Panbiogeography of Nothofagus (Nothofagaceae): analysis of the main species massings

Aim The aim of this paper is to analyse the biogeography of Nothofagus and its subgenera in the light of molecular phylogenies and revisions of fossil taxa. Location Cooler parts of the South Pacific: Australia, Tasmania, New Zealand, montane New Guinea and New Caledonia, and southern South America. Methods Panbiogeographical analysis is used. This involves comparative study of the geographic distributions of the Nothofagus taxa and other organisms in the region, and correlation of the main patterns with historical geology. Results The four subgenera of Nothofagus have their main massings of extant species in the same localities as the main massings of all (fossil plus extant) species. These main massings are vicariant, with subgen. Lophozonia most diverse in southern South America (north of Chiloé I.), subgen. Fuscospora in New Zealand, subgen. Nothofagus in southern South America (south of Valdivia), and subgen. Brassospora in New Guinea and New Caledonia. The main massings of subgen. Brassospora and of the clade subgen. Brassospora/subgen. Nothofagus (New Guinea–New Caledonia–southern South America) conform to standard biogeographical patterns. Main conclusions The vicariant main massings of the four subgenera are compatible with largely allopatric differentiation and no substantial dispersal since at least the Upper Cretaceous (Upper Campanian), by which time the fossil record shows that the four subgenera had evolved. The New Guinea–New Caledonia distribution of subgenus Brassospora is equivalent to its total main massing through geological time and is explained by different respective relationships of different component terranes of the two countries. Global vicariance at family level suggests that Nothofagaceae/Nothofagus evolved largely as the South Pacific/Antarctic vicariant in the breakup of a world‐wide Fagales ancestor.     

Nestedness of Southern Ocean island biotas: ecological perspectives on a biogeographical conundrum

Aim To use patterns of nestedness in the indigenous and non‐indigenous biotas of the Southern Ocean islands to determine the influence of dispersal ability on biogeographical patterns, and the importance of accounting for variation in dispersal ability in their subsequent interpretation, especially in the context of the Insulantarctic and multi‐regional hypotheses proposed to explain the biogeography of these islands. Location Southern Ocean islands. Methods Nestedness was determined using a new metric, d1 (a modification of discrepancy), for the indigenous and introduced seabirds, land birds, insects and vascular plants of 26 Southern Ocean islands. To assess the possible confounding effects of spatial autocorrelation on the results, islands were assigned to 11 major island groups and each group was treated as a single island in a following analysis. In addition, nestedness of the six Southern Ocean islands comprising the South Pacific Province (New Zealand islands) was analysed. All analyses were conducted for species and genera, for each of the taxa on its own, and for the complete data sets. Results Statistically significant nestedness was found in all of the taxa examined, with nestedness declining in the order seabirds > land birds > vascular plants > insects for the indigenous species. Vagility had a marked influence on nestedness and the biogeographical patterns shown by the indigenous species. This influence was borne out by additional analyses of marine taxa and small‐sized terrestrial species, both of which were more nested than the most nested group examined here, the seabirds. Assemblages of non‐indigenous species also showed nestedness, and nestedness was generally more pronounced than in the indigenous species. Surprisingly, vagility had a significant effect on nestedness in these assemblages too. Main conclusions Nestedness analyses provide a quantitative means of comparing biogeographical patterns for groups differing in vagility. These comparisons revealed that vagility has a considerable influence on biogeographical patterns and should be taken into account in analyses. Here, investigations of more vagile taxa support hypotheses for a single origin of the Southern Ocean island biota (the Insulantarctica scenario), whilst those of less mobile taxa support the more commonly held, multi‐regional hypothesis. All biogeographical analyses across the Southern Ocean (and elsewhere) will be influenced by the effects of dispersal ability, with composite analyses dominated by sedentary groups likely to favour multi‐regional scenarios, and those dominated by mobile groups favouring single origins. Mechanisms underlying nestedness in the region range from nested physiological tolerances in more mobile groups to colonization ability and patterns of speciation in less vagile taxa. Considerable nestedness in the non‐indigenous assemblages is largely a consequence of the fact that many of these species are European weedy species.     

Biogeographical and geological evidence for a smaller, completely-enclosed Pacific Basin in the Late Cretaceous

Aim To use biogeographical, palaeomagnetic, palaeosedimentary, and plate circuit data from Late Cretaceous regions in and around the Pacific to test the plate tectonic hypothesis of a pre‐Pacific superocean. Location East Asia, Australia, Antarctica, the western Americas, and the Pacific. Methods Literature surveys of the distributions of Cretaceous, circum‐Pacific taxa were compared with palaeomagnetic and palaeosedimentary data. Uncontroversial plate motions based on seafloor spreading data were also used to test the results of the biogeographical and palaeomagnetic analyses. Results The distributions of Cretaceous terrestrial taxa, mostly dinosaurs, imply direct, continental connections between Australia and East Asia, East Asia and North America, North America and South America, South America and Antarctica, and Antarctica and Australia. Palaeomagnetic, palaeosedimentary, and basic plate circuit analyses require little to no latitudinal motion of the Pacific plate with respect to the surrounding continents. Specifically, the data implies that western North America, East Asia, and the Pacific plate all increased in latitude by roughly the same amount (c. 11 ± 5°) since the Campanian – and that the Pacific Ocean Basin has increased in length north‐to‐south. Main conclusions Each of the analyses provides independent corroboration for the same conclusion: the Late Cretaceous Pacific plate was completely enclosed by the surrounding continents and has not experienced significant latitudinal motion with respect to North America, East Asia, or the Bering land bridge. This contrasts significantly with the plate tectonic history of the Pacific, implying instead that the Pacific plate formed in situ, pushing the continents apart as the plate and basin expanded. These results also substantiate recent biogeographical analyses that have concluded that a narrower Pacific Ocean Basin in the Mesozoic and early Tertiary provides the most reasonable explanation for the great number of trans‐Pacific disjunctions of poor dispersing taxa.     

NOTHOFAGUS AND PACIFIC BIOGEOGRAPHY

Abstract — Gondwanan biogeography, particularly the relationships between southern South America, New Zealand, Australia, New Guinea and New Caledonia, has been much studied. Nothofagus is often used as the "test taxon", and many papers have been directed at using Nothofagus to explain Gondwanan biogeography. Cladistic biogeographers, working on plant material, have generally failed to find congruence among taxa expected from the southern Pacific disjunctions. New morphological and molecular data on the phytogeny of Nothofagus have re-opened the issue, and we analysed these data to construct a new hypothesis of the biogeography of the genus. We assembled all plant taxa for which we could find reasonably robust phylogenetic hypotheses, and sought a parsimonious biogeographical pattern common to all. Two analyses, based on different assumptions, produced the same general areacladogram. We use the general area-cladogram, in conjunction with the fossil record of Nothofagus to construct a historical scenario for the evolution of the genus. This scenario indicates extensive extinction, but also suggests that Australia has a more recent relationship to New Zealand than to southern South America. This is not congruent with the current geological theories, nor with the patterns evident from insect biogeography. We suggest that concordant dispersal is an unlikely explanation for this pattern, and propose that the solution might be found in alternative geological hypotheses.     

Island evolutionary biogeography: analysis of the weevils (Coleoptera: Curculionidae) of the Falkland Islands (Islas Malvinas)

Aim I analysed distributional and phylogenetic information on weevils (Coleoptera: Curculionidae) from the Falklands, and integrated it with molecular, palaeontological and geological information to infer a geobiotic scenario. Location Falkland Islands (Islas Malvinas). Methods The panbiogeographical analysis was based on data on 23 Falkland species and their related taxa from southern South America. For the cladistic biogeographical analysis I analysed six weevil taxa for which phylogenetic hypotheses are available (the generic groups Cylydrorhinus, Strangaliodes and Falklandius, and the genera Antarctobius, Germainiellus and Puranius). Results from this analysis were compared with previous regionalizations. Cenocrons (sets of taxa that share the same biogeographical history) were identified by considering temporal information provided by fossils and molecular clocks. Finally, a geobiotic scenario was proposed by integrating the available information. Results Six generalized tracks were detected: Maule–Valdivian forests, Magellanic forest, Magellanic moorland, Falkland Islands, Magellanic forest–Magellanic moorland, and Magellanic forest–Falkland Islands. A node was identified in the Magellanic forest, based on the overlap of two generalized tracks. A single general area cladogram was obtained, implying the following sequence: (Magellanic moorland (Maule–Valdivian forests (Magellanic forest, Falkland Islands))). The Falklands are classified here as a biogeographical province in the Austral realm, Andean region and Subantarctic subregion. Falkland weevils seem to belong to a single Subantarctic cenocron. The sequence of events deduced implies the following steps: development of the Subantarctic biota in southern South America, arrival of the Falkland crustal block from South Africa in the Early Cretaceous, geodispersal of the Subantarctic cenocron from southern South America to the Falklands during the Early Oligocene, vicariance of the Magellanic moorland, vicariance of the Maule–Valdivian forests, and final vicariance between the Magellanic forest and the Falkland Islands. Main conclusions The biotic components identified support the connection of the Falkland weevils with the Magellanic forest. Falkland weevils belong to a single cenocron, dated to at least the Early Oligocene, when geodispersal from southern South America may have occurred. An older African cenocron may have been replaced completely by the Subantarctic one when the proto‐Falklands made contact with the Patagonian continental shelf. A geobiotic scenario implying vicariance events related to sea‐level variations could explain the distributional patterns analysed herein.     

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.     

Tackling the taxonomic impediment: a global assessment for ant‐nest beetle diversity (Coleoptera: Carabidae: Paussini)

We evaluated the completeness and historical trends of the taxonomic knowledge of the myrmecophilous ground beetle tribe Paussini (Coleoptera, Carabidae, Paussinae). Species accumulation curves were modelled using a logistic function. Similar analyses were conducted for genera and subgenera. Although not all biogeographical regions have been equally explored, accumulation curves reached a plateau in all cases. Our models predict that about 96% of the world fauna has been already described. However, the asymptotes calculated for the Australian and Oriental species should be interpreted as false plateaus because of the lack of recent research. Similarly, patterns of genera accumulation indicate that a plateau has been reached. As a result of continued debate on the use and validity of Paussini subgenera, the accumulation curves of subgenera showed stepped patterns, with no evidence of plateaus. Thus, although relatively few species are expected to be described in the future, the species accumulation curves indicate that the taxonomic inventory is not yet complete. Differences in accumulation patterns among biogeographical regions can be used to highlight the areas where more species are expected, and hence where taxonomical efforts should be concentrated. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 330–339.     

Biogeographical patterns in endoparasite communities of a marine fish (Sebastes capensis Gmelin) with extended range in the Southern Hemisphere

Aim The endoparasites of Sebastes capensis Gmelin are examined over most of its geographical range (coasts of Peru, Chile, Argentina and South Africa) to determine: (1) whether the endoparasite communities of this fish show zoogeographical patterns; and (2) if so, whether there are any relationships between spatial variations in the endoparasite fauna and known zoogeographical patterns for marine free‐living organisms (e.g. prey that are included in the life cycles of endoparasites). Location Fish were captured at nine localities along the Pacific coast of South America, from 11° S in the centre of the Peruvian coast, to 52° S in southern Chile, and also at two localities in the Atlantic Ocean, at 43° S in Argentina, and 34° S in South Africa. Methods From April to September 2003 and April to August 2004, 626 fish were captured. Endoparasites and diet were examined following traditional methods. Cluster analyses were used to evaluate the distribution patterns of the endoparasite communities, and to evaluate similarities in the prey composition per locality. Results The endoparasite fauna of S. capensis consisted of four species widely distributed along the Pacific coast: Ascarophis cf. sebastodis, Anisakis sp., Corynosoma australe, and Pseudopecoelus sp. Other parasites were distributed only in some geographical areas. The species richness of the parasite communities increased with latitude along the Pacific coast, while parasite communities from Argentina and South Africa showed low species richness. Cluster analyses based on endoparasite composition and on prey composition grouped localities in a way consistent with known biogeographical areas for marine free‐living organisms. Main conclusions The endoparasites of S. capensis exhibit a pattern associated with known biogeographical areas for free‐living organisms. The latitudinal increase in endoparasite community richness is associated with changes in prey composition (intermediate hosts) and also possibly with the presence of definitive hosts. Therefore, the biogeographical patterns of prey are considered key determinants of the endoparasite community structure of the host.     

Historical biogeography of Australian Rhamnaceae, tribe Pomaderreae

Aim To discover the pattern of relationships of areas of endemism for Australian genera in the plant family Rhamnaceae tribe Pomaderreae for comparison with other taxa and interpretation of biogeographical history. Location Australian mainland, Tasmania and New Zealand. Methods A molecular phylogeny and geographic distribution of species within four clades of Pomaderreae are used as a basis for recognition of areas of endemism and analysis of area relationships using paralogy‐free subtrees. The taxon phylogeny is the strict consensus tree from a parsimony analysis of 54 taxa, in four clades, and sequence data for the internal transcribed spacer regions of ribosomal DNA (ITS1‐5.8S‐ITS2) and the plastid DNA region trnL‐F. Results The biogeographical analysis identified five subtrees, which, after parsimony analysis, resulted in a minimal tree with 100% consistency and seven resolved nodes. Three sets of area relationships were identified: the areas of Arnhem and Kimberley in tropical north Australia are related based on the phylogeny of taxa within Cryptandra; the moister South‐west of Western Australia, its sister area the coastal Geraldton Sandplains, the semi‐arid Interzone region and arid Western Desert are related, based on taxa within Cryptandra, Spyridium, Trymalium and Pomaderris; and the eastern regions of Queensland, McPherson‐Macleay, south‐eastern New South Wales (NSW), Victoria, southern Australia, Tasmania and New Zealand are related based on Cryptandra, Pomaderris and Spyridium. Tasmania and NSW are related based entirely on Cryptandra, but the position of New Zealand relative to the other south‐eastern Australian regions is unresolved. Main conclusions The method of paralogy‐free subtrees identified a general pattern of geographic area relationships based on Australian Pomaderreae. The widespread distribution of clades, the high level of endemicity and the age of fossils for the family, suggest that the Pomaderreae are an old group among the Australian flora. Their biogeographical history may date to the early Palaeogene with subsequent changes through to the Pleistocene.     

The trans-Pacific zipper effect: disjunct sister taxa and matching geological outlines that link the Pacific margins

Aim To combine analyses of trans‐Pacific sister taxa with geological evidence in order to test the hypothesis of the existence of a Panthalassa superocean. Location The study is concerned with taxa, both fossil and extant, from East Asia, Australia, New Zealand, South America and North America. Methods Phylogenetic and distributional analyses of trans‐Pacific biota were integrated with geological evidence from the Pacific and circum‐Pacific regions. Results A series of recent biogeographical analyses delineates a zipper‐like system of sister areas running up both margins of the Pacific, with each section of western North and South America corresponding to a particular section from East Asia/Australia/New Zealand. These sister areas coincide neatly with a jigsaw‐like fit provided by the matching Mesozoic coastlines that bracket the Pacific. Main conclusions The young age (<200 Myr) of oceanic crust, the matching Mesozoic circum‐Pacific outlines, and a corresponding system of interlocking biogeographical sister areas provide three independent avenues of support for a closed Pacific in the Upper Triassic–Lower Jurassic. The hypothesis of the existence and subsequent subduction of the pre‐Pacific superocean Panthalassa is not only unnecessary, it conflicts with this evidence. Panthalassa‐based paleomaps necessitate the invention of dozens of additional hypotheses of species‐dependent, trans‐oceanic dispersal events, often involving narrow‐range taxa of notoriously limited vagility, in order to explain repeated examples of the same biogeographical pattern. Removing the vanished‐superocean hypothesis reunites both the matching geological outlines and all the disjunct sister taxa. In brief, what appears to be a multi‐era tangle of convoluted, trans‐oceanic distributions on Panthalassa‐based paleomaps is actually a relatively simple biogeographical pattern that is explainable by a single vicariant event: the opening and expansion of the Pacific.     

CHLOROPLAST-DNA PHYLOGENETICS AND BIOGEOGRAPHY IN A RETICULATING GROUP: STUDY IN POA (POACEAE)

Cladistic analysis of Poa chloroplast DNA (cpDNA) restriction sites tested previously hypothesized relationships within the genus. Forty-six taxa representing 19 sections or groups and three subgenera of Poa and two out-group genera, Puccinellia and Bellardiochloa, are analyzed. Five major and several minor cpDNA groups are identified. The cpDNA cladogram is generally congruent with the subgeneric taxonomy of Poa. Exceptions are reclassified or are discussed in terms of character incompatibilities and possible reticulation events. The cpDNA tree detected relationships among sections that were unresolved using traditional character sets and provides a basis for polarization of morphological character states. An assessment of biogeographic events based on the cpDNA tree suggests: 1) Poa originated in Eurasia; 2) at least six groups of species independently colonized North America; and 3) two of the latter groups colonized South America, and one closely related group colonized New Zealand and Australia. The cpDNA tree provided a conservative estimate of the number of amphi-neotropical disjunctions when compared to the known number of species disjunctions.     

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.     

The biogeography of Gunnera L.: vicariance and dispersal

Aim The genus Gunnera is distributed in South America, Africa and the Australasian region, a few species reaching Hawaii and southern Mexico in the North. A cladogram was used to (1) discuss the biogeography of Gunnera and (2) subsequently compare this biogeographical pattern with the geological history of continents and the patterns reported for other Southern Hemisphere organisms. Location Africa, northern South America, southern South America, Tasmania, New Zealand, New Guinea/Malaya, Hawaii, North America, Antarctica. Methods A phylogenetic analysis of twenty‐six species of Gunnera combining morphological characters and new as well as published sequences of the ITS region, rbcL and the rps16 intron, was used to interpret the biogeographical patterns in Gunnera. Vicariance was applied in the first place and dispersal was only assumed as a second best explanation. Results The Uruguayan/Brazilian Gunnera herteri Osten (subgenus Ostenigunnera Mattfeld) is sister to the rest of the genus, followed sequentially upwards by the African G. perpensa L. (subgenus Gunnera), in turn sister to all other, American and Australasian, species. These are divided into two clades, one containing American/Hawaiian species, the other containing all Australasian species. Within the Australasian clade, G. macrophylla Blume (subgenus Pseudogunnera Schindler), occurring in New Guinea and Malaya, is sister to a clade including the species from New Zealand and Tasmania (subgenus Milligania Schindler). The southern South American subgenus Misandra Schindler is sister to a clade containing the remaining American, as well as the Hawaiian species (subgenus Panke Schindler). Within subgenus Panke, G. mexicana Brandegee, the only North American species in the genus, is sister to a clade wherein the Hawaiian species are basal to all south and central American taxa. Main conclusions According to the cladogram, South America appears in two places, suggesting an historical explanation for northern South America to be separate from southern South America. Following a well‐known biogeographical pattern of vicariance, Africa is the sister area to the combined southern South America/Australasian clade. Within the Australasian clade, New Zealand is more closely related to New Guinea/Malaya than to southern South America, a pattern found in other plant cladograms, contradictory to some of the patterns supported by animal clades and by the geological hypothesis, respectively. The position of the Tasmanian G. cordifolia, nested within the New Zealand clade indicates dispersal of this species to Tasmania. The position of G. mexicana, the only North American species, as sister to the remaining species of subgenus Panke together with the subsequent sister relation between Hawaii and southern South America, may reflect a North American origin of Panke and a recolonization of South America from the north. This is in agreement with the early North American fossil record of Gunnera and the apparent young age of the South American clade.     

Globally basal centres of endemism: the Tasman-Coral Sea region (south-west Pacific), Latin America and Madagascar/South Africa

The New Zealand wrens (Acanthisittidae) are basal in passerine birds and in New Caledonia, the closest country to New Zealand, Amborella is basal in angiosperms. A review of molecular studies indicates that 29 other locally or regionally endemic clades around the Tasman and Coral Sea basins have cosmopolitan or globally widespread sister groups. Other areas that have local endemics basal to diverse global groups include Borneo, Madagascar/South Africa/Tanzania, southern China–Taiwan–Japan, and different parts of Latin America, especially the Guayana Plateau and western Mexico. Basal clades are widely interpreted as ancestral and their location is generally taken to represent a centre of origin for the group as a whole. In the present study, basal groups are treated simply as small (less speciose) sister groups. The Tasman and western Mexico/Caribbean regions have important biogeographic and tectonic ties with each other and with the central Pacific. Large igneous provinces (Ontong Java, Hikurangi‐Manihiki, and Gorgona Plateaus) formed in the central Pacific in the Cretaceous. Fossil wood is found on the Ontong Java Plateau, and formerly emergent seamounts up to 24 km across occur on Hikurangi Plateau. Many terranes in New Zealand, New Caledonia, New Guinea and western America represent former island arcs (or their products) that formed in the central Pacific and later accreted to the Pacific margins. Long‐term survival of taxa as metapopulations on the ephemeral volcanic islands and atolls of plate margins and fissures may explain the biogeographical connections among the Tasman region, the central Pacific, and the accreted terranes of western America. Branching sequences in cladograms and phylogenetic trees have been interpreted as dispersal events, but instead probably indicate the sequence of differentiation in an already widespread ancestor. Major disjunctions of tens of thousands of kilometres often occur between taxa at consecutive nodes on a tree and dispersal by physical movement is unlikely. The break between the globally basal centres and the rest of the world marks the initial site of differentiation of a widespread ancestor, with subsequent or more or less simultaneous differentiation occurring in other areas. Differentiation of the modern angiosperms, passerines and other groups first took place around the Tasman region, or at least the terranes now accumulated there, and then around other centres, especially Madagascar–South Africa and Mexico–north‐west South America. Angiosperms are now recognized as basal to extant gymnosperms and major tectonic dynamism around the globally basal centres during the Mesozoic, involving terrane accretion, orogeny, and rifting could have been involved with the last important modernization of angiosperms, birds and other groups. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96 , 222–245.     

Antitropicality and convergent evolution: a case study of Permian neospiriferine brachiopods

Antitropical distribution is a biogeographical pattern characterized by natural occurrences of the same species or members of the same clade in the middle‐ or middle‐to‐high‐latitudinal habitats of both hemispheres, either on land or in marine environments, without appearing in the intervening tropical environments. For most of the noted examples of Permian antitropical distribution, particularly in marine invertebrates, the causes of disjunctions have been mainly linked to either dispersal or vicariance models. Little attention has been paid to other possible mechanisms. This study investigated the antitropicality of some Permian neospiriferine brachiopods through detailed taxonomic revision, comparison of palaeobiogeographical distribution, and a phylogenetic analysis. Several species, previously assigned to Kaninospirifer, are here reassigned to other genera, especially to Fasciculatia in the northern hemisphere and to Quadrospira in the southern hemisphere during the Permian. Both Kaninospirifer and Fasciculatia appear to have been restricted to north‐western Pangea and north‐eastern Asia during the Permian, but there is no robust evidence to suggest their presence in the southern hemisphere to which Imperiospira and Quadrospira were confined. In spite of the distributional separation between the two pairs of neospiriferine genera in the Permian palaeobiogeographical regime, they share considerable numbers of morphological characters, such as a large shell, subdued fasciculation, and reduction of ventral adminicula. Notwithstanding these morphological similarities, our phylogenetic reconstruction of the neospiriferines does not support a close relationship between these genera. This therefore must indicate that these similar morphological features were independently acquired, probably with these taxa living in spatially separate but ecologically compatible environmental conditions in the mid‐latitudinal area of each hemisphere during the Permian. We regard this as an example of convergent evolution.     

The historical biogeography of the southern group of the sucker genus Moxostoma (Teleostei: Catostomidae) and the colonization of central Mexico

The historical biogeography of the southern group of Moxostoma Rafinesque, 1820, a genus of Nearctic freshwater fishes belonging to the Catostomidae, along its entire distribution in North America was inferred to: (1) determine the biogeographical events responsible for its current pattern of diversity and distribution; (2) correlate the climatic and geologic history of the region with the biogeographical pattern observed; and (3) trace the colonization route into central Mexico and the western Pacific slope drainages. The sequences of mitochondrial cytochrome b and the third intron of the growth hormone were obtained for the members of the southern group and related species of the Catostomidae. Phylogenetic analyses and relaxed molecular clock analyses were performed to determine the relatedness of the species and to estimate divergence times. To uncover biogeographical patterns, a dispersal–extinction–cladogenesis (DEC) analysis was conducted. The phylogenetic analyses were consistent with the historical hydrographic scenario in the region. The divergence times show that the southern group evolved during the Pliocene–Pleistocene. The DEC analyses showed that vicariance and dispersal played an important role in the current distribution patterns of the lineages in central Mexico, and allow us to trace an independent route of colonization from the northern areas of North America into central Mexico.     

Patterns of endemism in south-eastern Pacific benthic polychaetes of the Chilean coast

Aim In this study we evaluate patterns of endemism for benthic polychaete species along the southeastern Pacific coast of Chile. Our goals were (1) to describe latitudinal gradients of endemism and identify areas of high endemism, (2) to evaluate the effect of biogeographical limits on endemism patterns, and (3) to evaluate indirectly the role played by evolutionary dynamics on patterns of endemism. Location South‐eastern Pacific coast of Chile, ranging from Arica (18° S) to Cape Horn (56° S). Methods We used a list of 178 species of endemic, shallow benthic polychaetes to evaluate patterns of endemism. Parsimony analysis of endemicity (PAE) and the endemism index (EI) were used to evaluate hierarchical relationships of endemism between different latitudinal bands, and to identify areas with high degrees of endemism and differences in endemism. We evaluated the effect of biogeographical limits on endemic polychaete fauna by testing for the existence of geometric constraints (mid‐domain effect). The role of evolutionary dynamics on latitudinal patterns of endemism was evaluated with nestedness analysis (NA) using the temperature index. Results The PAE analysis indicated two large, separate areas of endemism: (1) the northern area between 18° S and 38° S, and (2) the southern area between 39° S and 56° S. The endemism index showed a maximum value (32 species) around 39°–41° S. Species‐richness curves of each 3° band of latitude showed a clear mid‐domain effect (69%), but the two maximum points of species richness at mid‐latitudes (36° S to 38° S and 39° S to 41° S) did not correspond to the mid‐domain peak in species richness, presenting a greater number of species than expected by the mid‐domain effect. The nestedness analysis showed that the number of genera reaches a maximum of 70 at mid‐latitudes (36°–41° S), decreasing towards both the northern and southern areas. The spatial distribution of the entire data set of endemic species showed a nested pattern (T° = 24.5°, P < 0.0001). Main conclusions Our results strongly support the existence of a latitudinal gradient of endemism for benthic polychaete species along the Chilean coast. The shape of this gradient is clearly non‐linear, with a marked peak of endemism occurring at mid‐latitudes (36°–41° S, endemism hotspot), which also corresponds to a peak in species richness. Furthermore, this hotspot is the midpoint separating two distinct areas of endemism to the north and south. We suggest that the observed pattern of endemism for benthic polychaete taxa of the Chilean coast can be explained by a combination of geometric constraints and historical mechanisms, such as the processes that affected the Chilean coast during the Neogene (e.g. ENSO, oxygen minimum zone, glaciations).     

The Neotropical genus Stibadocerina Alexander and its phylogenetic relationship to other Stibadocerinae genera: further evidence of an ancestral trans‐Pacific biota (Diptera: Cylindrotomidae)

Abstract Stibadocerina Alexander, a monotypic genus, includes the only known Neotropical species of the family Cylindrotomidae, S. chilensis Alexander, 1929 , from South Central Chile (ca. 36°50′S–42°17′S). In this paper, Stibadocerina chilensis is redescribed and illustrated in detail. A study of wing‐vein homology in the subfamily Stibadocerinae is provided, to identify the components of the reduced radial sector in Stibadocerina and related taxa. The proposed hypotheses of wing‐vein homology are tested, and the systematic position of Stibadocerina is assessed through a cladistic analysis of 13 characters of the male imago, scored for exemplar species of the four genera included in the Stibadocerinae. A single most parsimonious tree supports the monophyly of the Stibadocerinae and the following relationships among its included genera: Stibadocerodes [Stibadocera (Stibadocerella +Stibadocerina)]. The subfamily includes one example of a vicariant distribution with a sister‐group relationship between South Central Chilean and East Asian taxa, and supports a biogeographical interpretation of an ancestral trans‐Pacific biota.     

Biogeography of Nicotiana section Suaveolentes (Solanaceae) reveals geographical tracks in arid Australia

Aim Nicotiana section Suaveolentes is largely endemic to Australia but includes one species endemic to Africa, one to New Caledonia and Tongatupa, and one to the Marquesas Islands in the Pacific. Other sections of Nicotiana are found in the New World. In Australia, Suaveolentes is widespread across the continent, with many taxa adapted to the Eremean zone. We aim to analyse the biogeography of the Australian clade, both to shed light on the evolution of the group and to determine general area relationships that provide insight into the history of the arid‐zone biota. Location Mesic and arid regions of continental Australia, the Central–South Pacific and Namibia, Africa. Methods A phylogeny of Suaveolentes, based on morphology and molecular data, was used to analyse the relationships of areas in which the taxa occur. The section is monophyletic, and all but three taxa were included (25). The method of paralogy‐free subtree analysis was employed, with the basal taxon Nicotiana africana used as the outgroup. Results Paralogy‐free subtree analysis found five area subtrees that, when combined, resulted in a minimal area cladogram with six resolved nodes. Pacific and mesic eastern Australia (including Lord Howe Island) are at the base of the area cladogram, followed by the differentiation of North West Australia and later South East Australia. Arid regions of Australia are related, revealing three biogeographical tracks: a northern track including the Great Sandy Desert and Tanami, which are related to the Pilbara; a central track relating the Western Desert, Central Ranges, Eastern Desert and North East Interzone; and a southern track relating the South West Interzone, Nullarbor, Adelaide/Eyre and the South East Interzone. Plesiomorphic taxa with chromosome number n = 24–23 occur on the periphery of the continent, and derived taxa with n = 21, 20, 18, 16–15 identify the tracks across arid Australia. Main conclusions The patterns of distribution and differentiation of Suaveolentes in Australia show that the age of the clade is at least Early Miocene, dating to before the onset of aridification in Australia about 15 Ma. The patterns are also interpreted as evidence that it was vicariance that largely shaped speciation in the Eremean zone, with range expansion of some widespread taxa probably occurring in the most recent cycles of severe drying and mobilization of desert dune sands.     

Systematic placement and biogeographical relationships of the monotypic genera Gypothamnium and Oxyphyllum (Asteraceae: Mutisioideae) from the Atacama Desert

Gypothamnium and Oxyphyllum (Asteraceae) are two monotypic genera endemic to the Atacama Desert of northern Chile. We performed a phylogenetic analysis using published sequences of the plastid rbcL and ndhF genes, the trnL‐trnF region and the nuclear ribosomal internal transcribed spacer (ITS) to assess the systematic placement of the two genera within Mutisioideae. On the basis of the phylogenetic results, we constructed area cladograms to explore the biogeographical relationships and origin of the genera. The phylogenetic analysis showed that Gypothamnium is closely related to Aphylloclados, Plazia, Urmenetia, Lycoseris and Onoseris, whereas Oxyphyllum is closely related to Leucheria, Moscharia, Polyachyrus and, with low support, Jungia. These results do not differ substantially from those proposed in previous treatments based on morphological characters. The biogeographical analysis suggests that Gypothamnium in the coastal Atacama Desert is related to taxa that are currently distributed in eastern subtropical South America and in the Puna. Oxyphyllum may have originated from central Chile and other areas in southern South America, but its sister group (Leucheria + Polyachyrus) also reaches the Puna and the coastal Atacama Desert. Both groups show ancestral affinities with elements currently distributed in north‐western South America and Mesoamerica. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 32–51.