Biogeography of the Lizard Genus Tropidurus Wied-Neuwied, 1825 (Squamata: Tropiduridae): Distribution,Endemism, and Area Relationships in South America

Based on comprehensive distributional records of the 23 species currently assigned to the lizard genus Tropidurus, we investigated patterns of endemism and area relationships in South America. Two biogeographic methods were applied, Parsimony Analysis of Endemicity (PAE) and Brooks Parsimony Analysis (BPA). Two areas of endemism were detected by PAE: the first within the domains of the semiarid Brazilian Caatinga, which includes seven endemic species, and the second in the region of the Serranía de Huanchaca, eastern Bolivia, in which three endemic species are present. The area cladograms recovered a close relationship between the Atlantic Forest and areas of the South American open corridor. The results revealed a close relationship among the provinces Caatinga (Cerrado, Parana Forest (Pantanal+Chaco)). The uplift of the Brazilian Central Plateau in the Late Pliocene-Early Pleistocene (4-2 Myr BP) has been interpreted as a major event responsible for isolation and differentiation of biotas along these areas. However, we emphasize that without the establishment of a temporal framework concerning the diversification history of Tropidurus it is premature to correlate cladogenetic events with specific time periods or putative vicariant scenarios. The limiting factors hampering the understanding of the biogeographic history of this genus include (1) the absence of temporal references in relation to the diversification of distinct clades within Tropidurus; (2) the lack of an appropriate taxonomic resolution of the species complexes currently represented by widely distributed forms; and (3) the need for a comprehensive phylogenetic hypothesis. We suggest that these three important aspects should be prioritized in future investigations.     

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.     

The lizard assemblage from Seasonally Dry Tropical Forest enclaves in the Cerrado biome, Brazil, and its association with the Pleistocenic Arc

Aim  To determine if the distributions of lizard species from Seasonally Dry Tropical Forest (SDTF) enclaves within the Cerrado biome in central Brazil are associated with the Tropical Seasonal Forests Region, a recently proposed phytogeographic unit of South America, corroborating the existence of a Pleistocenic Arc of SDTFs.
Location  SDTF remnants in the Paranã River valley, municipality of São Domingos, Goiás, Brazil.
Methods  Lizards were extensively sampled using haphazard sampling, funnel traps, and pitfall traps with drift fences during four expeditions. The composition of the SDTF lizard assemblage was compared with those from other South American phytogeographic regions (Caatinga, Cerrado, Chaco, Llanos, and the dry forests of Colombia and Bolivia), based on the literature and our own unpublished data.
Results  The SDTF lizard assemblage contained 20 species, including 11 species with extensive distributions among the regions considered, seven species shared exclusively with Cerrado localities, a single species shared exclusively with other SDTFs, and one endemic species. The presence of Lygodactylus klugei (Smith, Martin & Swain, 1977), presumably endemic to the Pleistocenic Arc formed by the Tropical Seasonal Forests Region, considerably extends the known distribution of this species, suggesting historical connections between Caatinga and Cerrado SDTF enclaves.
Main conclusions  The composition of the lizard assemblage in Cerrado SDTF enclaves seems to corroborate the recent proposal that the SDTF should be recognized as a phytogeographic unit (or dominium). The presence of disjunct populations and endemic species highlights the urgency of considering the uniqueness of the Paranã River valley SDTFs and the importance of its conservation.     

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.     

The history of Seasonally Dry Tropical Forests in eastern South America: inferences from the genetic structure of the tree Astronium urundeuva (Anacardiaceae)

Today, the Seasonally Dry Tropical Forests (SDTF) of eastern South America occur as large, well-defined nuclei (e.g. Caatinga in the northeast) and as smaller enclaves within other vegetations (e.g. Cerrado and Chaco). In order to infer the way the present SDTF distribution was attained, the genetic structure of Astronium urundeuva, a tree confined to SDTF, was assessed using two chloroplast spacers and nine microsatellite loci. Five haplotypes were identified, whose distribution was spatially structured. The distribution of the two most common and divergent haplotypes suggested former vicariance and progressive divergence due to isolation. More recent range expansions of these two lineages subsequently occurred, leading to a secondary contact at the southern limit of the Caatinga SDTF nucleus. The multilocus-Bayesian approach using microsatellites consistently identified three groups of populations (Northeast, Central and Southwest). Isolation by distance was found in Northeast and Southwest groups whereas admixture was detected in the Central group, located at the transition between Caatinga and Cerrado domains. All together, the results support the existence of range expansions and secondary contact in the Central group. This study provides arguments that favour the existence of a previously more continuous formation of SDTF in eastern South America.     

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

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

Misconceptions about parsimony analysis of endemicity

Parsimony analysis of endemicity (PAE) has been widely criticized in the recent literature based on methodology rather than on theory. Here I argue that most of the criticisms of PAE result from confusion between the dynamic and static approaches of PAE, by both users and critics of the method. Originally, PAE (the dynamic approach) was proposed primarily for historical comparisons of biotic distributions based on geological and stratigraphical information; that is, the stratigraphical record of the biota within two or more horizons was used to evaluate changes (layer by layer) in their distributional patterns. This led to an analysis of the biota throughout space and through time. On the other hand, the static approach excluded the temporal component and based the analysis on a single geological horizon. Most problems exemplified and discussed in the literature refer to the static approach. In addition to this defence of the original PAE, I present some new criticisms regarding the application of PAE using artificially delimited areas (for example areas defined by geopolitical boundaries), which may lead to incorrect interpretations. Recently, several variations of static PAE have appeared: some designed to accommodate ecological data (e.g. parsimony analysis of distributions – PAD); others that incorporate phylogenetic content (e.g. cladistic analysis of distributions and endemism – CADE); and some that have been integrated with other historical methods (e.g. panbiogeography) in order to detect and evaluate hypotheses of biogeographical homologies. Biogeographers, both ecological and historical, should be aware of the problems and limitations of both dynamic and static PAE and evaluate new variations of PAE (PAD, CADE, etc.). Finally, I argue in favour of an independent and pluralist discipline of biogeography that treats biogeography as related to systematics but not dependent on it, as some scholars have assumed.     

Geographic variation and systematic review of the lizard genus Vanzosaura (Squamata,Gymnophthalmidae), with the description of a new species

Vanzosaura rubricauda (Boulenger, 1902) is a small‐bodied gymnophthalmid lizard widespread in dry biomes of South America. Throughout its distribution, V. rubricauda populations experience contrasting environmental conditions, and a marked disjunction occurs in the central portion of the Cerrado biome. Previous studies indicate that V. rubricauda may be a species complex, and here we used mitochondrial DNA data and external morphology to test for population differentiation and its systematic implications for the genus. We found three geographically cohesive groups recovered as well resolved, and strongly supported mitochondrial DNA (mtDNA) haploclades that are differentiated with respect to morphometry, meristics, and colour pattern. In light of the observed differences, we propose a new taxonomic rearrangement of the genus, where we: (1) restrict V. rubricauda to Chaco and western Cerrado regions; (2) resurrect V anzosaura multiscutata (Amaral, 1933) comb. nov. for Caatinga populations; and (3) describe V anzosaura savanicola sp. nov. for the eastern Cerrado region of Brazil. The new species is diagnosed from its congeners by having longer limbs and tail, fewer smooth subcaudals, and genetic distances ranging from 5 to 13%. The new species, along with other recent discoveries, highlights the Jalapão‐Serra Geral region as one of the most important areas of endemism for Cerrado squamates. © 2014 The Linnean Society of London     

Distributional modelling and parsimony analysis of endemicity of Senecio in the Mediterranean-type climate area of Central Chile

Aim Floristic blocks and areas of endemism resulting from a parsimony analysis of endemicity (PAE) using raw floristic data versus data generated from distributional modelling for 130 species in the genus Senecio Tourn. ex L. distributed in the Mediterranean‐type climate area of Central Chile were compared, and the results were used to identify conservation priorities for the flora of the region. Location Central Chile, between 30° and 38° S. Methods Using herbarium records, a species × area matrix consisting of presence/absence data was constructed from a 0.5° × 0.5° grid. Distributional modelling techniques incorporating vegetation formations, elevation and the contagion index were used to interpolate floristic composition of poorly known areas. Parsimony analysis of endemicity was used to identify floristic blocks and areas of endemism. Results Using the number of most parsimonious trees as an index, distributional modelling greatly optimized the results of the PAE analysis. Three floristic blocks and four areas of endemism were suggested based on the PAE results using potential distribution data not incorporating the contagion index, while four blocks and two areas of endemism were suggested from the PAE results using potential distribution data incorporating the contagion index. Floristic blocks for the northern coast, southern Andes, and northern/central Andes were found, with some blocks showing divisions within them representing distinct geographic subunits. Major breaks between and within floristic blocks were identified at 32.5°–33° S and 34.5°–35° S. Main conclusions The floristic blocks identified with the distributional modelling and PAE correspond well to results from some previous studies and support hypothesized biogeographic divisions within Central Chile. The results were similar to those obtained from parallel analysis of the entire tree flora of Central Chile. The vegetative formation‐based distributional modelling produced robust and reproducible results when used along with PAE, especially when the contagion index was incorporated, and is a useful technique for area classification. The results demonstrate the utility of Senecio as an indicator genus for biogeography and conservation in southern South America.     

Biogeographical patterns of the avifaunas of the Caribbean Basin Islands: a parsimony perspective

We analyzed the avifaunas of the Caribbean islands and nearby continental areas and their relationships using Parsimony Analysis of Endemicity (PAE), in order to assess biogeographical patterns and their concordance with geological and phylogenetic evidence. Using distributional information of birds obtained from published literature, a presence/absence matrix for 695 genera and 2026 species of land and freshwater birds was constructed and analyzed. Three different analyses were performed: for species, for genera, and for species and genera combined. In the combined analysis, the Lesser Antilles appear paraphyletic at the base of the cladogram. Then, two major clades are identified: South America (Andes, Venezuelan lowlands, Dutch West Indies and Trinidad and Tobago) and North America, including the Greater Antilles in a clade that is the sister area to Yucatan and the Central American countries nested from north to south. PAE results support Caribbean vicariant models and cladistic biogeographical hypotheses on area relationships, and show relative congruence with available phylogenetic data. Bird biogeography on the Caribbean islands appears to have been caused by both vicariance and dispersal processes. © The Willi Hennig Society 2007.     

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.     

Biogeographic regions of North American mammals based on endemism

Since the 19th Century, two regions have been recognized for North American mammals, which overlap in Mexico. The Nearctic region corresponds to the northern areas and the Neotropical region corresponds to the southern ones. There are no recent regionalizations for these regions under the criterion of endemism. In the present study, we integrate two methods to regionalize North America, using species distribution models of mammals: endemicity analysis (EA) and parsimony analysis of endemicity (PAE). EA was used to obtain areas of endemism and PAE was used to hierarchize them. We found 76 consensus areas from 329 sets classified in 146 cladograms, and the strict consensus cladogram shows a basal polytomy with 14 areas and 16 clades. The final regionalization recognizes two regions (Nearctic and Neotropical) and a transition zone (Mexican Transition Zone), six subregions (Canadian, Alleghanian, Californian‐Rocky Mountain, Pacific Central America, Mexican Gulf‐Central America, and Central America), two dominions (Californian and Rocky Mountain), and 23 provinces. Our analysis show that North America is probably more complex than previously assumed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 485–499.     

The implications of Tertiary and Quaternary sea level rise events for avian distribution patterns in the lowlands of northern South America

Aim To assess the correspondence between current avian distributions in the lowlands of northern South America with respect to the hypothesized importance of sea level rise and other events over the past 15 million years on speciation. Location Tropical lowlands of north‐western South America. Methods To establish which bird taxa may have originated in each area of endemism, I examined the ranges of all bird species occurring in the Pacific and the Caribbean lowlands. To determine land and sea distribution during a sea level rise of 100 m in north‐western South America and eastern Central America, I traced the 100 m contour line from the Geoatlas of Georama and the Global 30‐Arc‐Second Elevation Data (GTOPO30) produced by the US Geological Survey. Results During a sea level rise of ～100 m, marine incursions would have occurred from the Caribbean Sea and the Pacific Ocean. Several areas of tropical forest and dry/arid vegetation would have been isolated, currently known as the Darién, Chocó, Nechí, Catatumbo, Tumbesian and Guajiran areas of animal and plant endemism. Main conclusions A large part of the high diversity of forest and nonforest birds and other animals and plants in the Pacific rain forest and the Caribbean woodland likely arose as the result of sea level rises, dispersal and other geological and climatic events.     

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.     

Distributional patterns of the species of <Emphasis Type="Italic">Valeriana</Emphasis> (Valerianaceae) in southern South America

Distributional patterns of Valeriana species from southern South America were analyzed. We prepared a database with the records of 40 species from Argentina and Chile south of 33°S, obtained from herbarium specimens, published taxonomic studies and field trips. We undertook a track analysis and a parsimony analysis of endemicity (PAE), the latter with 134 grid cells of 1.5° latitude by 1.5° longitude. Three generalized tracks and one node were obtained, and three areas of endemism were identified. These general patterns of distribution in Valeriana were used to infer possible vicariance and dispersal events that might have shaped them. We identified a center of diversification in Central Chile and the Argentinean province of Neuquén. It represents a complex area that is related to both the Central Chilean and Subantarctic subregions.     

Areas of endemism in the Antarctic – a case study of the benthic hydrozoan genus Oswaldella (Cnidaria,Kirchenpaueriidae)

Aim The aim of this study is to investigate areas of endemism within the distribution of Oswaldella species in the Southern Ocean, thereby testing previous hypotheses and proposing alternative scenarios for Antarctic evolution. Location Southern Ocean, Antarctic and sub-Antarctic waters of southern South America. Methods We prepared a database for the 31 currently known species of the Antarctic genus Oswaldella, which includes geographical locations gathered from published taxonomic studies as well as materials from museums and expeditions. A parsimony analysis of endemicity (PAE) was used to test hypotheses of distribution patterns. Results Four areas of endemism are hypothesized: southern South America, two high Antarctic areas (eastern and western) and a larger area, mainly in western Antarctica at lower latitudes and including insular areas (but not the Balleny Islands). Main conclusions The results support, in part, previous hypotheses for the Southern Ocean region, while providing more detailed resolution. The areas of endemism may reflect both historical and ecological processes that influenced the Antarctic biota. The Magellanic area reflects the well-known affinities of the Antarctic biota with that of South America and may be a consequence of dispersal through deeper (and colder) waters, followed by speciation. The second area, the largest one, encompasses most of the insular faunas and may also be associated with deeper waters formed since 43 Ma. The third area may be explained by the development of seaways in the circum-Antarctic region beginning 50 Ma. Finally, the fourth zone, with a very poor fauna, coincides with the opening of the Tasman Strait and the formation of the Australo-Antarctic Gulf, associated with a minor wind-driven current.     

Distribution patterns and biogeographic analysis of Austral Polychaeta (Annelida)

Aim We investigate the biogeography of Austral Polychaeta (Annelida) using members of the families Eunicidae, Lumbrineridae, Oenonidae, Onuphidae, Serpulidae and Spionidae and Parsimony Analysis of Endemicity (PAE). We determine whether observed polychaete distribution patterns correspond to traditional shallow-water marine areas of endemism, estimate patterns of endemism and relationships between areas of endemism, and infer the biological processes that have caused these patterns. Location The study is concerned with extant polychaete taxa occupying shallow-water areas derived from the breakup of the Gondwana landmass (i.e. Austral areas). Methods Similarity was assessed using a significance test with Jaccard's indices. Areas not significantly different at 0.99 were combined prior to the PAE. Widespread species and genera (155 taxa) were scored for presence/absence for each area of endemism. PAE was used to derive hypotheses of area relationships. Hierarchical patterns in the PAE trees were identified by testing for congruence with patterns derived from cladistic biogeographic studies of other Gondwanan taxa and with geological evidence. Results The polychaete faunas of four area-pairs were not significantly different and the areas amalgamated: South-west Africa and South Africa, New Zealand South Island and Chatham Islands, Macquarie Island and Antipodean Islands, and West Antarctica and South Georgia. Areas with the highest levels of species endemism were southern Australia (67.0%), South-east South America (53.2%) and South Africa (40.4%). About 60% of species and 7.5% of genera occupied a single area of endemism. The remainder were informative in the PAE. Under a no long-distance dispersal assumption a single minimal-length PAE tree resulted (l=367; ci=0.42); under dispersal allowed, three minimal-length trees resulted (l=278; ci=0.56). In relation to the sister grouping of the New Zealand areas and Australia we find congruence between our minimal-length trees and those derived from a biogeographic study of land plants, and with area relationships predicted by the Expanding Earth Model. Main conclusions The polychaete distribution patterns in this study differ slightly from the classical areas of endemism, most notably in being broader, thereby bringing into question the value of using single provincial system for marine biogeographic studies. The Greater New Zealand region is found to be ‘monophyletic’ with respect to polychaetes, that is comprising a genuine biogeographical entity, and most closely related to the polychaete fauna of southern Australia. This finding is consistent with studies of land plants and with the Expanding Earth model, but disagrees with conventional geology and biogeographic hypothesis involving a ‘polyphyletic’ New Zealand. Both vicariance and concerted range expansion (=biotic dispersion) appear to have played important roles in shaping present-day distribution patterns of Austral polychaetes. Shallow-water ridge systems between the Australian and Greater New Zealand continental landmasses during the Tertiary are thought to have facilitated biotic dispersion.     

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

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

Vicariance and endemism in a Neotropical savanna hotspot: distribution patterns of Cerrado squamate reptiles

Aim To test predictions of the vicariance model, to define basic biogeographical units for Cerrado squamates, and to discuss previous biogeographical hypotheses. Location Cerrado; South American savannas south of the Amazon, extending across central Brazil, with marginal areas in Bolivia and Paraguay and isolated relictual enclaves in adjacent regions. Methods We compiled species occurrence records via field sampling and revision of museum specimens and taxonomic literature. All species were mapped according to georeferenced locality records, and classified as (1) endemic or non‐endemic, (2) typical of plateaus or depressions, and (3) typical of open or forested habitats. We tested predictions of the vicariance model using biotic element analysis, searching for non‐random clusters of species ranges. Spatial congruence of biotic elements was compared with putative areas of endemism revealed by sympatric restricted‐range species. Effects of topographical and vegetational mosaics on distribution patterns were studied according to species composition in biotic elements and areas of endemism. Results We recorded 267 Cerrado squamates, of which 103 (39%) are endemics, including 20 amphisbaenians (61% endemism), 32 lizards (42%) and 51 snakes (32%). Distribution patterns corroborated predictions of the vicariance model, revealing groups of species with significantly clustered ranges. An analysis of endemic species recovered seven biotic elements, corroborating results including non‐endemics. Sympatric restricted‐range taxa delimited 10 putative areas of endemism, largely coincident with core areas of biotic elements detected with endemic taxa. Distribution patterns were associated with major topographical and vegetational divisions of the Cerrado. Endemism prevailed in open, elevated plateaus, whereas faunal interchange, mostly associated with forest habitats, was more common in peripheral depressions. Main conclusions Our results indicate that vicariant speciation has strongly shaped Cerrado squamate diversity, in contrast to earlier studies emphasizing faunal interchange and low endemism in the Cerrado vertebrate fauna. Levels of squamate endemism are higher than in any other Cerrado vertebrate group. The high number of recovered endemics revealed previously undetected areas of evolutionary relevance, indicating that biogeographical patterns in the Cerrado were poorly represented in previous analyses. Although still largely undocumented, effects of vicariant speciation may be prevalent in a large fraction of Cerrado and Neotropical biodiversity.