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.     

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.     

West Wind Drift revisited: testing for directional dispersal in the Southern Hemisphere using event-based tree fitting

Aim Recent studies suggest that if constrained by prevailing wind or ocean currents dispersal may produce predictable, repeated distribution patterns. Dispersal mediated by the West Wind Drift (WWD) and Antarctic Circumpolar Current (AAC) has often been invoked to explain the floristic similarities of Australia, South America and New Zealand. If these systems have been important dispersal vectors then eastward dispersal – from Australia to New Zealand and the western Pacific to South America – is expected to predominate. We investigate whether phylogenies for Southern Hemisphere plant groups provide evidence of historical dispersal asymmetry and more specifically whether inferred asymmetries are consistent with the direction of the WWD/AAC. Location Southern Hemisphere. Methods We assembled a data set of 23 published phylogenies for plant groups that occur in New Zealand, Australia and/or South America. We used parsimony‐based tree fitting to infer the number and direction of dispersals within each group. Observed dispersal asymmetries were tested for significance against a distribution of expected values. Results Our analyses suggest that dispersal has played a major role in establishing present distributions and that there are significant patterns of asymmetry in Southern Hemisphere dispersal. Consistent with the eastward direction of the WWD/ACC, dispersal from Australia to New Zealand was inferred significantly more often than in the reverse direction. No significant patterns of dispersal asymmetry were found between the western Pacific landmasses and South America. However, eastward dispersal was more frequently inferred between Australia and South America, while for New Zealand–South American events westward dispersal was more common. Main conclusions Our results suggest that eastward circumpolar currents have constrained the dispersal of plants between Australia and New Zealand. However, the WWD/ACC appear to have had less of an influence on dispersal between the western Pacific landmasses and South America. This observation may suggest that differences in dispersal mechanism are important – direct wind or water dispersal vs. stepping‐stone dispersal along the Antarctic coast. While our analyses provide useful preliminary insights into dispersal asymmetry in the Southern Hemisphere we will need larger data sets and additional methodological advances in order to test fully these dispersal patterns and infer processes from phylogenetic data.     

New Zealand phylogeography: evolution on a small continent

New Zealand has long been a conundrum to biogeographers, possessing as it does geophysical and biotic features characteristic of both an island and a continent. This schism is reflected in provocative debate among dispersalist, vicariance biogeographic and panbiogeographic schools. A strong history in biogeography has spawned many hypotheses, which have begun to be addressed by a flood of molecular analyses. The time is now ripe to synthesize these findings on a background of geological and ecological knowledge. It has become increasingly apparent that most of the biota of New Zealand has links with other southern lands (particularly Australia) that are much more recent than the breakup of Gondwana. A compilation of molecular phylogenetic analyses of ca 100 plant and animal groups reveals that only 10% of these are even plausibly of archaic origin dating to the vicariant splitting of Zealandia from Gondwana. Effects of lineage extinction and lack of good calibrations in many cases strongly suggest that the actual proportion is even lower, in keeping with extensive Oligocene inundation of Zealandia. A wide compilation of papers covering phylogeographic structuring of terrestrial, freshwater and marine species shows some patterns emerging. These include: east–west splits across the Southern Alps, east–west splits across North Island, north–south splits across South Island, star phylogenies of southern mountain isolates, spread from northern, central and southern areas of high endemism, and recent recolonization (postvolcanic and anthropogenic). Excepting the last of these, most of these patterns seem to date to late Pliocene, coinciding with the rapid uplift of the Southern Alps. The diversity of New Zealand geological processes (sinking, uplift, tilting, sea level change, erosion, volcanism, glaciation) has produced numerous patterns, making generalizations difficult. Many species maintain pre‐Pleistocene lineages, with phylogeographic structuring more similar to the Mediterranean region than northern Europe. This structure reflects the fact that glaciation was far from ubiquitous, despite the topography. Intriguingly, then, origins of the flora and fauna are island‐like, whereas phylogeographic structure often reflects continental geological processes.     

Relaxed Molecular Clock Provides Evidence for Long-Distance Dispersal of Nothofagus (Southern Beech)

Nothofagus (southern beech), with an 80-million-year-old fossil record, has become iconic as a plant genus whose ancient Gondwanan relationships reach back into the Cretaceous era. Closely associated with Wegener's theory of “Kontinentaldrift”, Nothofagus has been regarded as the “key genus in plant biogeography”. This paradigm has the New Zealand species as passengers on a Moa's Ark that rafted away from other landmasses following the breakup of Gondwana. An alternative explanation for the current transoceanic distribution of species seems almost inconceivable given that Nothofagus seeds are generally thought to be poorly suited for dispersal across large distances or oceans. Here we test the Moa's Ark hypothesis using relaxed molecular clock methods in the analysis of a 7.2-kb fragment of the chloroplast genome. Our analyses provide the first unequivocal molecular clock evidence that, whilst some Nothofagus transoceanic distributions are consistent with vicariance, trans-Tasman Sea distributions can only be explained by long-distance dispersal. Thus, our analyses support the interpretation of an absence of Lophozonia and Fuscospora pollen types in the New Zealand Cretaceous fossil record as evidence for Tertiary dispersals of Nothofagus to New Zealand. Our findings contradict those from recent cladistic analyses of biogeographic data that have concluded transoceanic Nothofagus distributions can only be explained by vicariance events and subsequent extinction. They indicate that the biogeographic history of Nothofagus is more complex than envisaged under opposing polarised views expressed in the ongoing controversy over the relevance of dispersal and vicariance for explaining plant biodiversity. They provide motivation and justification for developing more complex hypotheses that seek to explain the origins of Southern Hemisphere biota.     

Relaxed molecular clock provides evidence for long-distance dispersal of Nothofagus (southern beech)

Nothofagus (southern beech), with an 80-million-year-old fossil record, has become iconic as a plant genus whose ancient Gondwanan relationships reach back into the Cretaceous era. Closely associated with Wegener's theory of “Kontinentaldrift”, Nothofagus has been regarded as the “key genus in plant biogeography”. This paradigm has the New Zealand species as passengers on a Moa's Ark that rafted away from other landmasses following the breakup of Gondwana. An alternative explanation for the current transoceanic distribution of species seems almost inconceivable given that Nothofagus seeds are generally thought to be poorly suited for dispersal across large distances or oceans. Here we test the Moa's Ark hypothesis using relaxed molecular clock methods in the analysis of a 7.2-kb fragment of the chloroplast genome. Our analyses provide the first unequivocal molecular clock evidence that, whilst some Nothofagus transoceanic distributions are consistent with vicariance, trans-Tasman Sea distributions can only be explained by long-distance dispersal. Thus, our analyses support the interpretation of an absence of Lophozonia and Fuscospora pollen types in the New Zealand Cretaceous fossil record as evidence for Tertiary dispersals of Nothofagus to New Zealand. Our findings contradict those from recent cladistic analyses of biogeographic data that have concluded transoceanic Nothofagus distributions can only be explained by vicariance events and subsequent extinction. They indicate that the biogeographic history of Nothofagus is more complex than envisaged under opposing polarised views expressed in the ongoing controversy over the relevance of dispersal and vicariance for explaining plant biodiversity. They provide motivation and justification for developing more complex hypotheses that seek to explain the origins of Southern Hemisphere biota.     

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.     

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.     

Falkland Islands biogeography: converging trajectories in the South Atlantic Ocean

Aim This paper describes the biogeographical setting of the Falkland Islands, in the context of the relationships of the islands’ biota to other sub‐Antarctic/cold temperate lands. Location The analysis focuses primarily on the Falklands biota, and explores its relationships to those of Patagonian South America and South Africa, other southern lands and the islands of the sub‐Antarctic Pacific, Indian and Atlantic Oceans. Methods The study derives largely from literature sources on the biota and geological history of the Falkland Islands. Results The animals and plants known from the Falkland Islands exhibit strong affinities with those of Patagonian South America, and especially Tierra del Fuego; additional affinities are with various remote islands of the sub‐Antarctic, as well as New Zealand and to a lesser extent Australia; often these are shared with Patagonia. While the biotic affinities might be interpreted, by some, as indicating a former Gondwanan/South American geological connection of the Falklands, geological evidence points to the Falklands formerly having a land connection to south‐eastern South Africa. Only faint hints of a South African biotic connection remain. The historical biotic and geological connections of the Falklands thus conflict. Moreover, the Falklands biota is so strongly Patagonian that derivation of that biota is best seen as resulting from dispersal, much of it probably recent. This dispersal biota appears to have replaced, and perhaps displaced, the South African biota present on the islands as they detached from South Africa and drifted across the south Atlantic Ocean, as it opened up as South America and Africa drifted apart.     

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.     

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.     

New Zealand's pteridophyte flora—Plants of ancient lineage but recent arrival?

A hypothesis is presented that most pteridophytes arrived in New Zealand relatively recently, by long-distance dispersal. The flora comprises 194 native species, of which 89 (46%) are endemic and 105 (54%) are widespread. Of the latter, 90% are shared with temperate Australasia, 53% with tropical regions, 14% with temperate southern Africa and 13% with the circum-Antarctic islands and South America. New Zealand has undergone such dramatic changes in location, land area, and topography since initial separation from Gondwana 85 Ma that it seems improbable that the 95 species shared with temperate Australasia could have remained conspecific throughout that time. Modern fossil and molecular evidence strongly suggest that many families of ferns had not even evolved prior to separation, and palynological evidence from New Zealand indicates that 78% of pteridophyte genera first appeared there only after separation from Gondwana. Present-day distributions in New Zealand suggest that ferns have greater dispersal potential than flowering plants, and that pteridophyte distributions are more heavily influenced by temperature, rainfall, and geothermal activity than by geological history. Most endemic pteridophyte species have a predominantly southern distribution pattern and are characteristic of cool, lowland to montane forest. Pteridophytes in the northern part of New Zealand show a lower level of endemism than elsewhere and tend to be widespread species that have arrived from temperate Australasian and tropical regions. There is also evidence that at least some pteridophytes have migrated from New Zealand to Australia. It is suggested that the hypothesis of long-distance dispersal of pteridophytes across the Tasman Sea could be tested by molecular techniques.     

Molecular dating of the 'Gondwanan' plant family Proteaceae is only partially congruent with the timing of the break-up of Gondwana

Aim  The flowering plant family Proteaceae is putatively of Gondwanan age, with modern and fossil lineages found on all southern continents. Here we test whether the present distribution of Proteaceae can be explained by vicariance caused by the break-up of Gondwana.
Location  Africa, especially southern Africa, Australia, New Zealand, South America, New Caledonia, New Guinea, Southeast Asia, Sulawesi, Tasmania.
Methods  We obtained chloroplast DNA sequence data from the rbc L gene, the rbc L- atp B spacer, and the atp B gene from leaf samples of forty-five genera collected from the field and from living collections. We analysed these data using Bayesian phylogenetic and molecular dating methods, with five carefully selected fossil calibration points to obtain age estimates for the nodes within the family.
Results  Four of eight trans-continental disjunctions of sister groups within our sample of the Proteaceae post-date the break-up of Gondwana. These involve independent lineages, two with an Africa-Australia disjunction, one with an Africa–South America disjunction, and one with a New Zealand–Australasia disjunction. The date of the radiation of the bird-pollinated Embothriinae corresponds approximately to the hypothesized date of origin of nectar-feeding birds in Australia.
Main conclusions  The findings suggest that disjunct distributions in Proteaceae result from both Gondwanan vicariance and transoceanic dispersal. Our results imply that ancestors of some taxa dispersed across oceans rather than rafting with Gondwanan fragments as previously thought. This finding agrees with other studies of Gondwanan plants in dating the divergence of Australian, New Zealand and New Caledonian taxa in the Eocene, consistent with the existence of a shared, ancestral Eocene flora but contrary to a vicariance scenario based on accepted geological knowledge.     

Timing transantarctic disjunctions in the Atherospermataceae (Laurales): evidence from coding and noncoding chloroplast sequences

Previous studies of the small Southern Hemisphere family Atherospermataceae have drawn contradictory conclusions regarding the number of transantarctic disjunctions and role of transoceanic dispersal in its evolution. Clarification of intergeneric relationships is critical to resolving (1) whether the two Chilean species, Laurelia sempervirens and Laureliopsis philippiana, are related to different Austral-Pacific species, implying two transantarctic disjunctions as suggested by morphology; (2) where the group is likely to have originated; and (3) whether observed disjunctions reflect the breakup of Gondwana. We analyzed chloroplast DNA sequences from six regions (the rbcL gene, the rpl16 intron, and the trnL-trnF, trnT-trnL, psbA-trnH, and atpB-rbcL spacer regions; for all six regions, 4,372 bp) for all genera and most species of Atherospermataceae, using parsimony and maximum likelihood (ML). The family's sister group, the Chilean endemic Gomortega nitida (Gomortegaceae), was used to root the tree. Parsimony and ML yielded identical single best trees that contain three well-supported clades (> or = 75% bootstrap): Daphnandra and Doryphora from south-eastern Australia; Atherosperma and Nemuaron from Australia-Tasmania and New Caledonia, respectively; and Laurelia novac-zelandiac and Laureliopsis philippiana from New Zealand and Chile, respectively. The second Chilean species, Laurelia sempervirens, is sister to this last clade. Likelihood ratio testing did not reject the molecular clock assumption for the rbcL data, which can therefore be used for divergence time estimates. The atherosperm fossil record, which goes back to the Upper Cretaceous, includes pollen, wood, and leaf fossils from Europe, Africa, South America, Antarctica, New Zealand, and Tasmania. Calibration of rbcL substitution rates with the fossils suggests an initial diversification of the family at 100-140 million years ago (MYA), probably in West Gondwana, early entry into Antarctica, and long-distance dispersal to New Zealand and New Caledonia at 50-30 MYA by the ancestors of L. novae-zelandiae and Nemuaron.     

Phylogeny and historical biogeography of Gondwanan moss-bugs (Insecta: Hemiptera: Coleorrhyncha: Peloridiidae)

The moss bugs of the Peloridiidae, a small group of cryptic and mostly flightless insects, is the only living family in Coleorrhyncha (Insecta: Hemiptera). Today 37 species in 17 genera are known from eastern Australia, New Zealand, New Caledonia and Patagonia, and the peloridiids are thereby a group with a classical southern Gondwanan distribution. To explicitly test whether the present-day distribution of the Peloridiidae actually results from the sequential breakup of southern Gondwana, we provide the first total-evidence phylogenetic study based on morphological and molecular characters sampled from about 75% of recognized species representing 13 genera. The results largely confirm the established morphological phylogenetic context except that South American Peloridium hammoniorum constitutes the sister group to the remaining peloridiids. A timescale analysis indicates that the Peloridiidae began to diversify in the land mass that is today's Patagonia in the late Jurassic (153 Ma, 95% highest posterior density: 78–231 Ma), and that splitting into the three extant well-supported biogeographical clades (i.e. Australia, Patagonia and New Zealand/New Caledonia) is consistent with the sequential breakup of southern Gondwana in the late Cretaceous, indicating that the current transoceanic disjunct distributions of the Peloridiidae are best explained by a Gondwanan vicariance hypothesis.     

East meets west: biogeology of the Campbell Plateau

The New Zealand Subantarctic Islands, emergent remnants of the Campbell Plateau, were given World Heritage status in 1998 in recognition of their importance to global biodiversity. We describe the flora and fauna of these islands and discuss the results of recent phylogenetic analyses. Part of the New Zealand Subantarctic biota appears to be relictual and to be derived from west Gondwana. The relictual element is characterized by genera endemic to the Campbell Plateau that show relationships with taxa of the southern South Island, New Zealand, southern South America, and the north Pacific. In contrast, a younger, east Gondwanan element is composed of species that are either taxonomically identical to widespread mainland species, or endemic species with close New Zealand relatives. Area cladograms support the inclusion of the southern South Island, New Zealand and Macquarie Island (although this is separate geologically) as parts of the Campbell Plateau, but suggest the Chatham Rise and Torlesse terranes of the eastern South Island, New Zealand were originally parts of east Gondwana. East and west Antarctica acted as independent plates during the breakup of Gondwana, and were separated by oceanic crust until a compressive phase sutured them along the trace of the trans‐Antarctic mountains during the early Tertiary. The Campbell Plateau microcontinent was connected to west Antarctica until its separation at 80 Mya, contemporaneous with the separation of the southern portion of the Melanesian rift from east Gondwana. Presently the Campbell Plateau is joined to the Melanesian Rift along the Alpine Fault. Cenozoic plate tectonic reconstructions place the Campbell Plateau adjacent to the Melanesian Rift throughout the rift–drift phase, relative motion being confined to strike–slip movement over the last 20 Myr. Our synthesis of phylogenetic and plate tectonic evidence suggests that the Alpine Fault is the most recent development of a much older extensional rift/basin boundary originally separating west and east Gondwana. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 95–115.     

A CLADISTIC ANALYSIS OF THE EVOLUTION AND BIOGEOGRAPHY OF DURVILLAEA (PHAEOPHYTA)1

The genus Durvillaea currently has four recognized species found along many exposed, rocky coastlines of the temperate to sub-Antarctic regions in the Southern Hemisphere. We propose that the current species distributions are related primarily to vicariance events and subsequent speciation associated with the breakup of Gondwana between 40 and 100 Ma. From an ancestral species, a stipitate species developed in the Tasman basin, with separation and speciation resulting in the D. potatorum/ D. willana complex in southeastern Australia and New Zealand. A second line of evolution led to D. chathamensis and D. antarctica characterized by a honeycombed medulla. The extensive distribution of D. antarctica throughout the Southern Hemisphere is related to both vicariance and dispersal events. The status of D. chathamensis as a species distinct from D. antarctica is questioned. The affinities of an as yet undescribed taxon from the Antipodes Islands are thought to be with the D. potatorum complex but require further study before they can be defined more precisely .     

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.     

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.