Extending the Namibian protected area network to safeguard hotspots of endemism and diversity

Namibia's state protected area network (PAN) covers 13.8% of the country's land area, but is seriously inadequate as a basis for effective biodiversity conservation. The early parks system was not designed with biological diversity in mind, and reflects instead a history of ideological, economic and veterinary considerations. Currently, parks in the Namib Desert biome make up 69% of the PAN, while savanna and woodland biomes are somewhat underrepresented (7.5 and 8.4% of their respective land areas), and the Karoo biome is badly underrepresented (1.6%). Four of 14 desert vegetation types are comprehensively protected, with 67 to 94% representation in the PAN, yet six savanna types have 0 to 2% representation by area. Mountain Savanna, a vegetation type unique to Namibia, is wholly unprotected. The status of two marine reserves, which in theory protect only 0.01% of Namibia's marine environment, needs clarification and augmentation with new reserves. Nearly 85% of Namibia's land is zoned for agriculture, so effective biodiversity protection means working outside the PAN to improve the sustainability and diversity of farming practices. Wildlife conservancies on commercial and communal farmlands show excellent potential to mitigate the ecological skew in the state PAN, with the ecological management of large areas being decentralized to rural communities in habitats otherwise neglected for conservation. Two important endemism zones, the Kaoko escarpment and coastal plain and the Sperrgebiet succulent steppe, plus the species-rich Caprivi area, offer three valuable opportunities for regional consolidation of protected areas into transboundary 'peace parks' or biosphere reserves.     

Origin and diversification of the Greater Cape flora: ancient species repository, hot-bed of recent radiation, or both?

Like island-endemic taxa, whose origins are expected to postdate the appearance of the islands on which they occur, biome-endemic taxa should be younger than the biomes to which they are endemic. Accordingly, the ages of biome-endemic lineages may offer insights into biome history. In this study, we used the ages of multiple lineages to explore the origin and diversification of two southern African biomes whose remarkable floristic richness and endemism has identified them as global biodiversity hotspots (succulent karoo and fynbos). We used parsimony optimization to identify succulent karoo- and fynbos-endemic lineages across 17 groups of plants, for which dated phylogenies had been inferred using a relaxed Bayesian (BEAST) approach. All succulent karoo-endemic lineages were less than 17.5 My old, the majority being younger than 10 My. This is largely consistent with suggestions that this biome is the product of recent radiation, probably triggered by climatic deterioration since the late Miocene. In contrast, fynbos-endemic lineages showed a broader age distribution, with some lineages originating in the Oligocene, but most being more recent. Also, in groups having both succulent karoo- and fynbos-endemic lineages, there was a tendency for the latter to be older. These patterns reflect the greater antiquity of fynbos, but also indicate considerable recent speciation, probably through a combination of climatically-induced refugium fragmentation and adaptive radiation.     

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.     

Biodiversity in the Gulf of Guinea: an overview

Patterns of species-richness and endemism in the Gulf of Guinea reflect the region's biogeographic history. Bioko is a continental-shelf island that was recently connected to the African mainland, whereas Príncipe, São Tomé and Annobón are truly oceanic and have never been connected with each other or with the mainland. As a result, Bioko supports a much more diverse flora and fauna but with relatively low levels of endemism at the species level, whereas the oceanic islands are relatively depauperate because of their isolation but rich in endemic taxa. Species endemism is 0–3% on Bioko for angiosperms, bats, birds, reptiles and amphibians, compared with much higher values on Principe for these same taxa of 8% (plants) to 100% (amphibians), on São Tomé between 14% (plants) and 100% (amphibians), and on Annobón 0% (bats) to 71% (reptiles). On a global scale, for their size both Príncipe and São Tomé support unusually high numbers of single-island endemic species of birds, reptiles and amphibia. For its tiny size, Annobón is also notable for its endemic birds and reptiles. Among terrestrial molluscs the rates of endemism are in general higher than for plants and vertebrates, from ca 50% on Bioko to ca 80% on the oceanic islands. In contrast and as might be expected, only Bioko supports a rich freshwater fish fauna and it contains many endemic taxa, whereas the oceanic islands support only a few salt-tolerant species. The Gulf of Guinea islands are also important for their marine organisms, amongst which coral reef fish and marginellid molluscs show high levels of endemism, though they are not especially species-rich. The Gulf of Guinea islands are of great interest to conservationists and evolutionary biologists. Each island, of greatly differing size and degree of isolation, has acquired its unique sub-set of plants and animals separately from the neighbouring mainland, followed by adaptive radiations in situ. For this reason the conservation value of the archipelago as a whole is greater than the sum of the biodiversity contained in its individual islands. Conservation initiatives in the Gulf of Guinea should therefore ensure that representative terrestrial, freshwater and marine habitats and groups of organisms are targeted in a co-ordinated manner among the islands.     

Avian diversity and endemism in Namibia: patterns from the Southern African Bird Atlas Project

Namibia supports a highly diverse avifauna of 644 species, including over 90 species endemic to the southern African subregion and 13 species endemic to the country. Patterns of species diversity in relation to protected areas and habitat types were analysed using data from the Southern African Bird Atlas Project. A modified Shannon index appropriate for atlas data was used to derive an index of diversity for all species, wetland, terrestrial, threatened, and regional endemic species. Species richness for Namibian endemics was mapped. Overall species diversity is highest in the northeast of Namibia where wetland and riparian habitats coincide. Both wetland and terrestrial species show highest diversity in this area. The greatest diversity of southern African endemics falls within the Savanna-Karoo systems. Several key areas are identified for red data species, including the Caprivi Strip, Kunene and Orange Rivers, coastal wetlands and ephemeral river mouths and pans. This highlights the pressures operating on wetland and riparian habitats in arid environments. Concentrations of Namibian endemics are found in the northwestern (Kaoko) escarpment of the country. Although much of the area of high diversity of wetland, terrestrial and red data species falls within protected areas, national and regional endemics are poorly represented within national parks.     

Species richness and conservationof Namibian freshwater macro-invertebrates,fish and amphibians

Namibia is an arid country but has a diverse array of wetland habitats ranging from ephemeral water bodies and rain-pools, artesian springs supporting small perennial pools and streams, to the large perennial rivers of the north-east with their associated floodplains. These rivers drain wetter areas north of Namibia and contribute many tropical species to Namibia's wetlands. There are 778 described species of macro-invertebrates representing eight phyla with 81 endemics. Many invertebrates still have to be identified or described but presently the greatest endemism occurs among the Ostracoda (18 species), Coleoptera (17), Diptera (14), Anostraca (six) and Amphipoda (five species). In total, Namibia has 50 species of frog with three endemics. No caecilians or salamanders occur in Namibia. There are 114 species of freshwater fish with five endemics. Most Namibian wetlands occur outside protected areas. Over-exploitation of wetland resources and flow regulation are currently major threats, but new environmental legislation being formulated is based on the goal of sustainable use.     

The species diversity, distributionand conservation of Namibian mammals

Namibia's extant mammal fauna of 250 species represents about 75% of the southern African region's species richness, 83% of generic richness and 98% of familial richness. Fourteen species are presently recognized as endemic (75% or more of the global population occurring within Namibian borders). These endemics occur in the Namib Desert, pro-Namib transition zone and adjoining escarpment, and are primarily rupicolous. The Namibian endemic mammal fauna is characterized by the monotypic Petromuridae, and the rodent genera Gerbillurus and Petromyscus. The distribution of smaller species has probably not changed significantly over the past 200 years, but species such as lion and plain zebra have undergone range reductions of 95% or more, and five species are listed as recently extinct. Approximately 50% of all Namibian mammal species are provisionally listed as 'secure'. However, due to patchy data, 94 species (38%) are classified as under possible or probable threat. Nineteen species (8%) are classified as under definite threat. Over 13% of Namibia is set aside by the state for conservation purposes. Ninety-five percent of mammal species occur in at least one park, over 80% occur in three or more parks, and 59 species (28%) occur in ten or more parks, although for most species nothing is known of their population viability there. Major threats to mammals in Namibia are invasive aliens, including the risk of genetic pollution, and habitat alteration, especially wetland degradation.     

Endemic vertebrates are the most effective surrogates for identifying conservation priorities among Brazilian ecoregions

Many studies have tested the performance of terrestrial vertebrates as surrogates for overall species diversity, because these are commonly used in priority‐setting conservation appraisals. Using a database of 3663 vertebrate species in 38 Brazilian ecoregions, we evaluated the effectiveness of various subsets for representing diversity of the entire vertebrate assemblage. Because ecoregions are established incorporating information on biotic assemblages, they are potentially more amenable to regional comparison than are national or state lists. We used 10 potential indicator groups (all species; all mammals, birds, reptiles, or amphibians; all endemic species; and endemic species within each class) to find priority sets of ecoregions that best represent the entire terrestrial vertebrate fauna. This is the first time such tests are employed to assess the effectiveness of indicator groups at the ecoregion level in Brazil. We show that patterns of species richness are highly correlated among mammals, birds, amphibians, and reptiles. Furthermore, we demonstrate that ecoregion sets selected according to endemic species richness captured more vertebrate species per unit area than sets based on overall vertebrate richness itself, or than those selected at random. Ecoregion sets based on endemic bird, endemic reptile, or endemic amphibian richness also performed well, capturing more species overall than random sets, or than those selected based on species richness of one or all vertebrate classes within ecoregions. Our results highlight the importance of evaluating biodiversity concordance and the use of indicator groups as well as aggregate species richness. We conclude that priority sets based on indicator groups provide a basis for a first assessment of priorities for conservation at an infracontinental scale. Areas with high endemism have long been highlighted for conservation of species. Our findings provide evidence that endemism is not only a worthwhile conservation goal, but also an effective surrogate for the conservation of all terrestrial vertebrates in Brazil.     

Comparative effects of urban and agricultural land transformation on Odonata assemblages in a biodiversity hotspot

Rivers of the Cape Floristic Region (CFR) biodiversity hotspot are threatened by land transformation. This region is a centre of endemism for many taxa, including Odonata. These insects are highly sensitive to changes in physical habitat structure, which makes them good bioindicators, and this led to the development of the Dragonfly Biotic Index (DBI). We investigated the effects of local agricultural and urban land transformations on Odonata species richness, assemblage composition and DBI scores in three CFR rivers. A total of 48 sites were selected and categorized as natural, agricultural or urban land use. Adult male Odonata and four environmental variables were recorded over two seasons. Land transformation significantly influenced Odonata assemblage composition but did not always significantly reduce species richness. Average vegetation height also affected Odonata assemblage composition and decreased species richness. Agricultural and urban sites had Odonata assemblages differing from those in the natural areas. Agricultural and urban local land use types reduced opportunities for some endemic species but provided for the persistence and establishment of widespread, generalist species, as indicated by great changes in DBI scores. Mitigating the adverse influences of land transformation through establishment of protected areas is essential for the conservation of rare taxa, particularly in an area with a high number of endemic species.     

Distribution patterns of biodiversity and the design of a representative reserve network in Portugal

Representativeness is a desirable property of conservation networks. In this paper an attempt is made to assess the efficiency of current conservation networks in Portugal in representing vertebrates (reptiles and amphibians) and plants (gymnosperms, pteridophytes and bryophytes). It was found that whilst the protected areas do not sample all species in the database they provide a better result than choosing areas at random. For the goal of maximizing representation of taxa per unit area hotspots and complementarity performed better. A pattern of over-representation of vertebrates in relation to lower plants was discovered among selected conservation areas in Portugal suggesting that charismatic organisms with large home ranges may not always be the most appropriate surrogates for biodiversity when representativeness is sought. When trying to fill the gaps in current protected areas with additional areas only complementarity performed better than choosing areas at random. Both rarity and richness hotspots gave worse results. Opportunistic administrative criteria such as supplementing the PAS with other conservation areas (CORINE Biotopes) without taking into account their contribution to a representation goal was nearly half as good as choosing areas at random. The results recall for the need of explicit goals and accountable methods in area selection for conservation and reinforce the role of complementarity for finding additional areas to protected areas when limited resources are available for ‘in situ’ conservation.     

Extinction Risks and the Conservation of Madagascar's Reptiles

Background

An understanding of the conservation status of Madagascar''s endemic reptile species is needed to underpin conservation planning and priority setting in this global biodiversity hotspot, and to complement existing information on the island''s mammals, birds and amphibians. We report here on the first systematic assessment of the extinction risk of endemic and native non-marine Malagasy snakes, lizards, turtles and tortoises.

Methodology/Principal Findings

Species range maps from The IUCN Red List of Threatened Species were analysed to determine patterns in the distribution of threatened reptile species. These data, in addition to information on threats, were used to identify priority areas and actions for conservation. Thirty-nine percent of the data-sufficient Malagasy reptiles in our analyses are threatened with extinction. Areas in the north, west and south-east were identified as having more threatened species than expected and are therefore conservation priorities. Habitat degradation caused by wood harvesting and non-timber crops was the most pervasive threat. The direct removal of reptiles for international trade and human consumption threatened relatively few species, but were the primary threats for tortoises. Nine threatened reptile species are endemic to recently created protected areas.

Conclusions/Significance

With a few alarming exceptions, the threatened endemic reptiles of Madagascar occur within the national network of protected areas, including some taxa that are only found in new protected areas. Threats to these species, however, operate inside and outside protected area boundaries. This analysis has identified priority sites for reptile conservation and completes the conservation assessment of terrestrial vertebrates in Madagascar which will facilitate conservation planning, monitoring and wise-decision making. In sharp contrast with the amphibians, there is significant reptile diversity and regional endemism in the southern and western regions of Madagascar and this study highlights the importance of these arid regions to conserving the island''s biodiversity.     

An assessment of the efficiency of protection status through determinations of biodiversity hotspots based on endemic bird species,Taiwan

Evaluations of species richness patterns have been performed at diverse scales, and biodiversity hotspots, especially endemism hotspots, have received much attention in conservation biology. We estimated the distributions of endemic bird species based on a 12-yr avian inventory project in Taiwan, identified biodiversity hotspots of endemism on a regional scale based on predictions from the ensemble forecasting framework and frequency histogram approach, and assessed the efficiency of protected areas. The results indicated that the predicted endemism hotspots were mostly located in mid- and high-elevation areas along the Central Mountain Range of Taiwan. An observed endemism hotspot was defined as one in which at least five of Taiwan's 17 endemic bird species were present. This criterion was used because the 5% of the sampled grid squares that were the richest in endemic bird species all had 5 endemic bird species or more. Seventy to seventy-one percent of the observed biodiversity hotspots matched the predicted biodiversity hotspots. This outcome was obtained whether the richness biodiversity in a grid square was based on summed predicted probability or summed predicted richness. The majority of the protected areas for these Taiwanese endemic bird species were national parks, protecting 24.1% of the predicted hotspot areas, whereas nature reserves and wildlife refuges protected less than 7%. Most of the predicted endemism hotspots were not adequately protected. We conclude that the ensemble forecasting framework and the frequency histogram approach are useful for selecting critical habitats and biodiversity hotspots for endemic species and for appraising the efficiency of the protection status provided by governments.     

Centres of plant endemism in China: places for survival or for speciation?

Aim This study aimed to identify the ‘centres of endemism’ of the Chinese spermatophyte flora in order to indirectly detect the locations of past glacial refugia. The role of these areas as places for plant survival (‘plant museums’) and/or areas for plant evolution and speciation (‘plant cradles’) was also assessed. Location China. Methods Distribution patterns of 555 plant endemic taxa, taken as a representative sample of the Chinese endemic flora, were mapped on a 1° × 1° latitude/longitude grid. For each grid cell, species richness (total count of species) and weighted richness (down‐weighting each species by the inverse of its range) were calculated. Grid cells within the top 5% of highest values of weighted richness were considered centres of endemism. Based on available information, all plant taxa included in this study were classified into palaeoendemics and neoendemics, and their distributional patterns were represented separately. Results Twenty areas of endemism were identified in central and southern China, roughly corresponding to mountain ranges, including the Hengduan and Daxue Mountains, the Yungui Plateau, central China Mountains, the Nanling Mountains, eastern China Mountains, and Hainan and Taiwan. Although almost all centres of endemism contained both palaeoendemic and neoendemic taxa, considerable differences in their respective numbers were recorded, with the majority of neoendemics on the eastern fringe of the Tibetan Plateau (Hengduan Mountains sensu lato) but more palaeoendemics towards the east. Main conclusions Owing to their varied topography, the mountainous regions of central and southern China have provided long‐term stable habitats, which allowed palaeoendemics to persist and facilitated the process of speciation. Contrasting patterns between the palaeoendemics and neoendemics within refugia might be attributable to the geological and tectonic history of specific areas. The eastern fringe of the Tibetan Plateau clearly constitutes the ‘evolutionary front’ of China, probably as a result of the uninterrupted uplift of the plateau since the late Neogene. In contrast, the tectonic stability of central and southern China during the Tertiary may have facilitated the persistence of relict plant lineages.     

The living heart: Climate gradients predict desert mountain endemism

Mountain regions are centers of biodiversity endemism at a global scale but the role of arid‐zone mountain ranges in shaping biodiversity patterns is poorly understood. Focusing on three guilds of taxa from a desert upland refugium in Australia, we sought to determine: (a) the relative extent to which climate, terrain or geological substrate predict endemism, and (b) whether patterns of endemism are complimentary across broad taxonomic guilds. We mapped regional endemism for plants, land snails, and vertebrates using combined Species Distribution Models (SDMs) for all endemic taxa (n = 82). We then modelled predictors of endemism using Generalised Additive Models (GAMs) and geology, terrain, and climate variables. We tested for the presence of inter‐ and intraguild hotspots of endemism. Many individual plant and land snail taxa were tightly linked with geology, corresponding to small distributions. Conversely, most vertebrate taxa were not constrained to specific geological substrates and occurred over larger areas. However, across all three guilds climate was the strongest predictor of regional endemism, particularly for plants wherein discrete hotspots of endemism were buffered from extreme summer temperatures. Land snail and vertebrate endemism peaked in areas with highest precipitation in the driest times of the year. Hotspots of endemism within each guild poorly predicted endemism in other guilds. We found an overarching signal that climatic gradients play a dominant role in the persistence of endemic taxa in an arid‐zone mountain range system. An association with higher rainfall and cooler temperatures indicates that continuing trends toward hotter and drier climates may lead to range contractions in this, and potentially other, arid‐zone mountain biotas. Contrasting patterns of endemism across guilds highlight the need to couple comprehensive regional planning for the protection of climate refugia, with targeted management of more localized and habitat specialist taxa.     

Elevational patterns of species richness and endemism for some important taxa in the Hengduan Mountains, southwestern China

We describe the elevational patterns of species richness and endemism of some important taxa in the Hengduan Mountains, southwest China. Species richness data came from publications, an online database, herbaria and field work. Species richness was estimated by rarefaction and interpolation. The Hengduan Mountains region was divided into a southern and northern subregion, and all species were assigned to four groups based on their distributional range within this region. The conditional autoregressive model (CAR) was used to relate species richness and explanatory variables. The elevational patterns of total, endemic and non-endemic species richness, at subregion and entire region scales, presented to be unimodal and peaked at similar elevations. Area size was strongly related with species richness, and was more powerful in explaining variation in species richness in the northern subregion than in the southern subregion. A single climatic variable (mean annual rainfall, potential evapotranspiration or moisture index) showed a weak relationship with the elevational pattern of species richness. Area and climatic variables together explained more than 67% of the variation in non-endemic richness, 53% in total richness, and 50% in endemic richness. There were three patterns of endemism at the generic level with increasing elevation: namely endemism increased, decreased, or peaked at middle elevations. All selected taxa have experienced rapid speciation and evolution within this region, which plays an important role in the uniform elevational patterns of total, endemic and non-endemic richness, and in the multiform elevational patterns of endemism. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Plant species richness, endemism, and genetic resources in Namibia

Namibia is a floristically diverse, arid to mesic country, with several highly distinct taxa. Including naturalized plants, there are about 4334 vascular plant species and infraspecific taxa within the country's borders, a substantial increase from the existing major reference work. Dominant families are the Poaceae (422species), Fabaceae (377), Asteraceae (385) and Mesembryanthemaceae (177). Freshwater algae and most other groups of lower plants remain poorly known. Concentrations of plant species richness are found in the Succulent Karoo biome, Kaokoveld, Otavi highland/Karstveld area, Okavango Basin, and Khomas highlands. Recent studies have led to a new estimate of 687 endemic plant species, defined as those contained wholly within Namibia's borders, amounting to about 17% of the Namibian flora. At least a further 275 species are Namib Desert endemics shared between the Kaokoveld and southern Angola (75spp.) and between the Succulent Karoo and northwestern South Africa (200spp.). Research on plant genetic resources is focused on species of potential or actual agricultural importance, such as pearl millet, Pennisetum glaucum, and cucurbits. Many wild plants have considerable genetic diversity and development potential. Primary threats to plant diversity fall in the category of poor land management and inappropriate development.     

Species richness and biogeography of non-acarine arachnids in Namibia

A total of 821 species, 296 genera, and 69 families of non-acarine arachnids (Araneae, Solifugae, Scorpiones, Pseudoscorpiones, Opiliones and Amblypygi) are presently known from Namibia. Patterns of spider, solifuge and scorpion species richness and endemism are summarized relative to the 14 major Namibian types of vegetation. Spiders are most speciose in the higher rainfall areas, solifuges in the dry areas, and scorpions in rocky areas. Namibia probably has the world's greatest diversity of solifuges. Many arachnids endemic to Namibia are associated with the Namib Desert sand dunes.     

Relationships between species richness of vascular plants and terrestrial vertebrates in China: analyses based on data of nature reserves

The relationship between plant diversity and animal diversity on a broadscale and its mechanisms are uncertain. In this study, we explored this relationship and its possible mechanisms using data from 186 nature reserves across China on species richness of vascular plants and terrestrial vertebrates, and climatic and topographical variables. We found significant positive correlations between species richness in almost all taxa of vascular plants and terrestrial vertebrates. Multiple regression analyses indicated that plant richness was a significant predictor of richness patterns for terrestrial vertebrates (except birds), suggesting that a causal association may exist between plant diversity and vertebrate diversity in China. The mechanisms for the relationships between species richness of plants and animals are probably dependent on vertebrate groups. For mammals (endothermic vertebrates), this relationship probably represents the integrated effects of plants on animals through trophic links (i.e. providing foods) and non-trophic interactions (i.e. supplying habitats), whereas for amphibians and reptiles (ectothermic vertebrates), this may be a result of the non-trophic links, such as the effects of plants on the resources that amphibians and reptiles require.     

Cladistic biogeography of the Mexican transition zone

Biogeographic relationships among nine montane areas of endemism across the transition zone between North and South America are analysed cladistically based on phylogenetic hypotheses of thirty‐three resident monophyletic taxa of insects, fish, reptiles, and plants. Areas of endemism include the Arizona mountains (AZ), Sonoran Desert (SD), Sierra Madre Occidental (OCC), southern Sierra Madre Occidental (SOC), Sierra Madre Oriental (ORI), Sierra Transvolcanica (TRAN), Sierra Madre del Sur (SUR), Chiapan‐Guatemalan Highlands (CGH), and Talamancan Cordillera (TC). Area relationships are summarized using Brooks Parsimony Analysis and Assumption 0, with the former resulting in more defensible biogeographic hypotheses. Areas of endemism are dividable into two monophyletic groups; a northern group including AZ, SD, OCC, and ORI, and a southern group consisting of TC, CGH, TRAN, SUR, and the isolated southern regions of the Sierra Madre Occidental (SOC). The northern set of areas are characterized by recent, probably Pleistocene, isolation and prevalent widespread species, whereas the southerly areas probably diverged after Pliocene closure of the Panamanian isthmus. The southern areas are redundantly represented on many of the taxon‐area cladograms by endemic species, indicative of much higher levels of endemism in the Sierra Transvolcanica and further south. Use of a general area cladogram in such a transition zone permits explicit exploration of biogeographic patterns and establishes a predictive framework for taxonomy and conservation prioritization.