Areas of endemism and spatial diversification of the Muscidae (Insecta: Diptera) in the Andean and Neotropical regions

Aim We present a biogeographical analysis of the areas of endemism and areas of diversification in the Muscidae. This analysis searched for geographical patterns in the Muscidae to reconstruct elements of the evolutionary biogeographical history of this insect family. Location Andean and Neotropical regions. Method We constructed a geographic database of 728 species from the literature and museum specimens. Areas of endemism were established by parsimony analysis of endemicity (PAE) based on grids of two different sizes: 5° (550 × 550 km) and 2° (220 × 220 km). Areas of diversification were delimited by track analysis that also included phylogenetic information. This process was independently applied to 11 genera. For each genus, we plotted generalized tracks generated by sister species on a map. When these generalized tracks supported inter‐generic nodes they were manually contoured and inferred to be areas of diversification for the Muscidae. Results Thirteen endemic areas were found using the 5° grid, and eight endemic areas resulted from the 2° grid. Ten areas were in agreement with previous studies, and 11 were new. Amazonian and Atlantic areas of diversification agreed with previous areas for the genus Polietina, and new areas of diversification were found in Panama and in central Chile. Main conclusions Six spatial patterns in the Muscidae were identified: (1) areas of endemism in both Pampa and Puna provinces were established with species whose distributions had not previously been analysed; (2) a new area of endemism was established in extreme southern South America, in Tierra del Fuego; (3) two new areas of diversification, which include Panama and central Chile, were identified; (4) a spatial association was identified between the separation of Chiloe Island from the continent and the diversification in Andean species; (5) a north–south track axis and latitudinal node intervals were identified, interpreted as spatial responses to glaciation or glacial retreat in the Andes; and (6) a spatial coincidence of areas of endemism, of diversification and high species richness in the Muscidae was discovered. The analysis of a complete database and the recognition of areas of diversification are extremely important in elucidating novel biogeographical patterns, which will in turn contribute to a better understanding of the geographical patterns of evolution in the Muscidae.     

Distribution patterns of the genus Pacifigorgia (Octocorallia: Gorgoniidae): track compatibility analysis and parsimony analysis of endemicity

Aim We analysed the distribution patterns of the eastern Pacific octocoral genus Pacifigorgia and deduced its ancestral distribution to determine why Pacifigorgia is absent from the Gulf of Mexico, the Caribbean of central America, and the Antilles. We also examined the current patterns of endemism for Pacifigorgia to look for congruence between hot spots of endemism in the genus and generally recognized areas of endemism for the eastern Pacific. Location The tropical eastern Pacific and western Atlantic, America. Methods We used track compatibility analysis (TCA) and parsimony analysis of endemicity (PAE) to derive ancestral distribution patterns and hot spots of endemism, respectively. Distributional data for Pacifigorgia were gathered from several museum collections and from fieldwork, particularly in the Pacific of Costa Rica and Panama. Results A single generalized track joined the three main continental eastern Pacific biogeographical provinces and the western Atlantic. This track can be included within a larger eastern Atlantic–eastern Pacific transoceanic track that may be the oldest transoceanic track occurring in the region. PAE results designate previously recognized eastern Pacific biogeographical provinces as Pacifigorgia hot spots of endemism. The number of endemic species, which for other taxonomic groups is similar among the eastern Pacific provinces, is higher in the Panamic province for Pacifigorgia. Main conclusions We propose that the absence of Pacifigorgia from the Gulf of Mexico, the Caribbean of central America, and the Antilles is the result of an ancient absence of the genus from these areas rather than the consequence of a major, recent, extinction episode. The Cortez province and the Mexican province appear together as a result of either non‐response to vicariance or dispersal across the Sinaloan Gap. We posit that the Central American Gap acts as a barrier that separates the Panamic province from the northern Cortez–Mexican province.     

Towards a generalized biogeography of the Southern Ocean benthos

Aim To investigate whether the biogeographical regions proposed by J. W. Hedgpeth and widely adopted by other authors hold true, are an oversimplification or with further data might show a unified Antarctic province. Location Southern Hemisphere. Methods The distributions of 1318 species of bivalves, 4656 species of gastropods, 1465 species of cheilostome and 167 species of cyclostome bryozoans were analysed for 29 regions in the Southern Hemisphere, including South American, South African, Tasmanian, New Zealand, sub‐Antarctic and Antarctic regions. We present data on species richness, rates of endemism, patterns of radiation, faunal similarities and multivariate biogeographical analyses. Results The most striking pattern to emerge from our data set of species counts per region was a strong east–west hemispheric asymmetry, with high species numbers in New Zealand, Tasmania and South Africa and low numbers in South America. In contrast, no difference was found in richness between the east and west parts of the Southern Ocean. We compared findings in our model taxa with published data on ascidians, cephalopods and pycnogonids. Further evidence of strong faunal links between the Antarctic and South America is reported in this study, although we found little evidence for a biogeographical relationship between the Antarctic or South America and New Zealand/Tasmania. Strong evidence exists for a long‐term influence of the Antarctic Circumpolar Current upon the distribution of Southern Ocean benthos. This is demonstrated by the reduced prevalence of South American species in the Antarctic and sub‐Antarctic with increasing distance from South America in the direction of the current. Three of our four study taxa (bivalves, cheilostomes and cyclostomes) show the Southern Ocean as a ‘single functional unit’ with no evidence for a biogeographical split between east and west. Main conclusions Unlike the biogeographical schemes previously proposed, we show that biogeographical regions in the Southern Ocean differ depending upon the class of animals being considered. Despite this we suggest that some general rules are viable, including species endemism rates of around 50%, a single Antarctic province and a definite distinction between the sub‐Antarctic islands influenced by South America and those of New Zealand.     

Atlantic reef fish biogeography and evolution

Aim To understand why and when areas of endemism (provinces) of the tropical Atlantic Ocean were formed, how they relate to each other, and what processes have contributed to faunal enrichment. Location Atlantic Ocean. Methods The distributions of 2605 species of reef fishes were compiled for 25 areas of the Atlantic and southern Africa. Maximum‐parsimony and distance analyses were employed to investigate biogeographical relationships among those areas. A collection of 26 phylogenies of various Atlantic reef fish taxa was used to assess patterns of origin and diversification relative to evolutionary scenarios based on spatio‐temporal sequences of species splitting produced by geological and palaeoceanographic events. We present data on faunal (species and genera) richness, endemism patterns, diversity buildup (i.e. speciation processes), and evaluate the operation of the main biogeographical barriers and/or filters. Results Phylogenetic (proportion of sister species) and distributional (number of shared species) patterns are generally concordant with recognized biogeographical provinces in the Atlantic. The highly uneven distribution of species in certain genera appears to be related to their origin, with highest species richness in areas with the greatest phylogenetic depth. Diversity buildup in Atlantic reef fishes involved (1) diversification within each province, (2) isolation as a result of biogeographical barriers, and (3) stochastic accretion by means of dispersal between provinces. The timing of divergence events is not concordant among taxonomic groups. The three soft (non‐terrestrial) inter‐regional barriers (mid‐Atlantic, Amazon, and Benguela) clearly act as ‘filters’ by restricting dispersal but at the same time allowing occasional crossings that apparently lead to the establishment of new populations and species. Fluctuations in the effectiveness of the filters, combined with ecological differences among provinces, apparently provide a mechanism for much of the recent diversification of reef fishes in the Atlantic. Main conclusions Our data set indicates that both historical events (e.g. Tethys closure) and relatively recent dispersal (with or without further speciation) have had a strong influence on Atlantic tropical marine biodiversity and have contributed to the biogeographical patterns we observe today; however, examples of the latter process outnumber those of the former.     

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.     

Input data, analytical methods and biogeography of Elegia (Restionaceae)

Aim The aim of this paper is to determine the optimal methods for delimiting areas of endemism for Elegia L. (Restionaceae), an endemic genus of the Cape Floristic Region. We assess two methods of scoring the data (presence–absence in regular grids, or in irregular eco‐geographical regions) and three methods for locating biogeographical centres or areas of endemism, and evaluate one method for locating biotic elements. Location The Cape Floristic Region (CFR), South Africa. Methods The distribution of all 48 species of Elegia was mapped as presence–absence data on a quarter‐degree grid and on broad habitat units (eco‐geographical areas). Three methods to delimit areas of endemism were applied: parsimony analysis of endemism (PAE), phenetic cluster analysis, and NDM (‘end em ism’). In addition, we used presence–absence clustering (‘Prabclus’) to delimit biotic elements. The performances of these methods in elucidating the geographical patterns in Elegia were compared, for both types of input data, by evaluating their efficacy in maximizing the proportion of endemics and the number of areas of endemism. Results Eco‐geographical areas perform better than quarter‐degree grids. The eco‐geographical areas are potentially more likely to track the distribution of species. The phenetic approach performed best in terms of its ability to delimit areas of endemism in the study area. The species richness and the richness of range‐restricted species are each highest in the south‐western part of the CFR, decreasing to the north and east. The phytogeographical centres identified in the present study are the northern mountains, the southern mountains (inclusive of the Riviersonderend Mountains and the Cape Peninsula), the Langeberg range, the south coast, the Cape flats, and the west coast. Main conclusions This study demonstrates that (1) eco‐geographical areas should be preferred over a grid overlay in the study of biogeographical patterns, (2) phenetic clustering is the most suitable analytical method for finding areas of endemism, and (3) delimiting biotic elements does not contribute to an understanding of the biogeographical pattern in Elegia. The areas of endemism in Elegia are largely similar to those described in other studies, but there are many detailed differences.     

Historical biogeography and diversification within the Neotropical parrot genus Pionopsitta (Aves: Psittacidae)

Aim We investigate spatial and temporal patterns of diversification within the Neotropical avifauna using the phylogenetic history of parrots traditionally belonging to the genus Pionopsitta Bonaparte 1854. This genus has long been of interest for those studying Neotropical biogeography and diversity, as it encompasses species that occur in most Neotropical forest areas of endemism. Location The Neotropical lowland forests in South and Central America. Methods Phylogenetic relationships were investigated for all species of the genus Pionopsitta and five other short‐tailed parrot genera using complete sequences of the mitochondrial genes cyt b and ND2 as well as 26 plumage characters. The resulting phylogeny was used to test the monophyly of the genus, investigate species limits, and as a framework for reconstructing their historical biogeography and patterns of diversification. Results We found that the genus Pionopsitta, as previously defined, is not monophyletic and thus the Chocó, Central American and Amazonian species will now have to be placed in the genus Gypopsitta. The molecular and morphological phylogenies are largely congruent, but disagree on the position of one of the Amazon basin taxa. Using molecular sequence data, we estimate that species within Gypopsitta diversified between 8.7 and 0.6 Ma, with the main divergences occurring between 3.3 and 6.4 Ma. These temporal results are compared to other taxa showing similar vicariance patterns. Main conclusions The results suggest that diversification in Gypopsitta was influenced mainly by geotectonic events, marine transgressions and river dynamics, whereas Quaternary glacial cycles of forest change seem to have played a minor role in the origination of the currently recognized species.     

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

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

Historical biogeography of cynolebiasine annual killifishes inferred from dispersal–vicariance analysis

Aim To analyse the biogeographical events responsible for the present distribution of cynolebiasine killifishes (Teleostei: Rivulidae: Cynolebiasini), a diversified and widespread Neotropical group of annual fishes threatened with extinction. Location South America, focusing on the main river basins draining the Brazilian Shield and adjacent zones. Methods Phylogenetic analysis of 214 morphological characters of 102 cynolebiasine species using tnt , in conjunction with dispersal–vicariance analysis (diva ) based on the distribution of cynolebiasine species among 16 areas of endemism. Results The basal cynolebiasine node is hypothesized to be derived from an old vicariance event occurring just after the separation of South America from Africa, when the terrains at the passive margin of the South American plate were isolated from the remaining interior areas. This would have been followed by geodispersal events caused by river‐capturing episodes from the adjacent upland river basins to the coastal region. Optimal ancestral reconstructions suggest that the diversification of the tribe Cynolebiasini in north‐eastern South America was first caused by vicariance events in the Paranã–Urucuia–São Francisco area, followed by dispersal from the São Francisco to the Northeastern Brazil area. The latter dispersal event occurred simultaneously in two different cynolebiasine clades, possibly as a result of a temporary connection of the São Francisco area before the uplift of the Borborema Plateau during the Miocene. The diversity of cynolebiasines inhabiting the Paraguay area is hypothesized to be derived from two processes: an older vicariance event (about 30 Ma) separating Paraguay from southern Amazonian areas (Guaporé–Xingu–Araguaia–Tocantins), and a series of more recent dispersal and vicariance events (about 15–11 Ma) caused by successive marine transgressions, which permitted alternating biotic exchange and isolation in the Paraguay, La Plata, Negro and Patos areas. Main conclusions diva indicates there to have been a series of vicariance events congruent with tectonic episodes in South America, but the present distribution of cynolebiasines has also been shaped by a series of dispersal events. The effects of the combined action of dispersal and vicariance events were more conspicuous in the Eastern Brazil and Paraguay areas, thus generating reticulate biogeographical scenarios.     

Exploring the Afromontane centre of endemism: Kniphofia Moench (Asphodelaceae) as a floristic indicator

Aim The genus Kniphofia contains 71 species with an African–Malagasy distribution, including one species from Yemen. The genus has a general Afromontane distribution. Here we explore whether Kniphofia is a floristic indicator of the Afromontane centre of endemism and diversity. The South Africa Centre of diversity and endemism was explored in greater detail to understand biogeographical patterns. Location Africa, Afromontane Region, southern Africa, Madagascar and Yemen. Methods Diversity and endemism for the genus were examined at the continental scale using a chorological approach. Biogeographical patterns and endemism in the South Africa Centre were examined in greater detail using chorology, phenetics, parsimony analysis of endemicity (PAE) and mapping of range‐restricted taxa. Results Six centres of diversity were recovered, five of which are also centres of endemism. Eight subcentres of diversity are proposed, of which only two are considered subcentres of endemism. The South Africa Centre is the most species‐rich region and the largest centre of endemism for Kniphofia. The phenetic analysis of the South Africa Centre at the full degree square scale recovered three biogeographical areas that correspond with the subcentres obtained from the chorological analysis. The PAE (at the full degree square scale) and the mapping of range‐restricted taxa recovered two and six areas of endemism (AOEs), respectively. These latter two approaches produced results of limited value, possibly as a result of inadequate collecting of Kniphofia species. Only two AOEs were identified by PAE and these are embedded within two of the six AOEs recovered by the mapping of range‐restricted taxa. All the above AOEs are within the three subcentres found by chorological and phenetic analysis (at the full degree square scale) for the South Africa Centre. Main conclusions The centres for Kniphofia broadly correspond to the Afromontane regional mountain systems, but with some notable differences. We regard Kniphofia as a floristic indicator of the Afromontane Region sensu lato. In southern Africa, the phenetic approach at the full‐degree scale retrieved areas that correlate well with those obtained by the chorological approach.     

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.     

Systematics and biogeography of Rhodniini (Heteroptera: Reduviidae: Triatominae) based on 16S mitochondrial rDNA sequences

Aim The tribe Rhodniini is one of six comprising the subfamily Triatominae (Heteroptera: Reduviidae), notorious as blood‐sucking household pests and vectors of Trypanosoma cruzi throughout Latin America. The human and economic cost of this disease in the American tropics is considerable, and these bugs are unquestionably of great importance to man. Studies of the evolution, phylogeny, biogeography, ecology, physiology and behaviour of the Rhodniini are needed to help improve existing Chagas’ disease control programmes. The objective of the study reported here was to propose biogeographical hypotheses to explain the modern geographical distribution of the species of Rhodniini. Location Neotropical region. Methods We employed mitochondrial rDNA sequences (16S) currently available in GenBank to align sequences of Rhodniini species using ClustalX. The analyses included 16S sequences from predatory reduviid subfamilies (Stenopodainae, Ectrichodiinae, Harpactorinae, Reduviinae and Salyavatinae) present in GenBank as an outgroup. Cladistic analysis used the program PAUP to derive trees based on maximum parsimony (MP) and maximum likelihood (ML). Known distribution data for Rhodniini species were obtained from reviews and plotted on maps of South and Central America using the program iMap. An area cladogram was derived from the cladistic result to show the historical connections among the studied taxa and the endemic areas. The program TreeMap (Jungle Edition) was used to deduce taxon–area associations where the optimal solutions to explain the biogeographical hypothesis of the Rhodniini in the Neotropics were those with lowest total cost. Results Parsimony and maximum‐likelihood analysis of 16S rDNA sequences included 14 species of Rhodniini, as well as five species of predatory Reduviidae representing five of the predatory subfamilies. Tanglegrams were used to show the relationship between the Neotropical areas of endemism and Rhodniini species. When TreeMap with codivergence (vicariance) events were weighted as 0 and duplication (sympatry), lineage losses (extinction) and host switching (dispersal) were all weighted as 1, 20 scenarios were found to explain the biogeographical history of Rhodniini in the Neotropical region. Twelve of the optimal solutions with the lowest total cost were used to explain the biogeography of the Rhodniini in the Neotropics. These optimal reconstructions require six vicariance events, 20 duplications (sympatry), at least three dispersals, and at least one extinction event. Main conclusions The Rhodniini have a complex biogeographical history that has involved vicariance, duplications (sympatry), dispersal and extinction events. The main geological events affecting the origin and diversification of the Rhodniini in the Neotropics were (1) uplift of the Central Andes in the Miocene or later, (2) break‐up of the Andes into three separate cordilleras (Eastern, Central and Western) in the Plio‐Pleistocene, (3) formation of a land corridor connecting South and North America in the Pliocene, and (4) uplift of the Serra do Mar and Serra da Mantiqueira mountain systems between the Oligocene and Pleistocene. The relationships and biogeographical history of the species of Rhodniini in the Neotropical region probably arose from the areas of endemism shown in our work.     

Phylogenetic biogeography of the leafy liverwort Herbertus (Jungermanniales, Herbertaceae) based on nuclear and chloroplast DNA sequence data: correlation between genetic variation and geographical distribution

Aim The cosmopolitan genus Herbertus is notorious for having a difficult taxonomy and for the fact that there is limited knowledge of species ranges and relationships. Topologies generated from variable molecular markers are used to discuss biogeographical patterns in Herbertus and to compare them with the geological history of continents and outcomes reported for other land plants. Location Africa, Asia, Azores, Europe, southern South America, northern South America, North America, New Zealand. Methods Phylogenetic analyses of nuclear ribosomal internal transcribed spacer and chloroplast (cp) trnL–trnF sequences of 66 accessions of Herbertus and the outgroup species Triandrophyllum subtrifidum and Mastigophora diclados were used to investigate biogeographical patterns in Herbertus. Areas of putative endemism were defined based on the distribution of species included in the analyses. Maximum parsimony analyses were undertaken to reconstruct ancestral areas and intraspecies migration routes. Results The analyses reveal species‐level cladograms with a correlation between genetic variation and the geographical distribution of the related accessions. The southern South American Herbertus runcinatus is sister to the remainder of the genus, which is split into two main clades. One contains the Neotropical–African Herbertus juniperoideus and the New Zealand/Tasmanian Herbertus oldfieldianus. An African accession of H. juniperoideus is nested within Neotropical accessions. The second main clade includes species that inhabit Asia, the Holarctic, Africa, and northern South America. Maximum parsimony analyses indicate that this clade arose in Asia. Herbertus sendtneri originated in Asia and subsequently colonized the Holarctic and northern South America. An Asian origin and colonization into Africa is indicated for H. dicranus. Main conclusions The current distribution of Herbertus cannot be explained by Gondwanan vicariance. A more feasible explanation of the range is a combination of short‐distance dispersal, rare long‐distance dispersal events (especially into regions that faced floral displacements as a result of climatic changes) extinction, recolonization, and diversification. The African Herbertus flora is a mixture of Asian and Neotropical elements. Southern South America harbours an isolated species. The molecular data indicate partial decoupling of molecular and morphological variation in Herbertus. Biogeographical patterns in Herbertus are not dissimilar to those of other groups of bryophytes, but elucidation of the geographical ranges requires a molecular approach. Some patterns could be the result of maintenance of Herbertus in the inner Tropics during glacial maxima, and dispersal into temperate regions in warm phases.     

Testing the island effect in adaptive radiation: rates and patterns of morphological diversification in Caribbean and mainland Anolis lizards

Many of the classic examples of adaptive radiation, including Caribbean Anolis lizards, are found on islands. However, Anolis also exhibits substantial species richness and ecomorphological disparity on mainland Central and South America. We compared patterns and rates of morphological evolution to investigate whether, in fact, island Anolis are exceptionally diverse relative to their mainland counterparts. Quite the contrary, we found that rates and extent of diversification were comparable--Anolis adaptive radiation is not an island phenomenon. However, mainland and Caribbean anoles occupy different parts of morphological space; in independent colonizations of both island and mainland habitats, island anoles have evolved shorter limbs and better-developed toe pads. These patterns suggest that the two areas are on different evolutionary trajectories. The ecological causes of these differences are unknown, but may relate to differences in predation or competition among mainland and island communities.     

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.     

ORIGIN AND EVOLUTION OF CONTINENTAL BIOTAS: SPECIATION AND HISTORICAL CONGRUENCE WITHIN THE AUSTRALIAN AVIFAUNA

Factors governing the origin and evolution of continental biotas were investigated using an analysis of speciation patterns within the Australian avifauna. Phylogenetic relationships within seven lineages of birds were analyzed by numerical cladistic techniques applied to data sets of morphological characters. These relationships revealed extensive congruence among the spatial and temporal histories of lineages whose species are endemic to common areas of endemism. A general hypothesis is constructed to explain this congruence in which widespread biotas are postulated to have been partitioned into areas of endemism by the origin of geomorphological and/or ecological-climatic barriers. Congruence in these phylogenetic patterns of differentiation suggests the following historical pattern of interrelationships for areas of endemism along the northern and eastern coasts of Australia: (Kimberley Plateau + Arnhem Land) + ([New Guinea + Cape York Peninsula] + [Atherton Plateau + Eastern Coastal Rainforest]). Likewise, this study indicates that the arid interior avifauna was segregated into two closely related biotas (Eastern and Western Desert biotas) by the Eyrean Barrier. These biotas are, in turn, related to a more mesic avifauna that was itself subdivided into areas of endemism located in the Southwest and Southeast corners of the continent.     

Neat and clear: 700 species of crane flies (Diptera: Tipulomorpha) link southern South America and Australasia

A biogeographical analysis of crane flies (Diptera, Tipulomorpha) in the southern hemisphere is used to test if their distribution patterns provide evidence of biogeographical homology (shared history) in the South Pacific. Crane fly distributions are interpreted in light of patterns of endemism and diversity and published phylogenetic studies. A panbiogeographical approach, assuming that repeating distribution patterns strongly suggest the existence of past connections between the areas (biogeographical homology), is used. A clear pattern is revealed in which crane fly taxa shared between southern South America, New Zealand and Australia are restricted to that region. Thirty genera and subgenera, together comprising about 700 species, occur in both South America and Australasia and only in these areas. This distribution defines the limits of the South Pacific Track, a standard biogeographical pattern displayed by many taxa, including the southern beeches (Nothofagus). Although the distribution of some taxa spans the entire track, others are present in parts of the areas only, forming a nested set of distributions. Within the surveyed genera and subgenera, all individual species are endemic to one single region or continent, suggesting vicariance as the main process behind crane fly disjunctions in this part of the world. The nested set of distribution patterns could be explained by extinctions in areas where taxa were present previously. Alternatively, it may indicate historical absences and the existence of a heterogeneous set of ancestral distributional ranges. ‘Gondwanan’ may not be the best term for the observed disjunctions, which should be labelled as trans‐Pacific instead. Recent molecular estimates of divergence times suggest a Permian origin of the earliest extant Diptera lineages such as the Tipulomorpha, followed by fast radiation in the Triassic. Although the differentiation of some crane fly groups occurring in the region may have been driven by recent Mesozoic and Cenozoic events of continental breakup, as least part of the fauna may have evolved allopatrically in response to older events. This may explain the overlapping distribution of subgenera in large genera such as Gynoplistia.     

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.     

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.