Areas of endemism of Mexican mammals: reanalysis applying the optimality criterion

In order to test Mexican areas of endemism of mammals identified by previous parsimony analyses of endemicity (PAEs), we applied the optimality criterion to three data matrices (based on point records, potential distributional models and the fill option in software NDM). We modelled the ecological niches of 429 terrestrial mammal species using the genetic algorithm for rule-set prediction (GARP) and models were projected as potential distributional areas. We overlapped the point occurrence data and the individual maps of potential distributions to a grid of 1° latitude–longitude. Three matrices of 247 grid cells (areas) and 429 species were built: (1) a binary matrix with '0' for absence and '1' for presence of at least one record of the species inside the grid-cell; (2) a three-state matrix similar to (1) but assigning the state '2' to the assumed presence in the model of potential distribution; and (3) a three-state matrix similar to (2), but applying the fill option of software NDM instead of using a model. The optimality criterion was performed in NDM version 2.7 and results were examined with VNDM version 2.7. The first and second matrices showed 13 areas of endemism and the third identified 16 areas of endemism. NDM provided a better resolution than PAE, allowing us to identify several new areas of endemism, previously undetected. Ecological niche models, projected as potential distributional areas, and the optimality criterion are very useful to identify areas of endemism, although they should be used with caution because they may overpredict potential distributional areas. PAE seems to underestimate the areas of endemism identified.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 468–478.     

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.     

Biogeographic analysis of endemic cacti of the Sierra Madre Oriental, Mexico

The distribution of cacti species that inhabit the Sierra Madre Oriental (SMO) was analysed. Grid-cells were analysed using parsimony analysis of endemicity (PAE) and endemism indices. Areas characterized by their diagnostic species were determined, aiming to propose areas for the conservation of threatened cacti. Distributional data were obtained from 1936 herbarium specimens, electronic information, and from field collections. Eight areas of endemism and three main clades were obtained from the grid-cell analysis. Areas obtained from the endemism indices are very similar to those obtained with the PAE, but differ in the association of grid-cells. PAE showed endemism patterns indicating that southern and central sections of the SMO province are the areas richest in geographically-restricted species. The results obtained with different endemism indices detected more or less the same areas, although the importance level is different. The corrected weighted endemism index can be considered as a reliable measure of endemism because it is unrelated to species richness. A regionalization of the SMO in three subprovinces is suggested, supported by characteristic cacti taxa and the existence of natural barriers.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 373–389.     

Track analysis of the North and Central American species of the Pantomorus–Naupactus complex (Coleoptera: Curculionidae)

We undertook a panbiogeographic analysis of the broad‐nosed weevils of the genera Naupactus Dejean, 1821, Pantomorus Schönherr, 1840 and Phacepholis Horn, 1876 (Coleoptera: Curculionidae) from North and Central America to propose a biogeographic scenario to explain their biotic diversification. Based on individual tracks of 30 species, we obtained six generalized tracks: Mesoamerican, Chiapas, Sierra Madre del Sur, Mexican Pacific Coast, Southern Great Plains and Northern Great Plains tracks. The Sierra Madre del Sur generalized track is the best supported, based on 10 species of the three genera. We found two nodes, one at the intersection of the Mesoamerican and Chiapas tracks, and another at the intersection of the Chiapas and Sierra Madre del Sur tracks. Species of Naupactus are primarily distributed in lowlands, associated mostly with dry forests and xeric environments. Species of Pantomorus and Phacepholis would have diversified from South American Naupactus‐like ancestors, mainly in montane habitats and lowlands of North and Central America, between sea level to about 2500 m of altitude.     

Patterns of Amazonian area relationships based on raw distributions of papilionid butterflies (Lepidoptera: Papilioninae)

Diversifications within a biota are due to several factors. Although some of these are untestable with current analytical methods, hierarchical congruence obtained with different cladistic methods and based on independent taxa are undoubtedly important. In the recent past, most hypotheses of historical biogeography (e.g. refugial, riverine, disturbance, vicariance) were tested on the Amazonian biota, selecting a number of diverse organisms such as plants, anurans, lizards, butterflies, birds and monkeys. In this study we used parsimony analysis of endemicity to infer historical relationships among 16 interfluvial areas in the Amazonian lowlands based on raw distributions of 114 Papilioninae (Lepidoptera). The analysis yielded two most parsimonious trees of area relationships. One tree was characterized by two main clusters of areas which showed a separation of Guyanan + south-east Amazonian interfluvial areas from western Amazonian interfluvial areas. The second tree showed the Guyanan interfluvial areas basal to a cluster which included all the other interfluvial areas. This latter cluster was subdivided into two main groups of areas separating the south-east Amazonian and the western Amazonian interfluvial areas. This result is discussed in the light of previous hypotheses obtained with the same method using some vertebrate taxa in the Amazonian lowlands. Likewise, comparisons with other hypotheses on lineages of birds, mammals and butterflies obtained applying cladistic biogeographical methods are made. The two alternative vicariant patterns presented for papilionid butterflies are strictly congruent with those for birds.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 345–357.     

Parsimony analysis of endemicity (PAE) of Mexican terrestrial mammals at different area units: when size matters

Abstract Aim Parsimony analysis of endemicity (PAE) is a biogeographical method that uses a parsimony algorithm to obtain an area cladogram, based on taxa inhabiting the study areas. We compare its performance at different geographical units (½° and 1° quadrats, ecoregions and biogeographical provinces) to analyse distributional patterns of Mexican terrestrial mammals, in order to assess the importance of the size of area units. Location The area analysed corresponds to Mexico. Methods Parsimony analyses were based on 56,859 collection records, corresponding to 703 genera, species and subspecies. Four data matrices were constructed for: (1) 716 quadrats of ½° latitude × ½° longitude, (2) 230 quadrats of 1° latitude × 1° longitude, (3) forty‐five ecoregions and (4) fourteen biogeographical provinces. Results For the ½° quadrat matrix, we obtained six cladograms of 17,138 steps. For the 1° quadrat matrix, we obtained five cladograms (strict consensus with 9394 steps). For the matrix of ecoregions, we obtained twelve cladograms (strict consensus cladogram with 3009 steps). For the provinces, we obtained a single cladogram with 1603 steps. Main conclusions The best results were obtained with natural areas instead of quadrats. There seems to exist a trend to decrease the absolute number of steps and an increase in the absolute and relative number of synapomorphies as the size of the area units decreases, although this does not necessarily occur for the number of cladograms.     

Cladistic analysis of distributions and endemism (CADE): using raw distributions of birds to unravel the biogeography of the South American aridlands

Aim To use published distributional and taxonomic information on Neotropical birds in a cladistic analysis of distributions and endemism (CADE) to generate a testable hypothesis of area‐relationships for the arid areas of endemism, particularly those of Central South America (the ‘arid diagonal’), and to clarify the different methodologies commonly associated with parsimony analysis of endemicity (PAE). Location South America. Methods Cladistic analysis of distributions and endemism. Results We obtain a tree where the relationships of most areas are resolved, and we find support for an exclusive clade of Central South American areas, with the Caatinga as sister to both the Chaco and Cerrado. Main conclusions There is a substantial amount of historical signal in avian distributions, when large numbers of taxa and multiple taxonomic levels are considered. Ecological noise and historical information are more easily distinguished in CADE analyses than they would be in PAE analyses. Based on our results we predict that among aridland birds, the Cerrado and Chaco species will be more closely related to each other than to Caatinga species.     

Patterns of species richness for blackflies (Diptera: Simuliidae) in the Nearctic and Neotropical regions

Abstract.  1. Patterns of simuliid species richness were examined over a variety of scales at 532 stream sites in the Nearctic (394) and Neotropical (138) regions. In Nearctic streams, species richness of immature blackflies both within and across ecoregions and over two seasons was examined. Stream variables at each site included seston, width, depth, velocity, discharge, conductivity, pH, dissolved oxygen, water temperature, dominant streambed-particle size, canopy cover, and riparian vegetation. These variables were subjected to a principal component analysis and derived principal components were related back to richness, using regression analysis. At the level of the stream reach, richness was not highly correlated with single-point measurements of stream conditions.
2. Using data from both Nearctic and Neotropical sites, the effect of regional richness on local richness was examined. As regional richness increased, local diversity reached an asymptote in which further increases in regional richness were not matched by increases in local richness. Hence, simuliid communities are best described as saturated (type II) communities, consistent with the current view of lotic communities as non-equilibrium systems.
3. The well-established pattern of greater species richness in tropical regions was not observed in this study. To the contrary, blackfly richness is higher in temperate streams than in tropical streams at both local and regional scales.     

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).     

Track analysis of the Nearctic region: Identifying complex areas with mammals

The Nearctic region is located on the North American plate. However, its tectonic history is related to convergence with other plates, which has promoted a complex topography. This complexity should be reflected by the distributional patterns of the biota. We used track analysis with 574 species of mammals to identify generalized tracks and panbiogeographic nodes in the Nearctic region and to propose an updated point of view of complex areas and their boundaries in North America. Seven generalized tracks with nested patterns (California, Columbia Plateau, Mesoamerican, Mexican Plateau, Neotropic, Southern Rocky Mountains, and Western Coast of USA) were identified using a parsimony analysis of endemicity with progressive character elimination. Nine panbiogeographic nodes were identified at the intersections of the generalized tracks, all of which were located in the Sierra of Chiapas and Central America physical features. A total of 192 nodes were identified for the nested patterns, located in only eight physical features. Our analysis revealed evolutionary patterns in generalized tracks, and the panbiogeographic nodes predicted areas with high evolutionary–geologic complexity, shared by other taxonomic groups.     

Biogeographical analysis of the flea beetle genus Chaetocnema in the Afrotropical Region: distribution patterns and areas of endemism

Aim The areal distributions of Chaetocnema species in the Afrotropical Region have been analysed with the aims of determining the distribution patterns (chorotypes) and identifying the most important areas of endemism for this flea beetle genus in sub‐Saharan Africa. Location Data were collected in sub‐Saharan Africa, including Madagascar. Methods The Afrotropical Region was divided into 103 5° quadrats (operative geographical units, or OGUs). A presence–absence matrix of the Afrotropical Chaetocnema species in the OGUs was analysed by cluster analysis (Baroni Urbani & Buser index and the WPGMA clustering method) to generate distribution pattern data based on similarity of distribution. The most important areas of endemism were identified by parsimony analysis of endemicity. Results The general distribution of Chaetocnema in the Afrotropical Region was found to be associated with moist environments and montane grasslands. Most species exhibit restricted geographical ranges. Cluster analysis revealed 120 spatial distributions that can be grouped into 13 distinct distribution patterns (chorotypes). The most important areas of endemism for Chaetocnema in sub‐Saharan Africa according to the present parsimony analysis of endemicity are: (1) central and eastern Madagascar [endemicity rate (ER) = 61.1%], (2) Western Cape Province (ER = 36.4%), (3) southern Drakensberg (ER = 26.7%), (4) the Shaba Region (ER = 16.7%), and (5) the North‐Kivu Region (ER = 5.0%). Main conclusions There are 123 known species of Chaetocnema in the Afrotropical Region, more than in any other zoogeographical region. About 91% of the species are endemic and they generally exhibit a restricted and often very localized geographical range. The remaining 9% of the species are represented by seven species that also inhabit northern Africa and/or the Arabian peninsula (C. bilunulata Demaison, C. ganganensis Bechyné, C. ljuba Bechyné, C. pulla Chapuis, C. tarsalis Wollaston, and C. wollastoni Baly), three species that widely inhabit the Palaearctic Region (C. conducta (Motschulsky), C. schlaeflini (Stierlin), and C. tibialis (Illiger)), and two species that were introduced (C. confinis Crotch, and C. picipes Stephens).     

中国斑腿蝗科特有种的分布及特有分布区划分

按照150km×150km对中国陆地部分进行栅格划分,收集整理了中国斑腿蝗科237个特有种在各栅格的分布信息,建立了物种(237)×栅格(168)数据矩阵,研究了中国斑腿蝗科特有种的分布规律并用PAE法进行特有分布区划分。结果表明:除天山山地分布的少数几个种外,中国斑腿蝗科特有种基本分布西南-东北走向。在该范围内南多北少。华南和西南是中国斑腿蝗科特有种最丰富的地区,在秦巴山区、藏东南和浙闽丘陵地带各有一个高密度的分布区。11个特有分布区中5个分布在东洋界部分,两个在中国东部古北和东洋界交界区,天山山地有一个分散的特有分布区,东北有2个特有分布区。特有分布区的分布和划分基本和中国动物地理区划及区系相一致,但台湾地区归入华中区;横断山和秦岭山地的栅格在支序图中多形成未决分类单元,特有分布区划分困难;华南和华中地区的特有分布区可以进一步划分成更小范围的特有分布区;从云南西北的腾冲、保山经过贵州南部到广西桂林的一个东西狭长的区域可以作为一个特有分布区     

Historical biogeographical patterns of the species of Bursera (Burseraceae) and their taxonomic implications

Aim The plant genus Bursera, with 104 species of trees and shrubs, has been used as a model for biogeographical analyses because of its high species richness and large number of endemic species. The biogeographical patterns of Bursera and their implications for its phylogenetic classification are reviewed in order that some hypotheses on the historical biogeography of tropical Mexico can be proposed. Location Bursera is found in the south‐western USA, most of Mexico, mainly below 1700 m elevation in tropical forests, with some species in xeric shrublands, diversifying along the Pacific slope, Central America, and north‐western South America. A few species occur on the Galapagos and Revillagigedo archipelagos, some of which are endemics, whereas in the Antilles species are distributed extensively, with several endemics in the Bahamas, Cuba, Jamaica, and Hispaniola. Methods Data from specimens in herbaria and the literature were used to construct a matrix of 104 species in 160 areas. Distributional patterns of the species of Bursera were inferred applying track analysis, parsimony analysis of endemicity (PAE), and Brooks parsimony analysis (BPA). Results Track analysis revealed four individual tracks: (1) a circum‐Caribbean track, comprising species of the Bursera simaruba species group; (2) an Antillean track, including species that have been transferred to Commiphora based on their pollen traits; (3) a Mexican Pacific track, including species of the B. fragilis, B. microphylla, and B. fagaroides species groups, called ‘cuajiotes’; and (4) a Neotropical Pacific track, including the two species groups assigned to section Bullockia, in which the individual track of the Bursera copallifera species group is nested within the track of the B. glabrifolia species group. The four tracks overlap in a node in the Mexican Pacific slope, where they are highly diversified. PAE allowed us to identify 22 areas of endemism: 12 in Mexico (11 along the Mexican Pacific slope), six in the Antilles, two in Central America, one in South America, and one in the Galapagos. The general area cladogram obtained by BPA has two main clades: one includes the greater Antilles; and the other, 12 Mexican areas of endemism. Main conclusions Bursera fragilis, B. microphylla, and B. fagaroides species groups can be treated together as a new section within Bursera, sect. Quaxiotea, because they are segregated from the other groups of sect. Bursera based on morphological, anatomical, molecular and geographical evidence.     

Distributional patterns of the Neotropical humid montane forest avifaunas

The study of biogeographical patterns of the Neotropical humid montane forest avifauna has been prevented due to the lack of phylogenetic hypotheses for most taxa, and to the paucity of detailed studies about the geographical distribution for most of the species. Distributional patterns of this avifauna were explored by integrating predictive distributional models and Parsimony Analysis of Endemicity (PAE). Distributional maps were generated using the Genetic Algorithms for Rule Set Prediction for 442 species; this information was transformed into a data matrix for analysis with PAE. Hierarchical information corresponding to the genus level was incorporated to help resolve the relationships between areas. A strict consensus cladogram showed a clear separation between the Mesoamerican and South American avifaunas. Within the Mesoamerican clade, three geographically structured groups were obtained, whereas the South American clade showed a polytomy of three groups, with two of them, the South Andean Yungas and the Tepuis, located outside the main clade. The third group is a well resolved clade, including areas from eastern Panama and northern Venezuela to central Bolivia. Area relationships suggest a mixed history of dispersal and vicariant events, with the latter being the most important for explaining the biogeographical patterns found.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 175–194.     

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.     

Historical biogeography of South American freshwater fishes

Aim To investigate biogeographical patterns of the obligate freshwater fish order Characiformes. Location South America. Methods Parsimony analysis of endemicity, likelihood analysis of congruent geographical distribution, and partition Bremer support were used. Results Areas of endemism are deduced from parsimony analysis of endemicity, and putative dispersal routes from a separate analysis of discordant patterns of distribution. Main conclusions Our results demonstrate the occurrence of 11 major areas of endemism and support a preferential eastern–western differentiation of the characiforms in the Amazonian region, contrasting with the southern–northern differentiation of terrestrial organisms. The areas of endemism identified seem to be deeply influenced by the distribution of the emerged land during the 100‐m marine highstand that occurred during the late Miocene and allow us to hypothesize the existence of eight aquatic freshwater refuges at that time. The raw distribution of non‐endemic species supports nine patterns of species distribution across the 11 areas of endemism, two of which support a southern–northern differentiation in the eastern part of the Amazon. This result shows that the main channel of the Amazon limited dispersal between tributaries from each bank of the river. The levels of endemism further demonstrate that the aquatic freshwater refuges promoted allopatric speciation and later allowed the colonization of the lowlands. By contrast, the biogeographical pattern found in the western part of the Amazon is identified as a result of the Miocene Andean foreland dynamic and the uplift of the palaeoarches that promoted allopatric divergence across several sedimentary basins by the establishment of disconnected floodplains. The assessment of conflicting species distributions also shows the presence of seven putative dispersal routes between the Amazon, Orinoco and Paraná rivers. Our findings suggest that, rather than there being a single predominant process, the establishment of the modern South American freshwater fish biotas is the result of an interaction between marine incursions, uplift of the palaeoarches, and historical connections allowing cross‐drainage dispersal.     

Modelling the spatial distribution of selected North American woodland mammals under future climate scenarios

1. North America has a diverse array of mammalian species. Model projections indicate significant variations in future climate conditions of North America, and the habitats of woodland mammals of this continent may be particularly sensitive to changes in climate.
2. We report on the potential spatial distributions of 13 wide-ranging, relatively common species of North American woodland mammals under future climate scenarios.
3. We examined the potential influence of the mean and seasonal climate variables on the distribution of species. Presence-only occurrence records of species, four predictor variables, two future climate scenarios (Representative Concentration Pathways 4.5 and 8.5), and two time steps (current and 2070) were used to build species’ distribution models using a maximum entropy algorithm (MaxEnt).
4. Our results suggested that overall, 11 of the 13 species are likely to gain climatically suitable space (regions where climate conditions will be similar to those of area currently occupied) at the continental scale, but American marten Martes americana and ‘woodland’ caribou Rangifer tarandus are likely to lose suitable climate range by 2070. Furthermore, climate space is likely to be expanding northwards under future climate scenarios for most of the mammals, and many jurisdictions in the border region between Canada and the USA are likely to lose iconic species, such as moose Alces alces. We identified regions as potential in situ and ex situ climate change refugia, which are increasingly considered to be important for biodiversity conservation.
5. The model results suggest significant implications for conservation planning for the 13 mammalian species under global climate change, especially at fine spatial scales. Numerous species that are presently common at their southern range edge will be functionally or completely extirpated in 50 years. The potential in situ and ex situ climate change refugia could provide an effective support for adaptive strategies aimed at species conservation planning.