Morpho-ecological study on the South American species of the genus Nothofagus

The morphology of the 11 taxa of South American Nothofagus are compared. Thirty eight characteristics were taken into account: 12 from leaves, 3 from stipules, 3 from buds, 6 from cupules, 7 from fruits, 4 from petioles and 3 from male flowers. The data matrix, with average values of 100 measurements per taxon, was analyzed with multivariate statistical methods of classification and polar and spatial ordination. Five groups were established: The first one (Nothofagus obliqua, N. obliqua var. macrocarpa, N. leoni and N. alessandrii), with deciduous leaves, is adapted to mediterranean climatic conditions. The second groups (N. dombeyi, N. nitida and N. antarctica), with small leaves which are evergreen for the first two species and deciduous for the last, develops under temperate humid conditions typical of the Valdivian region. The three remaining groups correspond to isolated species with different requirements: N. glauca, has large deciduous leaves and colonizes the most xerophytic biotope that a Nothofagus in central Chile can tolerate. N. pumilio, with mid-sized deciduous leaves, is adapted to the cold and dry zones of the southern Andes. Finally, N. betuloides, with small evergreen leaves, grows in the cold/humid somewhat boggy conditions of the Magellanic region.     

Foliar nitrogen and phosphorus dynamics of three Chilean Nothofagus (Fagaceae) species in relation to leaf lifespan

This study examined foliar nutrient dynamics and nutrient resorption (retranslocation) in three species of Chilean Nothofagus (Fagaceae) that differed in leaf lifespan and elevational distribution. In our central Chile study area the elevations at which these three species are most abundant increase from N. obliqua (deciduous) at low elevations to N. dombeyi at intermediate elevation and N. pumilio (deciduous) at higher elevations up to treeline. We sampled a single stand at 1680 m in which all three species co-occurred. Nothofagus dombeyi leaves were structurally heavier, with specific leaf mass approximately twice that of the two deciduous species. On a concentration basis, foliar N increased in the order N. dombeyi < N. pumilio < N. obliqua and foliar P increased in the order N. dombeyi < N. obliqua < N. pumilio. However, when the differences in specific leaf mass among species were taken into account by calculating N and P content on a leaf area basis, N. dombeyi had the greatest N and P content. N and P remained relatively constant throughout most of the 4-yr N. dombeyi leaf lifespan, then decreased prior to abscission. Nothofagus dombeyi resorbed significantly less N (44-50%) than did the two deciduous species (63-78%), both on proportional and absolute bases. In contrast, N. pumilio and N. dombeyi resorbed similar amounts of P prior to abscission (40-50%), whereas no significant resorption of P from leaves of N. obliqua was noted. We use these results to clarify the relative importance of environmental gradients associated with elevation vs. genetically fixed leaf lifespans in controlling the nutrient dynamics of these congeneric tree species.     

Axis differentiation in two South American Nothofagus species (Nothofagaceae)

An analysis was carried out on the length, diameter and number of leaves, and the ratios between these variables for current-year growth units (sibling growth units) derived from different nodes of previous-year growth units (parent growth units) of young Nothofagus dombeyi and Nothofagus pumilio trees. Changes in sibling growth unit length, diameter, and number of leaves with position on the parent growth unit were assessed. In both species, sibling-growth unit morphology varied according to both the axis type of the parent growth unit and the position of the sibling growth unit on its parent growth unit. For the largest parent growth units, the length, diameter and number of leaves of their sibling growth units decreased from distal to proximal positions on the parent growth unit. Distal sibling growth units had a more slender stem and longer internodes than proximal sibling growth units. Sibling growth units in equivalent positions tended to have a more slender stem for N. dombeyi than for N. pumilio. Long main-branch growth units of N. pumilio had longer internodes than those of N. dombeyi; the converse was true for shorter growth units. The growth unit diameter/leaf number ratio was consistently higher for N. pumilio than for N. dombeyi. Nothofagus pumilio axes would go through a faster transition from an 'exploring' morphology to an 'exploiting' morphology than N. dombeyi axes. Within- and between-species variations in growth unit morphology should be considered when assessing the adaptive value of the branching pattern of plants.     

Genetic variation in a geographically restricted and two widespread species of South American Nothofagus

The amount and distribution of genetic variation is compared using starch gel electrophoresis among populations of the three evergreen species of South American Nothofagus : the geographically restricted Nothofagus nitida (Phil.) Krasser and the widespread Nothofagus betuloides (Mirb.) Oerst. and Nothofagus dombeyi (Mirb.) Oerst. Ten enzyme systems were resolved coding for fifteen putative genetic loci. N. betuloides and N. dombeyi were more genetically variable than N. nitida ; they had higher total number of alleles, mean number of alleles per locus, percent of polymorphic loci, mean expected heterozygosity, and mean total genetic diversity. These results are in agreement with the hypothesis that widespread species are often associated with historically large and more continuously distributed populations which in turn can maintain higher levels of genetic variation. Conversely, the genetic structure of N. nitida including its low genetic identity with the other two species and the presence of four unique alleles support the hypothesis that N. nitida has been isolated from the other two species for a considerable period of time. Genetic data in concert with the fossil record indicate that these taxa may be members of a pre-Pleistocene flora and consequently their genetic structures reflect a more ancient evolutionary history than previously hypothesized.     

Mass loss and nutrient release from decomposing evergreen and deciduous Nothofagus litters from the Chilean Andes

Abstract Leaf litter decomposition experiments were conducted on two deciduous (Nothofagus obliqua (roble)) and Nothofagus pumilio (lenga)) and one evergreen (Nothofagus dombeyi) Nothofagus (Nothofagaceae) species from a single Chilean forest in order to understand how congeneric trees with differing leaf lifespans impact the soil in which they grow. Single‐species litter samples were decomposed in a mixed hardwood forest in Ohio and in a deciduous‐evergreen Nothofagus forest in Chile. In the Ohio forest, the two deciduous species’ litters decomposed at k ≈ 1.00 per year and the evergreen at k ≈ 0.75 per year. In Chile k ranged from k ≈ 0.06 (N. obliqua) to k ≈ 0.23 (N. pumilio) per year. In both experiments, N and P were released faster from the deciduous litters than from evergreen litter. In Ohio, evergreen litter immobilized more N and P for a longer time period than did deciduous litter. As N. dombeyi stands tend to have lower available soil N and P in this particular mixed Nothofagus forest, the increased time of N and P immobilization by N. dombeyi litter suggests a feedback role of the tree itself in perpetuating low N and P soil conditions.     

Relationships among Tasmanian Tertiary Nothofagus (Nothofagaceae) populations

Three leaf and two cupule forms of Nothofagus from the Early Oligocene deposit at Lea River are assigned to the previously described species Nothofagus tasmanica, N. lobata, N. gunnii, N glandularis and N. bulbosa. A single leaf specimen is possibly referable to Nothofagus subgenus Brassospora. Nothofagus microphylla , previously described from Monpeelyata, is separated into two species. These two species are N. microphylla and a new deciduous species named Nothojagus martinii. N. murophylla is transferred from the Nothofagus subgenus Lophozonia to subgenus Nothofagus , based upon cuticular characteristics. Nothw bulbosa and Nothojiis gunnii are also present at Monpeelyata. It is proposed that both N. lobata and N. microphylla have heterophyllous shoots and thus two leaf morphologies. Evolutionary and environmental relationships between Nothofagus species at Lea River, Monpeelyata and Little Rapid River are discussed. Large scale disturbance and increased seasonality are predicted to have caused the extinction of Nothofqus subgenus Nothofagus in Australia.     

A revised infrageneric classification of Nothofagus (Fagaceae)

HILL, R. S. & READ, J., 1991. A revised infrageneric classification of Nothofagus (Fagaceae). An examination of cupule morphology, leaf architecture and cuticular morphology of Nothofagus species demonstrates that the existing infrageneric classifications are inaccurate. Four subgenera are proposed, based on these characteristics as well as other morphological information. This analysis indicates that the traditional pollen groupings of Nothofagus closely reflect the infrageneric taxonomy. It is hypothesized that deciduousness is primitive in Nothofagus , and the evergreen habit has arisen more than once, and therefore the deciduous or evergreen habit is invalid as a primary taxonomic character.     

Insect antifeedant compounds from Nothofagus dombeyi and N. pumilio

A bioassay-guided purification of the extracts of Nothofagus dombeyi and N. pumilio leaves yielded several triterpenes and flavonoids including 2-O-acetylmaslinic acid, 3-O-acetyl 20,24,25-trihydroxydammarane, and 3,20,24,25-tetrahydroxydammarane as new natural products. All the isolated compounds were assessed for antifeeding activity against the 5th instar larvae of Ctenopsteustis obliquana. 12-Hydroxyoleanolic lactone and pectolinarigenin from N. dombeyi and dihydrooroxylin A from N. pumilio, showed significant antifeeding activity.     

Microsatellites for use in Nothofagus cunninghamii (Nothofagaceae) and related species

Nothofagus cunninghamii (myrtle) has a widespread distribution through the temperate rainforest of southeastern Australia with some disjunct populations existing in putative glacial refugia. Polymorphic nuclear markers are required to resolve the biogeographical history of the species and will also be useful for conservation and forestry applications in this and other species of Nothofagus. Fourteen polymorphic microsatellite loci were isolated from N. cunninghamii, with four to 10 alleles amplified in 15 individuals tested. Transferability to other species of Nothofagus was successful, with six loci transferring to all eight other species tested.     

Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients

Background and Aims

The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous–evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations.

Methods

Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous–evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra).

Key Results

Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood.

Conclusions

Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone.     

Post-glacial vegetation in the South Patagonian territory of the giant ground sloth, Mylodon

By comparison with the major modern plant communities of southern Patagonia, the changing post-glacial vegetation of the region is reconstructed from macrofossils derived from deposits in the Cueva del Mylodon, Ultima Esperanza, S. Chile. The oldest deposits, carbon-dated as c. 12,400 B.P., comprise dung of the extinct ground sloth Mylodon darwinii and show the animal to have fed entirely on Cyperaceae, Gramineae and species associated with these in the modern cool, wet sedge-grasslands of western Patagonia, communities which would be expected after the retreat of the ice. Overlying deposits of well-preserved, wind-blown leaf-litter permitted some quantitative analyses which show a rise of evergreen forest dominated by Nothofagus betuloides that reached its maximum c. 7000 B.P. and then declined as it was replaced by deciduous N. pumilio forest. A break in the fossil plant record, covering the human occupation about 5643 B.P. and the subsequent final appearance of Mylodon remains, is followed by evidence of mixed evergreen/deciduous forest in which Nothofagus pumilio gradually increases in importance to give deciduous forest some 2500 years ago similar to that found in the environs of the cave in historical times. Comparison with pollen diagrams shows that the modern climatic and vegetation difference between E. Fudgia and Ultima Eeperanza has persisted throughout the post-glacial period.     

Differential hypogeous sporocarp production from Nothofagus dombeyi and N. pumilio forests in southern Argentina

Mycorrhizal fungi that form hypogeous sporocarps are an important component of the temperate forest soil community. In many regions, such as the Nothofagus forest in the Patagonian Andes, this group of fungi has been poorly studied. Here we examined the spring and autumn community composition of "sequestrate fungi", based on sporocarp production in pure forests of Nothofagus dombeyi (evergreen) and N. pumilio (deciduous). We investigated the possible relationships between these communities and environmental factors over 2 y. The rarefaction curves and the minimal richness estimates converged at nearly the same level for each forest type, and the asymptotes suggested that the sampling effort was sufficient to capture most of the hypogeous sporocarp richness in these forest stands. In total 27 species were recovered. Basidiomycota, Ascomycota and Glomeromycota respectively accounted for nine, two and one genera. Species richness of hypogeous sporocarps varied in relation to forest type but not to season (fall and spring), whereas sporocarp biomass varied according to an interaction between season and forest type. Species richness and sporocarp biomass were positively correlated with rainfall and negatively correlated with altitude. In addition sporocarp species richness was positively related to number of trees per transect. We found that two different forest stands, each dominated by different species of Nothofagus, exhibited different hypogeous sporocarp communities.     

Micro-site performance of evergreen and deciduous dwarf shrubs in a subarctic heath in relation to nitrogen status

The relationship between micro-site nitrogen status (total soil N conc., total vegetation N, and leaf N conc.) and performance (biomass) for two evergreen and two deciduous dwarf shrubs was studied in a subarctic heath. One deciduous species, Vaccinium uliginosum , dominated on sites having the highest vegetation nitrogen content and highest leaf nitrogen concentrations. The evergreen Empetrum hermaphroditum dominated on sites with a low total vegetation nitrogen content and low leaf nitrogen concentrations. Two other species, V. myrtillus (deciduous) and V. vitis-idaea (evergreen), showed intermediate patterns that were not clearly separated from either V. uliginosum or Empetrum . Soil nitrogen content showed no significant differences between micro-sites. Possible physiological reasons for the observed patterns are discussed.     

Preformation and neoformation in shoots of Nothofagus antarctica (G. Forster) Oerst. (Nothofagaceae) shrubs from northern Patagonia

The size (length and diameter) and number of leaf primordia of winter buds of Nothofagus antarctica (G. Forster) Oerst. shrubs were compared with the size and number of leaves of shoots derived from buds in equivalent positions. Buds developed in two successive years were compared in terms of size and number of leaf primordia. Bud size and the number of leaf primordia per bud were greater for distal than for proximally positioned buds. Shoots that developed in the five positions closest to the distal end of their parent shoots had significantly more leaves than more proximally positioned shoots of the same parent shoots. The positive relationship between the size of a shoot and that of its parent shoot was stronger for proximal than for distal positions on the parent shoots. For each bud position on the parent shoots there were differences in the number of leaf primordia per bud between consecutive years. The correlations between the number of leaf primordia per bud and bud size, bud position and parent shoot size varied between years. Only shoots produced close to the distal end of a parent shoot developed neoformed leaves; more proximal sibling shoots consisted entirely of preformed leaves. Leaf neoformation, a process usually linked with high shoot vigour in woody plants, seems to be widespread among the relatively small shoots developed in N. antarctica shrubs, which may relate to the species' opportunistic response to disturbance.     

Age structure and dynamics of Patagonian beech forests in Torres del Paine National Park,Chile

This study documents the stem size and age-structure in forests dominated by different species of Nothofagus in Torres del Paine National Park (51° S), in the Chilean Patagonian region. We also explored the relationship between the various types of Nothofagus forest and postglacial succession. Pioneer stands on moraine fields 1–2 km of the glacier front are dominated by Nothofagus betuloides and Nothofagus antarctica. Moraines appear to be first colonized by the evergreen N. betuloides, followed within 5–7 years by deciduous N. antarctica. Nothofagus antarctica may replace the former species and develop monospecific stands on glacial valleys. Most trees in the N. antarctica stand studied were older than 40 years and floods may cause a significant mortality of young trees. Recruitment from seed seems to be infrequent. Old-growth stands dominated by deciduous Nothofagus pumilio occupy more stable substrates, and probably represent the last stage of postglacial succession. This long-lived tree species had recorded ages over 200 years. The canopy of N. pumilio forests appears to be a mosaic of even-aged, old-growth patches. We propose that regeneration episodes follow the blowdown of a large portion of the canopy, with long intervals with little or no regeneration. Windstorms may be an important force influencing the regeneration of N. pumilio. Exogenous disturbances, such as floods and windstorms, are an integral part of the forest cycle in the Patagonian region.     

Natural Hybrids between Ligularia vellerea and L.subspicata (Asteraceae: Senecioneae)

Three types of intermediate morphological individuals were found in the sympatric distribution areas of Ligularia vellerea and Lsubspicata. These intermediate individuals were hypothesized to be the hybrids of these two species based on the detailed comparison of their morphological characters. To test this hypothesis, we compared the DNA sequence of Lvellerea, Lsubspicata and the intermediate individuals by using the internal transcribed spacer region of nuclear ribosomal and atpB rbcL intergenic spacer region of the chloroplast. Direct sequencing of ITS clearly had additives and then cloned sequencing cloned out two kinds of sequences that were identical to those putative parental species, thus these putative hybrids were approved. With different morphological characters among three types, their plastid donors were all Lsubspicata according to the maternal sequences of atpB rbcL.     

Clarification of the name Nothofagus alpina and a new epithet for a Nothofagus hybrid

Arguments in favour of using Nothofagus alpina (Poepp. & Endl.) Oerst. (Fagaceae) rather than N. procera Oerst. or N. nervosa (Phil.) Krasser as the correct name for rauli are presented. We also refute suggestions that N. alpina is based on hybrid material. The hybrid of N. alpina with N. obliqua (Mirb.) Blume is formally described as N . ×  dodecaphleps Mike L. Grant & E. J. Clement and a key to the deciduous taxa of Nothofagus is provided.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 447–451.     

Morphological evidence for natural intergeneric hybridization in the New Zealand Gnaphalieae (Compositae): Anaphalioides bellidioides × Ewartia sinclairii

Wild intergeneric hybrids are recorded among six genera in the New Zealand Gnaphalieae, but little evidence substantiating their identity has been published. In this paper we evaluate the identity of putative hybrids between two species of Gnaphalieae endemic to New Zealand, Anaphalioides bellidioides and Ewartia sinclairii . The two species are occasionally sympatric in the north-east of the South Island and wild putative hybrids are rarely encountered. To test the hybridity hypothesis, specimens from nine putative hybrids were collected from one site and analysed for 76 morphological characters. The putative hybrids possessed predominantly intermediate or parental character states in comparison with A. bellidioides and E. sinclairii . Extreme traits were less common and a single novel character state was recorded in two hybrids only. Non-metric multidimensional scaling of dissimilarities derived from both continuous and discrete characters was strongly supportive of the hybridity hypothesis. Canonical discriminant analysis of the continuous characters was supportive for some putative hybrids only, in part reflecting the possession of extreme states in characters with high loadings. One of the putative hybrids was strongly indicated to be a backcross with A. bellidioides . The taxonomic treatment of cross-compatible genera is discussed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 59−75.     

Molecular tests of the proposed diploid hybrid originof Gilia achilleifolia (Polemoniaceae)

Gilia achilleifolia is a putative diploid hybrid species. Hybrid origin was hypothesized based on traditional biosystematicevidence (i.e., morphological, cytological, and crossability data),which may be insufficient to establish genealogical history. Here,phylogenetic analysis of sequence data from the internal transcribedspacer (ITS) regions is used to examine the relationship between theputative hybrid species and its proposed parents. Isozyme variation isassayed to test for genetic additivity in the putative hybrid taxon andmorphological data are analyzed cladistically to evaluate the charactersthat led to the original hypothesis of hybrid origin. The ITS-basedgene tree placed G. achilleifolia in two divergent clades, eachsister to one of the putative parental lineages. Little isozymeadditivity was observed and G. achilleifolia possessed sixunique alleles among 42 alleles observed. However, ITS and isozymetrees differed in their placement of the two lineages of G.achilleifolia; both lineages are closer to a third putative parentin the isozyme tree. Also, G. achilleifolia is intermediate orpolymorphic for all nine morphological characteristics differentiatingthe parental species. Sorting of ancestral polymorphisms cannot easilyaccount for expression patterns of seven of these characters. In ourview, these results fail to distinguish between alternative hypothesesof ancient hybrid origin and divergent evolution, belying the difficultyof detecting ancient hybrids.     

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.