Nothofagus Subgenus Research Articles

Two sides of the same coin? Transient hybridization in refugia and rapid postglacial ecological divergence ensure the evolutionary persistence of sister Nothofagus.

Glacial periods have been considered as inhospitable environments that consist of treeless vegetation at higher latitudes. The fossil record suggests many species survived the Last Glacial Maximum within refugia, usually at lower latitudes. However, phylogeographic studies have given support to the existence of previously unknown high-latitude refugia that were not detected in the fossil record. Here, we test the hypothesis that cold-tolerant trees of Patagonia survived cold periods in microclimatically favourable locales where hybridization occurred between sister taxa. To study local presence through glacial periods in multiple refugia, we used pollen records and genetic information (isozymes, microsatellites, and combined nuclear and chloroplast DNA sequences) of population pairs of Nothofagus antarctica and N. pumilio that belong to the ancient subgenus Nothofagus which can potentially hybridize in nature, along their entire latitudinal range in Patagonia. Studied species share the N. dombeyi type pollen, which was abundant at >20% in the northernmost latitudinal bands (35-43°S), even during the Last Glacial Maximum. Mid- and southern latitudinal records (44-55°S) yielded lower abundances of ~10% that increased after c. 15.0 cal. ka BP. Therefore, fossil pollen evidence suggests a long-lasting local presence of Nothofagus throughout glacial-interglacial cycles but mostly as small populations between 44°S and 51°S. We found species-specific and shared genetic variants, the latter of which attained relatively high frequencies, thus providing evidence of ancestral polymorphisms. Populations of each species were similarly diverse, suggesting survival throughout the latitudinal range. Estimates of coalescent divergence times were broadly synchronous across latitudes, suggesting that regional climates similarly affected populations and species that hybridized through climate cycles, fostering local persistence.

Plastomes of Nothofagus reflect a shared biogeographic history in Patagonia.

Plastomes are used in phylogenetic reconstructions because of their relatively conserved nature. Nonetheless, some limitations arise, particularly at lower taxonomic levels due to reduced interspecific polymorphisms and frequent hybridization events that result in unsolved phylogenies including polytomies and reticulate evolutionary patterns. Next-generation sequencing technologies allow access to genomic data and strongly supported phylogenies, yet biased topologies may be obtained due to insufficient taxon sampling. We analyze the hypothesis that intraspecific plastome diversity reflects biogeographic history and hybridization cycles among taxa. We generated 12 new plastome sequences covering distinct latitudinal locations of all species of subgenus Nothofagus from North Patagonia. Chloroplast genomes were assembled, annotated, and searched for simple sequence repeats (SSRs). Phylogenetic reconstructions included species and sampled locations. The six Nothofagus species analyzed were of similar size and structure; only Nothofagus obliqua of subgenus Lophozonia, used as an outgroup, presented slight differences in size. We detected a variable number of SSRs in distinct species and locations. Phylogenetic analyses of plastomes confirmed that subgenus Nothofagus organizes into two monophyletic clades each consisting of individuals of different species. We detected a geographic structure within subgenus Nothofagus and found evidence of local chloroplast sharing due to past hybridization, followed by adaptive introgression and ecological divergence. These contributions enrich the comprehension of transversal evolutionary mechanisms such as chloroplast capture and its implications for phylogenetic and phylogenomic analyses.

Palaeoecological evidence of pollen morphological changes: A climate change adaptation strategy?

Pollen records have been used to reconstruct changes in climate, based on the presence of taxa as proxies for environmental conditions. Pollen grains have morphological features that allow for the identification of different species, genera, and families of flowering plants. However, ecologically distinct species share similar pollen types that may make biogeographic and climate reconstructions problematic. Also, the responses of populations and species to climate changes adjusted through either plasticity or adaptation have been overlooked. Such adjustments are most probably the product of long-lasting in situ persistence in local refugia under favourable microclimates despite hostile regional climate. In southern South America three species of the dominant tree cover of the subgenus Nothofagus are present, two deciduous (N. antarctica and N. pumilio) and one evergreen (N. betuloides), identified in the pollen group Nothofagus dombeyi type. Here we present a pollen record from Punta Arenas (~53°S) that documents significant changes in pollen size that took place during the Late glacial and Early Holocene. The occurrence of smaller Nothofagus dombeyi type pollen grains during the Late glacial may indicate rapid local adaptation to colder and possibly wetter conditions. Our findings are supported by Ecological niche modelling which suggests limited availability of Nothofagus refugia during the Last Glacial Maximum. The local refugia may also have fostered interbreeding between congeneric species that led to unique morphological features that increased the fitness of the recipient pool. We postulate that such modifications were due to the potential transient hybridization between species that later diverged by backcrossing with each deciduous and evergreen taxa.

Biogeographically marginal: Source of evolutionary novelties and future potential

Towards distribution margins, populations are thought to be genetically impoverished due to isolation and genetic drift acting in often small border populations. Here we analyse the hypothesis that contrary to expectations, populations towards the dry Nothofagus pumilio range are relicts of a past distribution, and thus hold comparable levels of genetic diversity than central ones using conserved chloroplast DNA (cpDNA) sequences. We sampled fresh leaf tissue from a total of 70 natural populations along a precipitation gradient in Northern Patagonia, Argentina, from central (N = 33), i. e. humid, to marginal (N = 37), i. e. dry, locations which were analysed by cpDNA sequencing and phylogeographic methods. We also analysed the fate of populations under global scenarios by ecological niche models. The analysis of a total of 186 cpDNA sequences yielded nine different haplotypes, eight and six of which were present in marginal and central regions, respectively. We identified four new haplotypes for the species, three were exclusive to the marginal region and one was also new for the Nothofagus subgenus. Haplotypes were latitudinally structured as previously documented, yet those from marginal and central regions diverged synchronically at ~3.5 Ma. Coalescence analyses yielded larger effective population sizes for marginal populations, except for the northernmost group, and divergence between regions occurred from early to late Pleistocene. Neutrality tests developed to infer past population size changes showed population stability for marginal and central regions yet current range retraction. Future distributions by ecological niche models predict population decay in both central and marginal regions with a stronger effect in the latter. Our results show that marginal populations are as genetically diverse and demographically stable as central ones and their genetic makeup may assure their long-lasting persistence. Unique variants and genetically-based adaptive traits of the former may be used as germplasm sources for restoration under forecasted droughts.

Fossil Woods from the Middle Miocene (Río Correntoso Formation) of Patagonia (Northern Santa Cruz Province, Argentina)

We collected nine samples from the middle Miocene of Patagonia, in outcrops of the Rio Correntoso Formation (Rio Zeballos Group) in north Santa Cruz Province, Argentina. Fossil woods have variable preservation and some are poorly preserved. Six samples are silicified, and three are totally carbonized. Two taxonomic units were confidently described, Cupressinoxylon sp., a conifer with Cupressaceae or Podocarpaceae affinity and Nothofagoxylon scalariforme with Nothofagus subgenus Nothofagus affinity of the Nothofagaceae. Other samples are indeterminate angiosperms and conifers. Growth rings of two samples indicate seasonality and galleries with frass and coprolites found in an indeterminate conifer are similar to those made by coleoptera and acari. The assemblage taxonomic composition is similar to other Miocene wood floras of Patagonia with conifers and Nothofagaceae.

Hybrid identification in Nothofagus subgenus using high resolution melting with ITS and trnL approach.

The genus Nothofagus is the main component of southern South American temperate forests. The 40 Nothofagus species, evergreen and deciduous, and some natural hybrids are spread among Central and Southern Chile, Argentina, New Zealand, Australia, New Guinea and New Caledonia. Nothofagus nervosa, Nothofagus obliqua and Nothofagus dombeyi are potentially very important timber producers due to their high wood quality and relative fast growth; however, indiscriminate logging has degraded vast areas the Chilean forest causing a serious state of deterioration of their genetic resource. The South of Chile has a large area covered by secondary forests of Nothofagus dombeyi. These forests have a high diversity of species, large amount of biomass and high silvicultural potential. This work shows a case of hybrid identification in Nothofagus subgenus in different secondary forests of Chile, using high resolution melting. Unknown samples of Nothofagus subgenus are genetically distinguishable with the ITS region of Nothofagus antarctica, Nothofagus nitida and N. obliqua species. It was not possible to distinguish between unknown samples of Andean versus coastal origin. Melting curves with ITS approach of unknown material are genetically similar, positioned between N. dombeyi and N. antarctica and distant from N. nitida. The unknown samples are genetically very close to Nothofagus dombeyi. This suggests the presence of hybrid individuality between species (N. dombeyi × N. antarctica) with the possibility of introgression towards the gene pool of N. antarctica, producing the deciduous foliage that is both present. The trnL locus has no distinction between the N. dombeyi and N. antarctica species, since a similar melting curve is present and equal Tm (80.00 °C). The trnL locus cannot be genetically distinguished from one unknown sample of Nothofagus to another, as highlighted in this study.

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

Nothofagus 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.

Characterisation of seven polymorphic microsatellites for Nothofagus subgenus Fuscospora from New Zealand

We generated a genomic DNA library for Nothofagus solandri using 454 sequencing. Sequences from this library were screened for simple sequence repeat (microsatellite) motifs. Polymerase chain reaction (PCR) primer pairs were designed for 44 candidates and tested on a selection of samples collected from the wild. Seven of these PCR primer combinations produced interpretable polymorphic products. These have been assayed in 176 plants representing all the New Zealand species of Nothofagus subgenus Fuscospora (Nothofagus fusca, Nothofagus truncata and Nothofagus solandri). These markers will be applied to taxonomic, evolutionary and ecological studies of New Zealand beech.

Patterns of fungal diversity in New Zealand Nothofagus forests

The development of protocols for the conservation of fungi requires knowledge of the factors controlling their distribution, diversity, and community composition. Here we compare patterns of variation in fungal communities across New Zealand's Nothofagus forests, reportedly the most myco-diverse in New Zealand and hence potentially key to effective conservation of fungi in New Zealand. Diversity of leaf endophytic fungi, as assessed by culturing on agar plates, is assessed for three Nothofagus sp. growing in mixed stands from four sites. Host species was found to have a greater influence on fungal community assemblage than site. The leaf endophyte communities associated with Nothofagus solandri and Nothofagus fusca (both Nothofagus subgenus Fuscopora), were more similar to each other than either were to the community associated with Nothofagus menziesii (Nothofagus subgenus Lophozonia). The broad taxonomic groups isolated, identified on the basis of internal transcribed spacer (ITS) sequences, were similar to those found in similar studies from other parts of the world, and from an earlier study on the endophyte diversity in four podocarp species from New Zealand, but there were few matches at species level. Average levels of endophyte species diversity associated with single Nothofagus species and single podocarp species were similar, despite historical literature and collection data recording more than twice as many fungal species on average from the Nothofagus species. The significance of these findings to fungal conservation is discussed.

Phylogeographically concordant chloroplast DNA divergence in sympatric Nothofagus s.s. How deep can it be?

• Here, we performed phylogenetic analyses and estimated the divergence times on mostly sympatric populations of five species within subgenus Nothofagus. We aimed to investigate whether phylogenetic relationships by nuclear internal transcribed spacer (ITS) and phylogeographic patterns by chloroplast DNA (cpDNA) mirror an ancient evolutionary history that was not erased by glacial eras. Extant species are restricted to Patagonia and share a pollen type that was formerly widespread in all southern land masses. Weak reproductive barriers exist among them. • Fifteen cpDNA haplotypes resulted from the analysis of three noncoding regions on 330 individuals with a total alignment of 1794 bp. Nuclear ITS data consisted of 822 bp. We found a deep cpDNA divergence dated 32 Ma at mid-latitudes of Patagonia that predates the phylogenetic divergence of extant taxa. Other more recent breaks by cpDNA occurred towards the north. • Complex paleogeographic features explain the genetic discontinuities. Long-lasting paleobasins and marine ingressions have impeded transoceanic dispersal during range expansion towards lower latitudes under cooler trends since the Oligocene. • Cycles of hybridization-introgression among extant and extinct taxa have resulted in widespread chloroplast capture events. Our data suggest that Nothofagus biogeography will be resolved only if thorough phylogeographic analyses and molecular dating methods are applied using distinct genetic markers.

Phylogeny of Cyttaria inferred from nuclear and mitochondrial sequence and morphological data

Cyttaria species (Leotiomycetes, Cyttariales) are obligate, biotrophic associates of Nothofagus (Hamamelididae, Nothofagaceae), the southern beech. As such Cyttaria species are restricted to the southern hemisphere, inhabiting southern South America (Argentina and Chile) and southeastern Australasia (southeastern Australia including Tasmania, and New Zealand). The relationship of Cyttaria to other Leotiomycetes and the relationships among species of Cyttaria were investigated with newly generated sequences of partial nucSSU, nucLSU and mitSSU rRNA, as well as TEF1 sequence data and morphological data. Results found Cyttaria to be defined as a strongly supported clade. There is evidence for a close relationship between Cyttaria and these members of the Helotiales: Cordierites, certain Encoelia spp., Ionomidotis and to a lesser extent Chlorociboria. Order Cyttariales is supported by molecular data, as well as by the unique endostromatic apothecia, lack of chitin and highly specific habit of Cyttaria species. Twelve Cyttaria species are hypothesized, including all 11 currently accepted species plus an undescribed species that accommodates specimens known in New Zealand by the misapplied name C. gunnii, as revealed by molecular data. Thus the name C. gunnii sensu stricto is reserved for specimens occurring on N. cunninghamii in Australia, including Tasmania. Morphological data now support the continued recognition of C. septentrionalis as a species separate from C. gunnii. Three major clades are identified within Cyttaria: one in South America hosted by subgenus Nothofagus, another in South America hosted by subgenera Nothofagus and Lophozonia, and a third in South America and Australasia hosted by subgenus Lophozonia, thus producing a non-monophyletic grade of South American species and a monophyletic clade of Australasian species, including monophyletic Australian and New Zealand clades. Cyttaria species do not sort into clades according to their associations with subgenera Lophozonia and Nothofagus.

Cophylogeny and biogeography of the fungal parasite Cyttaria and its host Nothofagus, southern beech

The obligate, biotrophic association among species of the fungal genus Cyttaria and their hosts in the plant genus Nothofagus often is cited as a classic example of cophylogeny and is one of the few cases in which the biogeography of a fungus is commonly mentioned or included in biogeographic analyses. In this study molecular and morphological data are used to examine hypotheses regarding the cophylogeny and biogeography of the 12 species of Cyttaria and their hosts, the 11 species of Nothofagus subgenera Lophozonia and Nothofagus. Our results indicate highly significant overall cophylogenetic structure, despite the fact that the associations between species of Cyttaria and Nothofagus usually do not correspond in a simple one to one relationship. Two major lineages of Cyttaria are confined to a single Nothofagus subgenus, a specificity that might account for a minimum of two codivergences. We hypothesize other major codivergences. Numerous extinction also are assumed, as are an independent parasite divergence followed by host switching to account for C. berteroi. Considering the historical association of Cyttaria and Nothofagus, our hypothesis may support the vicariance hypothesis for the trans-Antarctic distribution between Australasian and South American species of Cyttaria species hosted by subgenus Lophozonia. It also supports the hypothesis of transoceanic long distance dispersal to account for the relatively recent relationship between Australian and New Zealand Cyttaria species, which we estimate to have occurred 44.6–28.5 mya. Thus the history of these organisms is not only a reflection of the breakup of Gondwana but also of other events that have contributed to the distributions of many other southern hemisphere plants and fungi.

Evidence of chloroplast capture in South American Nothofagus (subgenus Nothofagus, Nothofagaceae)

Subgenus Nothofagus, although geographically restricted at present to temperate areas of South America, has captured much attention in discussions of plant biogeography due to its widespread distribution through Gondwanan continents during the Tertiary. However, phylogenetic relationships within the subgenus Nothofagus have not yet been resolved. We examined geographic patterns of intraspecific and interspecific genetic variation to detect whether incongruences in nuclear or plastid DNA phylogenies occur, in order to better understand the evolutionary history of the subgenus Nothofagus. We conducted spatially-explicit sampling at 10 distinct locations throughout the range of austral South American forests and sampled all present Nothofagus species. We used ITS and chloroplast DNA sequences to estimate phylogenetic relationships. A phylogeny constructed from nuclear genes resolved the subgenus Nothofagus as monophyletic. We found that N. antarctica was a sister to a clade of evergreen species (N. betuloides, N. dombeyi, and N. nitida), while N. pumilio likely diverged earlier. Nine cpDNA haplotypes were distinguished in the subgenus Nothofagus which were associated to geographic locations rather than to taxonomic relationships. This species-independent cpDNA phylogeographic structures within the subgenus Nothofagus may be related to repeated chloroplast capture events over geological time in Patagonia.

The taphonomy of a remarkable leaf bed assemblage from the Late Oligocene–Early Miocene Gore Lignite Measures, southern New Zealand

A diverse assemblage of fossil leaves showing cellular detail is reported from the Late Oligocene–Early Miocene Gore Lignite Measures, southern New Zealand. The leaf-remains include at least five conifers, such as the genera Agathis, Dacrycarpus, Phyllocladus and Dacrydium, as well as a number of angiosperms including Gymnostoma, Nothofagus Subgenus Brassospora, Phormium, Proteaceae, Sapindaceae and Ericaceae. A parallel palynological study has identified a number of the same elements. The leaf layers represent litter horizons laid down in pools on the surface of a subtropical ombrotrophic forest mire that formed on an extensive low-lying coastal plain. Highly acidic water ponded in tree-fall depressions prevented microbial decay of the foliage. Taxa represented by both leaves and pollen are considered to have been components of the autochthonous swamp forest. Some other pollen grains are thought to represent local elements, based on the pollination biology of their nearest living relatives. Other taxa whose extant representatives are wind-pollinated may have grown further away.

Phytogeography and climate analysis of Nothofagus subgenus Brassospora in New Guinea and New Caledonia

Nothofagus subgenus Brassospora now occurs only in New Guinea and New Caledonia, but is well known from fossil deposits of South America, New Zealand, Antarctica and Australia. It is commonly used for palaeoclimatic interpretation, but the climate characteristics of the extant species have not been described. In this paper we used the climatic estimation software, BIOCLIM, to derive a climate profile of 24 variables for each of the 14 species of Nothofagus native to New Guinea, and lapse rates and isohyet maps to describe the annual mean temperature and rainfall range of the five species native to New Caledonia. The New Guinea species occur at annual mean temperatures ranging from 10.6 to 23.5°C, with annual precipitation of 1762–7733 mm. The first three axes of a principal components analysis explained 85% of the total variation, the first axis comprising temperature variables, the second comprising precipitation range and precipitation of the wet season, and the third axis comprising dry-season precipitation and annual and diurnal temperature range. Some species had distinct combinations of positions along these component axes, indicating clear niche differentiation with respect to climate. The New Caledonian species occur at annual mean temperatures of 14.5–23.5°C, and annual precipitation of c. 1500–3500 mm. Although there was no significant difference in annual mean temperature and precipitation between the New Guinea and New Caledonian species, comparison of median values across species suggests specialisation of most New Caledonian species towards slightly drier conditions than the New Guinea species that occur at similarly high annual mean temperatures. Use of subgenus Brassospora to interpret palaeoclimates should take into account the variation in climate experienced across the range of extant species.

New Zealand climate in the Neogene and implications for global atmospheric circulation

New Zealand has a good Neogene plant fossil record. During the Miocene it was without high topography and it was highly maritime, meaning that its climate, and the resulting vegetation, would be controlled dominantly by zonal climate conditions. Its vegetation record during this time suggests the climate passed from an ever-wet and cool but frostless phase in the Early Miocene in which Nothofagus subgenus Brassospora was prominent. Then it became seasonally dry, with vegetation in which palms and Eucalyptus were prominent and fires were frequent, and in the mid-Miocene, it developed a dry-climate vegetation dominated by Casuarinaceae. These changes are reflected in a sedimentological change from acidic to alkaline chemistry and the appearance of regular charcoal in the record. The vegetation then changed again to include a prominent herb component including Chenopodiaceae and Asteraceae. Sphagnum became prominent, and Nothofagus returned, but mainly as the subgenus Fuscospora (presently restricted to temperate climates). This is interpreted as a return to a generally wet, but now cold climate, in which outbreaks of cold polar air and frost were frequent. The transient drying out of a small maritime island and the accompanying vegetation/climate sequence could be explained by a higher frequency of the Sub-Tropical High Pressure (STHP) cells (the descending limbs of the Hadley cells) over New Zealand during the Miocene. This may have resulted from an increased frequency of ‘blocking’, a synoptic situation which occurs in the region today. An alternative hypothesis, that the global STHP belt lay at a significantly higher latitude in the early Neogene (perhaps 55°S) than today (about 30°S), is considered less likely because of physical constraints on STHP belt latitude. In either case, the difference between the early Neogene and present situation may have been a response to an increased polar–equatorial temperature gradient. This contrasts with current climate models for the geological past in which the latitude of the High Pressure belt impact is held invariant though geological time.

Ontogeny of pistillate flowers and inflorescences in Nothofagus subgenus Lophozonia (Nothofagaceae)

Cupule and pistillate inflorescence morphology is described in detail for the first time for a number of species in Nothofagus subgenus Lophozonia. The architecture and temporal appearance of prophylls, flowers, and cupule valves is dichasial; the cupule valves are third and higher order shoot systems. Comparative studies demonstrate that the dorsal rows of appendages on the cupule valves have a similar arrangement, and each row exhibits the same temporal pattern of development. This suggests that the cupule valves in all species have a modular construction and consist of rows of homologous appendages. This study clearly demonstrates the uniform groundplan of the pistillate inflorescence in this subgenus.

Holocene vegetation and climate change in southern New Zealand: linkages between forest composition and quantitative surface moisture reconstructions from an ombrogenous bog

AbstractThis paper presents a Holocene pollen record from an ombrotrophic bog in Southland, New Zealand, together with multiproxy data (testate amoebae, peat humification and plant macrofossils) from the same core to establish an independent semiquantitative record of peatland surface moisture. Linkages between reconstructed peatland surface moisture and regional forest composition are investigated using redundancy analysis of the forest pollen data constrained with predicted bog water‐table depths. Over 32% of the pollen data variance can be explained by surface moisture changes in the bog, suggesting a common cause of water‐table and regional vegetation change. Water tables were higher during the early to mid‐Holocene when the forest was dominated by podocarp taxa. Water tables lowered after about 3300 cal. yr BP coevally with the expansion of Nothofagus species, culminating with the dominance of Nothofagus subgenus Fuscospora in the past 1200 cal. yr BP. This is in apparent opposition to the warm/dry to cool/wet trend suggested by subjective interpretation of pollen data alone, from this and other studies. We suggest that during the late Holocene, drier summers associated with shifts in solar insolation caused reduced surface wetness and summer humidity, which together with a trend to cooler winters, apparently favoured the regeneration of Nothofagus species. Copyright © 2002 John Wiley & Sons, Ltd.

Nothofagus Cupules from Oligocene–Early Miocene Sediments at Balfour, Northwest Tasmania, Australia

The acceptable macrofossil record of Nothofagus (Nothofagaceae) is relatively sparse, with most published records represented by poorly preserved specimens. The Tasmanian fossil record is an exception to this, with an abundance of well‐preserved Nothofagus remains. Oligocene–Early Miocene sediments near Mount Balfour in northwest Tasmania preserve cupules of five species of Nothofagus in three of the four extant subgenera. Subgenus Lophozonia is represented by Nothofagus glandularis, subgenus Nothofagus by Nothofagus bulbosa, and subgenus Brassospora by Nothofagus peduncularis and the new species Nothofagus balfourensis and Nothofagus robustus. Subgenus Lophozonia is still present in Tasmania, but subgenus Nothofagus is now restricted to South America and subgenus Brassospora to New Guinea and New Caledonia. These three subgenera have only been recorded together previously in the Early Oligocene in Tasmania. This record extends that combination, indicating a long and presumably stable ecological relations...

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.