while another 28% is poorly collected and only 2% can be considered relatively well collected. The Thiessen polygon network represents an improvement in the presentation of collecting intensity compared with square grids. Main conclusions The maps of botanical collecting effort in the Neotropics should be used for visually correcting phytogeographical interpretations. With the help of GIS applications the observed spatial bias in collecting effort can be utilized in estimating the likelihood of occurrence of species in a repeatable manner. These estimates, in turn, can be used for various purposes in basic and applied science as well as in decision‐making. The biased collecting effort should, in the long run, be corrected by further field work in unexplored areas, which can be identified with the maps presented here.     

Areas of endemism in the Espinhaço Range in Minas Gerais, Brazil

The Espinhaço Range, a mountain chain located in the Brazilian states of Minas Gerais and Bahia, contains one of the richest floras in Brazil, with a high frequency of endemic species. Since 2005 it is designated as UNESCO biosphere reserve and is situated at the joint border of two global hotspots for biodiversity conservation. Endemic species with congruent occurrence patterns were identified in order to establish areas of higher endemism within the Espinhaço Range. Taxonomic reviews were analyzed in order to identify endemic taxa and a dataset was elaborated containing 1765 records from 178 endemic species of vascular plants, representing 17 families and including the geographic coordinates for each record. Two maps were produced showing species richness and collection effort in 15′ quadrats. The congruent occurrences were identified and a third map was provided, delimiting 10 candidate areas of endemism for a “parsimony analysis of endemicity” (PAE). One most parsimonious cladogram is then retrieved, evidencing three major clades corresponding to the northern, central and southern portions of the Espinhaço, in addition to four subclades included into the central clade. We finally identified six major areas of endemism. Furthermore, there is a strong correlation between species richness and collector effort, revealing which areas are in need of further field inventories.     

Using Museum Collections to Estimate Diversity Patterns along Geographical Gradients

Quantifying species-richness patterns along geographical gradients (typically latitude and elevation) has a long history in ecology and can be based on more-or-less complete censuses from a specified area (plot sampling), selective collection within a specified area (e.g. museum collections), or general information about species distributions (e.g. observations of extremes along the gradient, distribution maps). All these approaches require complete sampling to give the true richness in an area, but the richness pattern (i.e., the relative changes in richness along the gradient) may be estimated without complete sampling, although equal sampling between areas is necessary. This is relatively easy to do for fine-scale plot sampling, but rarely easy for other types of data. For data extracted from museum collections, a correct perception of the species richness pattern therefore depends on post-sampling treatment of data. Two commonly applied techniques for quantifying species richness patterns with these types of data are discussed, namely interpolation of species ranges and rarefaction. Such treatment may correct for unequal sampling in some instances, but may in other cases introduce artificial patterns. With incomplete sampling interpolation introduces an artificial humped pattern and rarefaction requires similar species abundance distributions to make unbiased comparisons among samples. One must therefore be cautious when applying these methods for estimating species richness patterns along geographical gradients.     

Taxic Richness Patterns and Conservation Evaluation of Madagascan Tiger Beetles (Coleoptera: Cicindelidae)

Distributional ranges of 17 genera and 172 species of Malagasy tiger beetles (Coleoptera, Cicindelidae) have been compiled to determine patterns of species richness and endemism. These patterns reveal large sampling gaps, and potential priority areas for conservation action. Northern and south-western parts of the island are richer in genera, whereas eastern and especially northern parts of the rainforest show higher species richness, due to extensive radiations within the genera Pogonostoma and Physodeutera. A set of 23 areas are identified in this study as priority foci for tiger beetle conservation, and six general regions are bioinventory priorities.     

HISTORICAL DIVERSIFICATION OF BIRDS IN NORTHWESTERN SOUTH AMERICA: A MOLECULAR PERSPECTIVE ON THE ROLE OF VICARIANT EVENTS

Studies of South American biodiversity have identified several areas of endemism that may have enhanced historical diversification of South American organisms. Hypotheses concerning the derivation of birds in the Chocó area of endemism in northwestern South America were evaluated using protein electrophoretic data from 14 taxonomically diverse species groups of birds. Nine of these groups demonstrated that the Chocó area of endemism has a closer historical relationship to Central America than to Amazonia, a result that is consistent with phytogeographic evidence. Within species groups, genetic distances between cis-Andean (east of the Andes) and trans-Andean (west of the Andes) taxa are, on average, roughly twice that between Chocó and Central American taxa. The genetic data are consistent with the hypotheses that the divergence of most cis-Andean and trans-Andean taxa was the result of either the Andean uplift fragmenting a once continuous Amazonian-Pacific population (Andean Uplift Hypothesis), the isolation of the two faunas in forest refugia on opposite sides of the Andes during arid climates (Forest Refugia Hypothesis), or dispersal of Amazonian forms directly across the Andes into the trans-Andean region (Across-Andes Dispersal Hypothesis). Disentangling these hypotheses is difficult due to the complexity of the Andean uplift and to the scant geologic and paleoclimatic information that elucidates diversification events in northwestern South America. Regarding the divergence of cis- and trans-Andean taxa, the genetic, geologic, and paleoclimatic data allow weak rejection of the Andean Uplift Hypothesis and weak support for the Forest Refugia and Andean Dispersal Hypotheses. The subsequent diversification of Chocó and Central American taxa was the result of Pleistocene forest refugia, marine transgressions, or parapatric speciation.     

Subsampling Herbarium Collections to Assess Geographic Diversity Gradients: A Case Study with Endemic Orchidaceae and Rubiaceae in Cameroon

We compiled herbarium specimen data to provide an improved characterization of geographic patterns of diversity using indices of species diversity and floristic similarity based on rarefaction principles. A dataset of 3650 georeferenced plant specimens belonging to Orchidaceae and Rubiaceae endemic to Atlantic Central Africa was assembled to assess species composition per half‐degree or one‐degree grid cells. Local diversity was measured by the expected number of species (S_k) per grid cell found in subsamples of increasing size and compared with raw species richness (S_R). A nearly unbiased estimator of the effective number of species per grid cell was also used, allowing quantification of ratios of ‘true diversity’ between grid cells. Species turnover was measured using a presence/absence‐based similarity index (Sørensen) and an abundance‐based index that corrects for sampling bias (NNESS). Our results confirm that the coastal region of Cameroon is more diverse in endemic species than those more inland. The southern part of this coastal forest is, however, as diverse as the more intensively inventoried northern part, and should also be recognized as an important center of endemism. A strong congruence between Sørensen and NNESS similarity matrices lead to similar delimitations of floristic units. Hence, heterogeneous sampling seems to confer more bias when measuring patterns of local diversity using raw species richness than species turnover using Sørensen index. Overall, we argue that subsampling methods represent a useful way to assess diversity gradients using herbarium specimens while correcting for heterogeneous sampling effort. Abstract in French is available in the online version of this article.     

Species distribution modelling as a macroecological tool: a case study using New World amphibians

Although species distribution modelling (SDM) is widely accepted among the scientific community and is increasingly used in ecology, conservation biology and biogeography, methodological limitations generate potential problems for its application in macroecology. Using amphibian species richness in North and South America, we compare species richness patterns derived from SDM maps and ‘expert’ maps to evaluate if: 1) richness patterns derived from SDM are biased toward climate‐based explanations for diversity when compared to expert maps, since SDM methods are typically based on climatic variables; and 2) SDM is a reliable tool for generating richness maps in hyperrich regions where point occurrence data are limited for many species. We found that although three widely used SDM methods overestimated amphibian species richness in grid cells when compared to expert richness maps in both North and South America due to systematic overestimation of range sizes, diversity gradients were reasonably robust at broad scales. Further, climatic variables statistically explained patterns of richness at similar levels among the different richness sources, although climatic relationships were stronger in the much better known North America than in South America. We conclude that in the face of the high deforestation rates coupled with incomplete data on species distributions, especially in the tropics, SDM represents a useful macroecological tool for investigating broad‐scale richness patterns and the dynamics between species richness and climate.     

Estimated plant richness pattern across northwest South America provides similar support for the species-energy and spatial heterogeneity hypotheses

Explaining geographic variation in plant species richness at broad spatial scales has long been a major challenge. Many hypotheses have been proposed during the last 200 yr, but recent work has focused on a few major alternatives. Among these, two hypotheses contend that plant species richness reflects 1) variation in energy and water availability among sampling units (the species-energy hypothesis) and 2) habitat and topographic heterogeneity within sampling units (the spatial heterogeneity hypothesis). We used a large botanical database and regression models to simultaneously confront the predictions from both hypotheses against an estimate of vascular plant richness across northwest South America. This estimate provided similar support for both hypotheses, a result that may be seen as contrasting with the notion that variation in energy and water availability among sampling units is the main determinant of plant species richness. We discuss potential explanations for this apparent discrepancy. Regression models that incorporated the relative contributions of both hypotheses predicted that the highest plant species richness in northwest South America is found in topographically complex areas. In contrast to several of the most recent mapping efforts, lowland Amazonia was predicted to be a plant richness trough in the study region. We suggest that diverging portrayals of plant richness across northwest South America result from differences in estimates of the relative importance of the species-energy and the spatial heterogeneity hypotheses.