Southern hemisphere biogeography inferred by event-based models: plant versus animal patterns

The Southern Hemisphere has traditionally been considered as having a fundamentally vicariant history. The common trans-Pacific disjunctions are usually explained by the sequential breakup of the supercontinent Gondwana during the last 165 million years, causing successive division of an ancestral biota. However, recent biogeographic studies, based on molecular estimates and more accurate paleogeographic reconstructions, indicate that dispersal may have been more important than traditionally assumed. We examined the relative roles played by vicariance and dispersal in shaping Southern Hemisphere biotas by analyzing a large data set of 54 animal and 19 plant phylogenies, including marsupials, ratites, and southern beeches (1,393 terminals). Parsimony-based tree fitting in conjunction with permutation tests was used to examine to what extent Southern Hemisphere biogeographic patterns fit the breakup sequence of Gondwana and to identify concordant dispersal patterns. Consistent with other studies, the animal data are congruent with the geological sequence of Gondwana breakup: (Africa(New Zealand(southern South America, Australia))). Trans-Antarctic dispersal (Australia <--> southern South America) is also significantly more frequent than any other dispersal event in animals, which may be explained by the long period of geological contact between Australia and South America via Antarctica. In contrast, the dominant pattern in plants, (southern South America(Australia, New Zealand)), is better explained by dispersal, particularly the prevalence of trans-Tasman dispersal between New Zealand and Australia. Our results also confirm the hybrid origin of the South American biota: there has been surprisingly little biotic exchange between the northern tropical and the southern temperate regions of South America, especially for animals.     

Plant dispersal N.E.W.S from New Zealand

Although New Zealand separated from Gondwana during the late Cretaceous (80 million years ago) it shares strong floristic affinities with other Southern Hemisphere landmasses. For 150 years, biogeographers have debated whether these similarities reflect the ancient Gondwanan connection or subsequent dispersal events. Molecular phylogenies are providing new insights into the history of Southern Hemisphere plant groups. These studies show that many plant lineages are recent arrivals in New Zealand, diversifying rapidly and then travelling to other Southern Hemisphere landmasses.     

Gondwanan radiation of the Southern Hemisphere crayfishes (Decapoda: Parastacidae): evidence from fossils and molecules

Aim The sequential break‐up of Gondwana is thought to be a dominant process in the establishment of shared biota across landmasses of the Southern Hemisphere. Yet similar distributions are shared by taxa whose radiations clearly post‐date the Gondwanan break‐up. Thus, determining the contribution of vicariance versus dispersal to seemingly Gondwanan biota is complex. The southern freshwater crayfishes (family Parastacidae) are distributed on Australia and New Guinea, South America, Madagascar and New Zealand and are unlikely to have dispersed via oceans, owing to strict freshwater limitations. We test the hypotheses that the break‐up of Gondwana has led to (1) a predominately east–west (((Australia, New Zealand: 80 Ma) Madagascar: 160–121 Ma) South America: 165–140 Ma), or (2) a southern (((Australia, South America: 52–35 Ma) New Zealand: 80 Ma) Madagascar: 160–121 Ma) pattern for parastacid crayfish. Further, we examine the evidence for a complete drowning of New Zealand and subsequent colonization by freshwater crayfish. Location Southern Hemisphere. Methods The evolutionary relationships among the 15 genera of Parastacidae were reconstructed using mitochondrial [16S, cytochrome c oxidase subunit I (COI)] and nuclear (18S, 28S) sequence data and maximum likelihood and Bayesian methods of phylogenetic reconstruction. A Bayesian (multidivtime ) molecular dating method using six fossil calibrations and phylogenetic inference was used to estimate divergence time among crayfish clades on Gondwanan landmasses. Results The South American crayfish are monophyletic and a sister group to all other southern crayfish. Australian crayfish are not monophyletic, with two Tasmanian genera, Spinastacoides and Ombrastacoides, forming a clade with New Zealand and Malagasy crayfish (both monophyletic). Divergence of crayfish among southern landmasses is estimated to have occurred around the Late Jurassic to Early Cretaceous (109–178 Ma). Main conclusions The estimated phylogenetic relationships and time of divergence among the Southern Hemisphere crayfishes were consistent with an east–west pattern of Gondwanan divergence. The divergence between Australia and New Zealand (109–160 Ma) pre‐dated the rifting at around 80 Ma, suggesting that these lineages were established prior to the break‐up. Owing to the age of the New Zealand crayfish, we reject the hypothesis that there was a complete drowning of New Zealand crayfish habitat.     

Low genetic differentiation between two geographically separated populations of demersal gadiform fishes in the Southern Hemisphere

The distribution patterns of many fishes between the three continents (Africa, Australia, and South America) in the Southern Hemisphere have been uncovered to be influenced by mostly vicariance or historical dispersal. Although some demersal fishes with intercontinental distribution are suggested to be more influenced by current/recent dispersal, few genetic studies have been made for demersal fishes so far. To provide more information for such fishes, genetic divergence was analyzed for two pairs of gadiform species and subspecies distributed around Australasia and South America: the blue grenadier, Macruronus novaezelandiae (from New Zealand) and the Patagonian grenadier, M. magellanicus (from South America) as well as two subspecies of the southern blue whiting, Micromesistius australis pallidus (from New Zealand) and M. a. australis (from South America). The sequence analyses of two mitochondrial DNA regions showed no divergence between Australasian and South American populations of the grenadiers and the southern blue whiting. The microsatellite DNA analysis also indicated significant but very minimal genetic differentiation between the two geographic populations of each pair. These results imply rather recent separation of the two geographic populations. Current/recent dispersal may be an important common factor for determining the distribution of demersal fishes in the Southern Hemisphere. Nonetheless, low but significant genetic differentiation observed requires treating the two populations of the economically important grenadiers and southern blue whiting, respectively, as different stocks for proper resource management.     

Process and pattern in the biogeography of New Zealand – a global microcosm?

Aim  To describe New Zealand's historical terrestrial biogeography and place this history in a wider Southern Hemisphere context.
Location  New Zealand.
Methods  The analysis is based primarily on literature on the distributions and relationships of New Zealand's terrestrial flora and fauna.
Results  New Zealand is shown to have a biota that has broad relationships, primarily around the cool Southern Hemisphere, as well as with New Caledonia to the north. There are hints of ancient Gondwanan taxa, although the long-argued predominance of taxa derived by vicariant processes, driven by plate tectonics and the fragmentation of Gondwana, is no longer accepted as a principal explanation of the biota's origins and relationships.
Main conclusions  Most of the terrestrial New Zealand flora and fauna has clearly arrived in New Zealand much more recently than the postulated separation of New Zealand from Gondwana, dated at c. 80 Ma. There is a view that New Zealand may have disappeared completely beneath the sea in the early Cenozoic, and acceptance of this would mean derivation of the entire biota by transoceanic dispersal. However, there are elements in the biota that seem to have broad distributions that date back to Gondwanan times, and also some that are thought unlikely to have been able to disperse to New Zealand across ocean gaps, especially freshwater organisms. Very strong connections to the biota of Australia, rather than to South America, are inconsistent with the timing of New Zealand's ancient and early separation from Gondwana and seem likely to have resulted from dispersal.     

Historical transoceanic dispersal of a freshwater shrimp: the colonization of the South Pacific by the genus Paratya (Atyidae)

Aim To infer the phylogenetic relationships within the freshwater shrimp genus Paratya Miers, 1882 (Atyidae) and to use these data to answer biogeographical questions about the location, timing and form of evolution of this genus in the South Pacific. Location Paratya are spread throughout various freshwater habitats in the western Pacific, with a disjunct northern range in the North Pacific (Japan, Korea, Ryukyu Islands, Siberia) and South Pacific (Australia, New Zealand, New Caledonia, Lord Howe, Norfolk Island). Methods Specimens were obtained from throughout its range. Mitochondrial sequences of cytochrome oxidase subunit I and 16S ribosomal DNA were analysed using phylogenetic techniques to identify whether landmasses are monophyletic and what the relationships are between landmasses. Molecular clock dating methods were used to date divergences between taxa. Results Each landmass was recovered as monophyletic. Japan/Ryukyu Islands is the most basal group, followed by New Zealand. Australian specimens form a sister group to a clade made up of two groups (New Caledonia and Lord Howe/Norfolk Island). The oldest divergence within the genus (between North and South Pacific) took place 12–19 Ma. Main conclusions The geographical origin of the genus (either Gondwana or Laurasia) is unclear. Dispersal occurred between the North and South Pacific long after the split up of Gondwana. Dispersal likely explains the presence of Paratya on each landmass in the South Pacific, from continent to isolated oceanic island. This dispersal is conjectured to have taken place through oceanic currents because of the amphidromous life cycle of some taxa of Paratya, given that amphyidromy is plesiomorphic in atyid shrimp.     

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.     

From East Gondwana to Central America: historical biogeography of the Alstroemeriaceae

Aim The Alstroemeriaceae is among 28 angiosperm families shared between South America, New Zealand and/or Australia; here, we examine the biogeography of Alstroemeriaceae to better understand the climatic and geological settings for its diversification in the Neotropics. We also compare Alstroemeriaceae with the four other Southern Hemisphere families that expanded from Patagonia to the equator, to infer what factors may have permitted such expansions across biomes. Location South America, Central America, Australia and New Zealand. Methods Three chloroplast genes, one mitochondrial gene and one nuclear DNA region were sequenced for 153 accessions representing 125 of the 200 species of Alstroemeriaceae from throughout the distribution range; 25 outgroup taxa were included to securely infer evolutionary directions and be able to use both ingroup and outgroup fossil constraints. A relaxed‐clock model relied on up to three fossil calibrations, and ancestral ranges were inferred using statistical dispersal–vicariance analysis (S‐DIVA). Southern Hemisphere disjunctions in the flowering plants were reviewed for key biological traits, divergence times, migration directions and habitats occupied. Results The obtained chronogram and ancestral area reconstruction imply that the most recent common ancestor of Colchicaceae and Alstroemeriaceae lived in the Late Cretaceous in southern South America/Australasia, the ancestral region of Alstroemeriaceae may have been South America/Antarctica, and a single New Zealand species is due to recent dispersal from South America. Chilean Alstroemeria diversified with the uplift of the Patagonian Andes c. 18 Ma, and a hummingbird‐pollinated clade (Bomarea) reached the northern Andes at 11–13 Ma. The South American Arid Diagonal (SAAD), a belt of arid vegetation caused by the onset of the Andean rain shadow 14–15 Ma, isolated a Brazilian clade of Alstroemeria from a basal Chilean/Argentinean grade. Main conclusions Only Alstroemeriaceae, Calceolariaceae, Cunoniaceae, Escalloniaceae and Proteaceae have expanded and diversified from Patagonia far into tropical latitudes. All migrated northwards along the Andes, but also reached south‐eastern Brazil, in most cases after the origin of the SAAD. Our results from Alstroemeria now suggest that the SAAD may have been a major ecological barrier in southern South America.     

Biogeography of the Monimiaceae (Laurales): a role for East Gondwana and long‐distance dispersal,but not West Gondwana

Aim The biogeography of the tropical plant family Monimiaceae has long been thought to reflect the break‐up of West and East Gondwana, followed by limited transoceanic dispersal. Location Southern Hemisphere, with fossils in East and West Gondwana. Methods We use phylogenetic analysis of DNA sequences from 67 of the c. 200 species, representing 26 of the 28 genera of Monimiaceae, and a Bayesian relaxed clock model with fossil prior constraints to estimate species relationships and divergence times. Likelihood optimization is used to infer switches between biogeographical regions on the highest likelihood tree. Results Peumus from Chile, Monimia from the Mascarenes and Palmeria from eastern Australia/New Guinea form a clade that is sister to all other Monimiaceae. The next‐deepest split is between the Sri Lankan Hortonia and the remaining genera. The African Monimiaceae, Xymalos monospora, then forms the sister clade to a polytomy of five clades: (I) Mollinedia and allies from South America; (II) Tambourissa and allies from Madagascar and the Mascarenes; (III) Hedycarya, Kibariopsis and Leviera from New Zealand, New Caledonia and Australia; (IV) Wilkiea, Kibara, Kairoa; and (V) Steganthera and allies, all from tropical Australasia. Main conclusions Tree topology, fossils, inferred divergence times and ances‐tral area reconstruction fit with the break‐up of East Gondwana having left a still discernible signature consisting of sister clades in Chile and Australia. There is no support for previous hypotheses that the break‐up of West Gondwana (Africa/South America) explains disjunctions in the Monimiaceae. The South American Mollinedia clade is only 28–16 Myr old, and appears to have arrived via trans‐Pacific dispersal from Australasia. The clade apparently spread in southern South America prior to the Andean orogeny, fitting with its first‐diverging lineage (Hennecartia) having a southern‐temperate range. The crown ages of the other major clades (II–V) range from 20 to 29 Ma, implying over‐water dispersal between Australia, New Caledonia, New Zealand, and across the Indian Ocean to Madagascar and the Mascarenes. The endemic genus Monimia on the Mascarenes provides an interesting example of an island lineage being much older than the islands on which it presently occurs.     

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.     

Out of the Palaeotropics? Historical biogeography and diversification of the cosmopolitan ectomycorrhizal mushroom family Inocybaceae

Aim The ectomycorrhizal (ECM) mushroom family Inocybaceae is widespread in north temperate regions, but more than 150 species are encountered in the tropics and the Southern Hemisphere. The relative roles of recent and ancient biogeographical processes, relationships with plant hosts, and the timing of divergences that have shaped the current geographic distribution of the family are investigated. Location Africa, Australia, Neotropics, New Zealand, north temperate zone, Palaeotropics, Southeast Asia, South America, south temperate zone. Methods We reconstruct a phylogeny of the Inocybaceae with a geological timeline using a relaxed molecular clock. Divergence dates of lineages are estimated statistically to test vicariance‐based hypotheses concerning relatedness of disjunct ECM taxa. A series of internal maximum time constraints is used to evaluate two different calibrations. Ancestral state reconstruction is used to infer ancestral areas and ancestral plant partners of the family. Results The Palaeotropics are unique in containing representatives of all major clades of Inocybaceae. Six of the seven major clades diversified initially during the Cretaceous, with subsequent radiations probably during the early Palaeogene. Vicariance patterns cannot be rejected that involve area relationships for Africa–Australia, Africa–India and southern South America–Australia. Northern and southern South America, Australia and New Zealand are primarily the recipients of immigrant taxa during the Palaeogene or later. Angiosperms were the earliest hosts of Inocybaceae. Transitions to conifers probably occurred no earlier than 65 Ma. Main conclusions The Inocybaceae initially diversified no later than the Cretaceous in Palaeotropical settings, in association with angiosperms. Diversification within major clades of the family accelerated during the Palaeogene in north and south temperate regions, whereas several relictual lineages persisted in the tropics. Both vicariance and dispersal patterns are detected. Species from Neotropical and south temperate regions are largely derived from immigrant ancestors from north temperate or Palaeotropical regions. Transitions to conifer hosts occurred later, probably during the Palaeogene.     

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.     

Circum-Antarctic continental distribution patterns in pteridophyte species

Four major austral continental distribution patterns are evident in pteridophytes. Twenty-two species are completely circum-Antarctic. Another 39 species are partially circum-Antarctic, occurring in Australasia (Australia and New Zealand) and Africa (including Madagascar) but not South America, while 29 are in Africa and South America but not Australasia, and 13 are in South America and Australasia but not Africa. Two hypotheses are considered as explanations for the patterns: continental drift following the breakup of Gondwana and long-distance dispersal. Fossil evidence indicates that the majority of pteridophyte families involved appeared after the southern continents had drifted apart, so long-distance dispersal is likely to explain the distribution of species in these families on now widely separated continents. For those families extant before the break-up, there is no indication in the fossil record that the species involved were present in Gondwana. Aspects of the ecology of the species that are partly or completely circum-Antarctic indicate that long-distance dispersal, rather than continental drift, is a likely explanation for the patterns.     

Frugivore gape size and the evolution of fruit size and shape in southern hemisphere floras

Abstract The present study uses differences among frugivore faunas of the southern hemisphere landmasses to test whether frugivore characteristics have influenced the evolution of fruit traits. Strong floristic similarities exist among southern landmasses; for example, 75% of New Zealand vascular plant genera also have species in Australia. However, plants in Australia and South America have evolved in the presence of a range of mammalian frugivores, whereas those in New Zealand, New Caledonia and the Pacific Islands have not. In addition, the avian frugivores in New Zealand and New Caledonia are generally smaller than those of Australia. If frugivore characteristics have influenced the evolution of fruit traits, predictable differences should exist between southern hemisphere fruits, particularly fruit size and shape. Fruit dimensions were measured for 77 New Zealand species and 31 Australian species in trans‐Tasman genera. New Zealand fruits became significantly more ellipsoid in shape with increasing size. This is consistent with frugivore gape size imposing a selective pressure on fruit ingestability. This result is not a product of phylogenetic correlates, as fruit length and width scaled isometrically for Australian species in genera shared with New Zealand. Within‐genus contrasts between New Zealand and Australian species in 20 trans‐Tasman genera showed that New Zealand species have significantly smaller fruits than their Australian counterparts. Within‐genus contrasts between New Zealand and South American species in nine genera gave the same result; New Zealand species had significantly smaller fruits than their South American counterparts. No difference was found in fruit size or shape between New Zealand and New Caledonia congeneric species from 12 genera. These results are consistent with the broad characteristics of the frugivore assemblage influencing the evolution of fruit size and shape in related species. The smaller‐sized New Zealand frugivore assemblage has apparently influenced the evolution of fruit size of colonizing taxa sometimes within a relatively short evolutionary timeframe.     

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.     

The first records of Stylodrilus heringianus (Oligochaeta: Lumbriculidae) from the Southern Hemisphere

Abstract

The oligochaete family Lumbriculidae is well represented in the Northern Hemisphere, but for the Southern Hemisphere only Lumbriculus variegatus (Müller) is recorded, from Africa, Australia, and New Zealand; no species are known from South America (Brinkhurst & Jamieson 1971). According to Brinkhurst (1971), L. variegatus may be a recent introduction to New Zealand, where it is now widely distributed in a range of inland waters.     

Cladogenesis as the result of long-distance rafting events in South Pacific topshells (Gastropoda, Trochidae)

We used DNA sequences of lecithotrophic monodontine topshells, belonging to the genera Diloma, Melagraphia, and Austrocochlea, to ascertain how this group became established over a large area of the South Pacific Ocean. The phylogeny of the topshells was estimated using portions of two mitochondrial genes (16S and cytochrome oxidase 1) and one nuclear gene (actin). A range of divergence rates was used to estimate the approximate timing of cladogenetic events within their phylogenetic tree. These estimates allow us to unambiguously reject vicariant explanations for several major divergence events and to infer several dispersal events across wide stretches of ocean. The first were two initial dispersal events from Australia (1) to an area between Samoa and Japan and (2) to New Zealand. Subsequently, at least one, and possibly two, recent eastward dispersals took place from New Zealand to Chile and the Juan Fernandez Islands, and one further dispersal occurred from somewhere in the tropical Pacific to Samoa. Moreover, owing to the short-lived nature of the topshell larvae, transoceanic larval dispersal is unlikely. The apparent paradox of a short larval phase and broad geographic range suggests that dispersal most probably occurred by rafting of adults on a suitable platform such as macroalgae; indeed, naturally buoyant bull kelp is the natural habitat of the most geographically widespread species in this group. Our molecular phylogenies imply that, despite of being an unlikely event, adult rafting in ocean currents has occurred on several occasions throughout the evolutionary history of topshells, resulting in their wide present-day distribution.     

Molecular evidence for long distance dispersal across the Southern Hemisphere in the Ganoderma applanatum-australe species complex (Basidiomycota)

We examined phylogeographic relationships in the cosmopolitan polypore fungus Ganoderma applanatum and allies, and conservatively infer a possible age of origin for these fungi. Results indicate that it is very unlikely that members of this species complex diversified before the break-up of Gondwana from Laurasia ca 120 M years ago, and also before the final separation of the Gondwanan landmasses from each other that was achieved about 66 M years ago. An earliest possible age of origin of 30 M years was estimated from nucleotide substitution rates in the 18S rDNA gene. Phylogenetic reconstruction of a worldwide sampling of ITS rDNA sequences reveals at least eight distinct clades that are strongly correlated with the geographic origin of the strains, and also correspond to mating groups. These include one Southern Hemisphere clade, one Southern Hemisphere–Eastern Asia clade, two temperate Northern Hemisphere clades, three Asian clades, and one neotropical clade. Geographically distant collections from the Southern Hemisphere shared identical ITS haplotypes, and an ITS recombinant was noted. Nested clade analysis of a parsimony network among isolates of the Southern Hemisphere clade indicated restricted gene flow with isolation-by-distance among the New Zealand, Australia–Tasmania, Chile–Argentine, and South Africa populations, suggesting episodic events of long-distance dispersal within the Southern Hemisphere. This study indicates that dispersal bias plays a more important role than generally admitted to explain the Southern Hemisphere distribution of many taxa, at least for saprobic fungi.     

Phylogeny and biogeography of Caltha (Ranunculaceae) based on chloroplast and nuclear DNA sequences

The genus Caltha (Ranunculaceae) consists of 10 species of low-growing, perennial herbs distributed throughout the moist temperate and cold regions of both the Northern and Southern Hemispheres. Traditionally, the species have been divided into two sections: section Psychrophila in the Southern Hemisphere with diplophyllous leaves and section Caltha in the Northern Hemisphere with leaves lacking inflexed appendages. This study uses chloroplast and nuclear DNA sequences to determine the relationships among the 10 species, test the monophyly of sections Psychrophila and Caltha, trace the evolutionary history of diplophylly, and explore biogeographical hypotheses for the genus. Analysis of these data resulted in a well-resolved and well-supported phylogeny. Section Psychrophila (C. sagittata, C. appendiculata, C. dionaeifolia, C. obtusa, C. introloba, and C. novae-zelandiae) was resolved as monophyletic, indicating a single origin of diplophylly. The species of section Caltha (C. natans, C. scaposa, C. palustris, and C. leptosepala) formed a paraphyletic grade. The resulting phylogeny strongly supports a Northern Hemisphere origin for Caltha, followed by dispersal to the Southern Hemisphere (Gondwanaland). A vicariance model is invoked to explain present-day distributions in South America, Australia, and New Zealand.     

Biogeography and evolution of the Galapagos: integration of the biological and geological evidence

Biogeographic tracks are mapped for Galapagos endemics representing 25 plant and animal taxa and including organisms with good and poor means of dispersal. These patterns confirm standard biogeographic tracks linking Galapagos with Central America, western North and South America, the Caribbean, Asia and Australasia. Discovery of the Galapagos Gore in the 1970s corroborates the biogeographic prediction for a major tectonic centre associated with the Galapagos. The biogeographic model developed by Croizat in 1958 of Galapagos colonization involving an ancestral biota inhabiting eastern Pacific geosynclinal forelands is congruent with plate tectonic models supporting a Pacific island arc origin for western American terranes. American relatives of Galapagos endemics may have originated within an eastern Pacific paleogeography rather than representing centres of origin for dispersal to the Galapagos. Galapagos colonization by an eastern Pacific biota between late Cretaceous and mid-Tertiary has significant implications for understanding the tempo and mode for both the origins of island biota and general models of evolutionary differentiation. Popular assertions that overwater dispersal represents the only viable origin for the entire Galapagos biota is no longer biogeographically or geologically tenable.