Molecular phylogeny ofNothofagus (Nothofagaceae) based on theatpB-rbcL intergenic spacer of the chloroplast DNA

The genusNothofagus is distributed in the Southern Hemisphere from South America to Oceania, and its distribution has been assumed to be formed by continental drift by means of Gondwana break-up during the Mesozoic era. The phylogeny of the genus was elucidated by the sequences ofatpB-rbcL intergenic spacer of cpDNA for the better understanding of its evolution and biogeography. The phylogeny ofNothofagus corresponded completely to the pollen morphology which recognizes four pollen types in extant species, and agrees well with the taxonomic system of Hill and Read (1991) although there, the subgenusNothofagus showed in unresolved polytomy. The topology of the phylogenetic tree reveals that subgenusLophozonia was derived first, and thenFuscospora, Nothofagus andBrassospora. Species from South America and New Zealand were assigned to each cluster according to their pollen morphology. Therefore, diversification ofNothofagus should have already proceeded at the subgenus level before the completion of Gondwana break-up Tropical species distributed in New Guinea and New Caledonia whose evolutionary history has been controversial were revealed to be a derived group. All five New Caledonian species formed a monophyletic group with very few sequence divergences in the intergenic spacer of cpDNA, thus showing rapid adaptive radiation in the island. Evolutionary trends of several morphological traits ofNothofagus are discussed. The evolution of valve number of cupules, number of nuts per cupule, and habit of leaf-fall (evergreen or deciduous) which are diversified in the genus, were revealed as having occurred several times as the result of convergence.     

Nothofagus subgenus Brassospora (Nothofagaceae) leaf fossils from New Zealand: a link to Australia and New Guinea?

N othofagus palustris sp. nov. is the first record of well‐preserved leaves of Nothofagus subgenus Brassospora in New Zealand, and is described from an Oligo–Miocene leaf bed in the Gore Lignite Measures of the South Island. Nothofagus palustris is represented by relatively small and variably toothed leaves with cuticular remains that possess all the characteristic features of the subgenus, including the presence of variably sized stomata that are randomly arranged within areoles, hydathodes along the major veins and ‘bulging cells’ within the areoles on the adaxial side. Phylogenetic assessment shows that the leaves are similar to those of Australian Oligocene and Miocene species and may belong to the same clade of Brassospora. Most notably, these species share the derived feature of abundant leaf wax, a feature that is now only well developed in two New Guinean species. This and other evidence allows the possibility that the ancestor of N. palustris reached New Zealand from Australia. However, it is improbable that N. palustris or a similar species was the common ancestor of the clade of Brassospora that is now confined to New Caledonia. Ecologically, N. palustris is unusual among extant and previously described macrofossil species of Brassospora in being found in a relatively open, swampy habitat. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 503–515.     

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

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

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.     

Late Cenozoic diversification of the austral genus Lagenophora (Astereae,Asteraceae)

Lagenophora (Astereae, Asteraceae) has 14 species in New Zealand, Australia, Asia, southern South America, Gough Island and Tristan da Cunha. Phylogenetic relationships in Lagenophora were inferred using nuclear and plastid DNA regions. Reconstruction of spatio‐temporal evolution was estimated using parsimony, Bayesian inference and likelihood methods, a Bayesian relaxed molecular clock and ancestral area and habitat reconstructions. Our results support a narrow taxonomic concept of Lagenophora including only a core group of species with one clade diversifying in New Zealand and another in South America. The split between the New Zealand and South American Lagenophora dates from 11.2 Mya [6.1–17.4 95% highest posterior density (HPD)]. The inferred ancestral habitats were openings in beech forest and subalpine tussockland. The biogeographical analyses infer a complex ancestral area for Lagenophora involving New Zealand and southern South America. Thus, the estimated divergence times and biogeographical reconstructions provide circumstantial evidence that Antarctica may have served as a corridor for migration until the expansion of the continental ice during the late Cenozoic. The extant distribution of Lagenophora reflects a complex history that could also have involved direct long‐distance dispersal across southern oceans. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 78–95.     

Nothofagus (Fagaceae) and its invertebrate fauna—an overview and preliminary synthesis

The invertebrates directly associated with the southern hemisphere tree genus Nothofagus are described and discussed in terms of their organization into guilds and attributes of their hostplants. Using literature records, comparisons are made of the guild composition between host species and provenances; a full invertebrate-hostplant list is appended. Sap-sucking Homoptera are particularly diverse and usually host-species specific whereas defoliating insects seem less prevalent and less specific. Few seed-eating insects are reported. Some provenances and hostplants exhibit local radiations in certain taxa, such as Eriococcidae in New Zealand, Notophorina psyllids in South America and scolytid beetles on N. dombeyi. Environmental stresses such as drought, disturbance and landslips, can induce outbreaks of certain scale insects and borers, affecting the dynamics of the forest. Generally, however, the influence of invertebrates appears benign, but occasional large-scale defoliation is observed on N. solandri var. cliffortioides, N. moorei, N. antarctica and N. pumilio. Collectively this fauna has two elements: a geographically recurrent Gondwanic element, usually monophagous, and a provenance-specific element (often polyphagous) locally recruited following vicariance of the host genus. The fauna of Nothofagus can be in stark contrast to that of adjacent forest types. In Australia, the insect profile of Nothofagus has more in common with fagaceous trees on other continents than the more recent Eucalyptus forests which surround it. There is almost no evidence of fauna transfer from Nothofagus to Eucalyptus. Relative to New Zealand, the Australian Nothofagus fauna is depauperate and may reflect truncation in Pleistocene glaciations. Typically low insect populations on these trees may partly explain the limited range of insectivorous birds and spiders observed in Australia. Much more data on the phytophagous insects of this biome is needed for comparative purposes. The fauna of the montane tropical subgenus Brassospora pollen group is almost unknown but could hold the key to the origin of fagaceous insect communities more generally. The conservation value of remaining Nothofagus forests is enhanced by recognition of their co-adapted fauna, some of which are arranged in stable functional groups likely to give valuable insight into invertebrate-hostplant interactions originating in the Cretaceous. Elucidation of its role in the processes of these unique forests remains a fertile field for endeavour in the future.     

Nothofagus Biogeography Revisited with Special Emphasis on the Enigmatic Distribution of Subgenus Brassospora in New Caledonia

Dispersals versus vicariance events and the presence of subgenus Brassospora in New Caledonia are two riddles of Nothofagus biogeography, a genus also distributed in New Guinea, New Zealand, South America, Southeast Australia, and Tasmania. Within a cladistic framework using the software COMPONENT 2.0, we demonstrate that most parsimonious area cladograms (areagrams) sensu cladistic biogeography need not always be the most plausible explanation nor reflect alternative geological hypotheses. The most parsimonious Nothofagus history sensu historical biogeography is reconstructed where a minimum of dispersed taxa is hypothesized and vicariance events are identified. A fully resolved well-established Nothofagus phylogeny was reconciled with three geological hypotheses (geograms) of East Gondwana break-up: (a) the conventional view, (b) an Australian—New Caledonian relationship, and (c) a biotic interchange between New Guinea and New Caledonia. Fossils determined to subgenus were optimized to the predicted lineages in the reconciled tree. Due to extensive extinctions, a maximum of three vicariance events are inferred, all being basal in the subgenera, an indication of subgeneric diversification prior to the break-up of Gondwana. Two taxa, N. gunnii and N. menziesii, are hypothesized as being long-distance dispersed. The most parsimonious solution suggests a close relationship between New Guinea and New Caledonia, supporting a Brassospora colonization route, but this hypothesis fails to predict numerous extinct lineages observed in the fossil record and thus must be rejected. The traditional break-up sequence of Gondwana is not the most parsimonious solution, indicating one incongruent node, but causes no overall incongruence with the fossil record. Considering all parameters, the occurrence of Brassospora in New Caledonia is most parsimoniously explained as a single colonization event from New Zealand where the subgenus subsequently went extinct in the Pliocene.     

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.     

Systematics of Nothofagus (Nothofagaceae) based on rDNA spacer sequences (ITS): taxonomic congruence with morphology and plastid sequences

Phylogenetic relationships were examined within the southern beech family Nothofagaceae using 22 species representing the four currently recognized subgenera and related outgroups. Nuclear ribosomal DNA sequences encoding the 5.8s rRNA and two flanking internal transcribed spacers (ITS) provided 95 phylogenetically informative nucleotide sites from a single alignment of ~588 bases per species. Parsimony analysis of this variation produced two equally parsimonious trees supporting four monophyletic groups, which correspond to groups designated by pollen type. These topologies were compared to trees from reanalyses of previously reported rbcL sequences and a modified morphological data set. Results from parsimony analysis of the three data sets were highly congruent, with topological differences restricted to the placement of a few terminal taxa. Combined analysis of molecular and morphological data produced six equally parsimonious trees. The consensus of these trees suggests two basal clades within Nothofagus. Within the larger of the two clades, tropical Nothofagus (subgenus Brassospora) of New Guinea and New Caledonia are strongly supported as sister to cool-temperate species of South America (subgenus Nothofagus). Most of the morphological apomorphies of the cupule, fruit, and pollen of Nothofagus are distributed within this larger clade. An area cladogram based on the consensus of combined data supports three trans-Antarctic relationships, two within pollen groups and one between pollen groups. Fossil data support continuous ancestral distributions for all four pollen groups prior to continental drift; therefore, vicariance adequately explains two of these disjunctions. Extinction of trans-Antarctic sister taxa within formerly widespread pollen groups explains the third disjunction; this results in a biogeographic pattern indicative of phylogenetic relationship not vicariance. For the biogeographically informative vicariant clades, area relationships based on total evidence support the recently advanced hypothesis that New Zealand and Australia share a unique common ancestry. Contrary to previous thought, the distribution of extant Nothofagus is informative on the area relationships of the Southern Hemisphere, once precise phylogenetic relationships are placed in the context of fossil data.     

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.     

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.     

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.     

Pollen morphology and fossil record of the feathery mistletoe family Misodendraceae

Misodendraceae is a small family of mistletoes in the order Santalales. Its distribution is restricted to the southern South American temperate forests. The family comprises the sole genus Misodendrum with eight species of hemiparasitic shrubs, mainly parasitising the southern beech Nothofagus. This contribution presents palynological evidence from seven species, using light microscopy and scanning electron microscopy. Pollen grains are consistently small, periporate and echinate, although differences in the length of echini and number and size of pores were noted. Pollen features can be used to distinguish groups of species and, in some cases, individual species. Cluster analysis of pollen characters differentiates two main groups: one includes M. brachystachyum, M. oblongifolium and M. quadriflorum; and the other includes M. gayanum, M. linearifolium, M. punctulatum and M. angulatum. Palynological results are compared with previous systematic studies of the family. The South American fossil pollen record is summarised and characters of the fossil pollen are analysed using UPGMA to test the relationships between extant and fossil species. Miocene pollen resulted similar to species of subgenus Angelopogon while Eocene pollen is disimilar to extant species of Misodendraceae.     

Not so ancient: the extant crown group of Nothofagus represents a post-Gondwanan radiation

This study uses a molecular-dating approach to test hypotheses about the biogeography of Nothofagus. The molecular modelling suggests that the present-day subgenera and species date from a radiation that most likely commenced between 55 and 40 Myr ago. This rules out the possibility of a reconciled all-vicariance hypothesis for the biogeography of extant Nothofagus. However, the molecular dates for divergences between Australasian and South American taxa are consistent with the rifting of Australia and South America from Antarctica. The molecular dates further suggest a dispersal of subgenera Lophozonia and Fuscospora between Australia and New Zealand after the onset of the Antarctic Circumpolar Current and west wind drift. It appears likely that the New Caledonian lineage of subgenus Brassospora diverged from the New Guinean lineage elsewhere, prior to colonizing New Caledonia.The molecular approach strongly supports fossil-based estimates that Nothofagus diverged from the rest of Fagales more than 84 Myr ago. However, the mid-Cenozoic estimate for the diversification of the four extant subgenera conflicts with the palynological interpretation because pollen fossils, attributed to all four extant subgenera, were widespread across the Weddellian province of Gondwana about 71 Myr ago. The discrepancy between the pollen and molecular dates exists even when confidence intervals from several sources of error are taken into account. In contrast, the molecular age estimates are consistent with macrofossil dates. The incongruence between pollen fossils and molecular dates could be resolved if the early pollen types represent extinct lineages, with similar types later evolving independently in the extant lineages.     

Small Celastraceae and Polygonaceae twigs from the Upper Cenozoic (Ituzaingó Formation) of the La Plata Basin,Argentina

Two new wood types from the Late Cenozoic of the Ituzaingó Formation, La Plata Basin, Northeast Argentina add to our knowledge of South American Cenozoic plants. The materials were preserved by siliceous cellular permineralization, and they were prepared for microscopic examination by surface polishing and in thin sections. The anatomy of these new species was described. The relationship and comparison with the nearest living relatives (NLRs) are discussed. Maytenoxylon perforatum Franco gen. and sp. nov. is described as the first fossil wood referable to Celastraceae from South America. This new fossil species is related to extant Maytenus Molina. The other fossil twig, Ruprechtioxylon breae Franco sp. nov., has features of the Polygonaceae family and particularly resembles the extant specie Ruprechtia laxiflora Meisn. The occurrence of these fossil woods in south-eastern South America suggests that a relatively warm and dry to seasonally dry climate prevailed over this region of Gondwana during the Upper Cenozoic. It also provides new evidence for the hypothesis of the more wide distribution of Seasonally Dry Tropical Forest (SDTF) during the Upper Cenozoic.     

Paleofloristic assemblage from the Paleogene Río Guillermo Formation,Argentina: preliminary results of phylogenetic relationships of Nothofagus in South America

Hünicken's paleobotanical collection is one of the most important in South America because it was the first one to be related to a documented stratigraphic profile. This floristic assemblage (Oligocene) recovered from Río Guillermo Formation, at Estancia Tres Marías, Río Guillermo Valley, Santa Cruz, Argentina, is described by the first time in this paper. Five species of genus Nothofagus (N. subferruginea, N. serrulata, N. crenulata, N. elongata and N. variabilis) together with Myrcia bagualense are components of the assemblage. The presence of Acaena brandmayri is confirmed for the first time for the area. Paleoenviromental and paleoclimate implications are also discussed. Characteristic species from a temperate-cold climate such as the genus Nothofagus are present in the material studied. A phylogenetic study of the fossil species herein described and the extant South American species of Nothofagus is presented. The preliminary results support the hypothesis that fossil species of Nothogafus are closely related to the modern species.     

Molecular evidence for long-distance dispersal in the New Zealand pteridophyte flora

Aim To examine the relative importance of long‐distance dispersal in shaping the New Zealand pteridophyte (ferns and lycophytes) flora and its relationships with other floras, with the null hypothesis that the extant New Zealand pteridophyte flora has been isolated since New Zealand’s separation from Gondwana. Location New Zealand. Methods rbcL DNA sequences were assembled for 31 New Zealand pteridophyte genera, with each genus represented by one New Zealand species and the most closely related non‐New Zealand species for which data were available. Maximum‐likelihood, maximum‐parsimony, and Bayesian analysis phylograms were constructed and used as input for r 8s molecular dating, along with 23 fossil calibrations. Divergence estimates less than conservatively recent ages for New Zealand’s geological isolation, namely H_o > 30 Ma for pairs involving New Caledonian and Norfolk Island species and H_o > 55 Ma for all others, were taken as rejection of the null hypothesis. Results The null hypothesis was rejected for all pairs except, under some parameter conditions, for those involving the New Zealand species Cardiomanes reniforme, Lindsaea trichomanoides, Loxsoma cunninghamii, Lygodium articulatum, Marattia salicina, and Pteris comans. However, the Lindsaea and Pteris results probably reflect the absence in the analyses of closely related non‐New Zealand samples, while the Marattia divergence was highly contingent on which fossil calibrations were used. Main conclusions Rejection of the null hypothesis for the majority of pairs implies that the extant New Zealand lineage has undergone long‐distance dispersal either into or out of New Zealand. The notion of a long isolation since geological separation can, therefore, be dismissed for much of New Zealand’s pteridophyte flora. The analyses do not identify the direction of the long‐distance dispersal, and these New Zealand lineages could have had vicariant origins with subsequent long‐distance emigration. However, the alternative that many extant New Zealand pteridophyte lineages only arrived by long‐distance immigration after geological isolation seems likely.     

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 phylogenetic placement and biogeographical origins of the New Zealand stick insects (Phasmatodea)

The Lanceocercata are a clade of stick insects (Phasmatodea) that have undergone an impressive evolutionary radiation in Australia, New Caledonia, the Mascarene Islands and areas of the Pacific. Previous research showed that this clade also contained at least two of the nine New Zealand stick insect genera. We have constructed a phylogeny of the Lanceocercata using 2277 bp of mitochondrial and nuclear DNA sequence data to determine whether all nine New Zealand genera are indeed Lanceocercata and whether the New Zealand fauna is monophyletic. DNA sequence data were obtained from mitochondrial cytochrome oxidase subunits I and II and the nuclear large subunit ribosomal RNA and histone subunit 3. These data were subjected to Bayesian phylogenetic inference under a partitioned model and maximum parsimony. The resulting trees show that all the New Zealand genera are nested within a large New Caledonian radiation. The New Zealand genera do not form a monophyletic group, with the genus Spinotectarchus Salmon forming an independent lineage from the remaining eight genera. We analysed Lanceocercata apomorphies to confirm the molecular placement of the New Zealand genera and to identify characters that confirm the polyphyly of the fauna. Molecular dating analyses under a relaxed clock coupled with a Bayesian extension to dispersal‐vicariance analysis was used to reconstruct the biogeographical history for the Lanceocercata. These analyses show that Lanceocercata and their sister group, the Stephanacridini, probably diverged from their South American relatives, the Cladomorphinae, as a result of the separation of Australia, Antarctica and South America. The radiation of the New Caledonian and New Zealand clade began 41.06 million years ago (mya, 29.05–55.40 mya), which corresponds to a period of uplift in New Caledonia. The main New Zealand lineage and Spinotectarchus split from their New Caledonian sister groups 33.72 (23.9–45.62 mya) and 29.9 mya (19.79–41.16 mya) and began to radiate during the late Oligocene and early Miocene, probably in response to a reduction in land area and subsequent uplift in the late Oligocene and early Miocene. We discuss briefly shared host plant patterns between New Zealand and New Caledonia. Because Acrophylla sensu Brock & Hasenpusch is polyphyletic, we have removed Vetilia Stål from synonymy with Acrophylla Gray.     

The endemic plant families and the palms of New Caledonia: a biogeographical analysis

This paper provides a panbiogeographical analysis of the endemic plant families and the palms of New Caledonia. There are three endemic plant families in New Caledonia and several genera that were previously recognized as endemic families. Of these taxa, some are sister to widespread Northern Hemisphere or global groups (Canacomyrica, Austrotaxus, Amborella). The others belong to trans‐Indian Ocean groups (Strasburgeria), trans‐tropical Pacific groups (Oncotheca) or Tasman Sea/Coral Sea groups (Phelline, Paracryphia) that are sister to widespread Northern Hemisphere or global groups. In palms, the four clades show allopatric regional connections in, respectively: (1) western Indonesia, Malaysia and Thailand; (2) Vanuatu/Fiji and the southern Ryukyu Islands near Taiwan; (3) the western Tasman/Coral Sea (eastern Australia, New Guinea and the Solomon Islands); and (4) the eastern Tasman/Coral Sea (Lord Howe and Norfolk Islands, New Zealand, Vanuatu, Fiji and the Solomon Islands). The four clades thus belong to different centres of endemism that overlap in New Caledonia. The patterns are attributed not to chance dispersal and adaptive radiation but to the different histories of the eight terranes that fused to produce modern New Caledonia. Trans‐tropical Pacific connections can be related to the Cretaceous igneous plateaus that formed in the central Pacific and were carried, with plate movement, west to the Solomon Islands and New Zealand, and east to Colombia and the Caribbean.