Natural hybridization between a deciduous (Nothofagus antarctica, Nothofagaceae) and an evergreen (N. dombeyi) forest tree species: evidence from morphological and isoenzymatic traits

BACKGROUND AND AIMS: Trees with a partial leaf-shedding pattern and other morphological features a priori considered intermediate between those of the deciduous Nothofagus antarctica (G. Forster) Oersted and the evergreen N. dombeyi (Mirb.) Oersted (Nothofagaceae) were found in natural stands. The hybridization between a deciduous and an evergreen species of Nothofagus has not been reported so far in natural communities. METHODS: The putative hybrids and the two presumed parental species were compared using 14 enzyme systems as well as shoot, leaf and reproductive morphology. KEY RESULTS: Six enzyme systems showed good resolution (MDH-B, IDH, SKDH, 6-PGDH, GOT and PGI) and in four of them (PGI, MDH-B, SKDH and 6-PGDH) the putative hybrids showed intermediate zymogram patterns between N. antarctica and N. dombeyi. Both principal coordinates analysis on isozyme data and principal components analysis (PCA) on quantitative morphological traits of shoots and leaves separated both parental species and located the putative hybrids closer to N. antarctica than to N. dombeyi. In the PCA, the number of basal cataphylls and the length : width ratio of leaves were the variables most discriminating among shoots of the three entities. The putative hybrids were intermediate between both species regarding leaf vernation, outline and venation, variation in leaf shape (length/width) with position on the parent shoot and in staminate inflorescence and cupule morphology. For other morphological traits, the putative hybrids resembled one of the parental species or differed from both species (e.g. valve morphology). CONCLUSIONS: Isoenzymatic and morphological data sets support the idea of the hybrid nature (probably F1 generation) of the semi-deciduous trees found. Nothofagus antarctica and N. dombeyi are probably more closely related than previously assumed. The relevance of pollen type in revealing evolutionary relationships between Nothofagus species is supported, and that of leaf-shedding pattern is rejected.     

Relative extents of preformation and neoformation in tree shoots: Analysis by a deconvolution method

BACKGROUND AND AIMS: Neoformation is the process by which organs not preformed in a bud are developed on a growing shoot, generally after preformation extension. The study of neoformation in trees has been hindered due to methodological reasons. The present report is aimed at assessing the relative importance of preformation and neoformation in the development of shoots of woody species. METHODS: A deconvolution method was applied to estimate the distribution of the number of neoformed organs for eight data sets corresponding to four Nothofagus species and a Juglans hybrid. KEY RESULTS: The number of preformed organs was higher and less variable than the number of neoformed organs. Neoformation contributed more than preformation to explain full-size differences between shoots developed in different positions within the architecture of each tree species. CONCLUSIONS: Differences between the distributions of the numbers of preformed and neoformed organs may be explained by alluding to the duration of differentiation and extension for each of these groups of organs. The deconvolution of distributions is a useful tool for the analysis of neoformation and shoot structure in trees.     

Genetic diversity in Nothofagus alessandrii (Fagaceae), an endangered endemic tree species of the coastal maulino forest of Central Chile

BACKGROUND AND AIMS: The endemic tree Nothofagus alessandrii (Fagaceae) has been historically restricted to the coastal range of Region VII of central Chile, and its forests have been increasingly destroyed and fragmented since the end of the 19th century. In this study, the patterns of within- and among-population genetic diversity in seven fragments of this endangered narrowly endemic tree were examined. METHODS: Allozyme electrophoresis of seven loci of N. alessandrii was used to estimate genetic diversity, genetic structure and gene flow. KEY RESULTS: High levels of genetic diversity were found as shown by mean expected heterozygosity (H(e) = 0.182 +/- 0.034), percentage of polymorphic loci (P(p) = 61.2 %), mean number of alleles per locus (A = 1.8) and mean number of alleles per polymorphic locus (A(p) = 2.3). Genetic differentiation was also high (G(ST) = 0.257 and Nm = 0.7). These values are high compared with more widespread congeneric species. CONCLUSIONS: Despite its endemic status and restricted geographical range N. alessandrii showed high levels of genetic diversity. The observed patterns of diversity are explained in part by historical processes and more recent human fragmentation.     

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.     

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.     

Climatic relationships of some New Zealand forest tree species

Abstract. A dataset of some 10 000 plots was used to describe the climatic relationships of 33 widespread New Zealand tree species. Estimates of mean annual temperature, temperature seasonality, mean annual solar radiation, and moisture balance were derived from mathematical surfaces fitted to climate station data. Plots were also categorized into five lithological classes and three drainage classes. Generalized additive models were used to examine species/environment relationships. Mean annual temperature and mean annual solar radiation are most strongly correlated with current tree distributions, followed by moisture balance, temperature seasonality, lithology, and drainage. Most broad-leaved tree species other than Nothofagus spp. reach their greatest levels of occurrence in warm, moist environments with high solar radiation. In contrast, Nothofagus spp. generally reach their greatest levels of occurrence in cooler and/or lower insolation environments, and all have lower levels of occurrence on rhyolitic substrates which have resulted from large-scale geomorphic disturbance, mostly over the past few thousand years. Although coniferous species have widely differing climatic optima, many are biased towards lithological classes characterized either by large-scale geomorphic disturbance or harsh edaphic conditions. The relevance of these results to particular synecological questions is briefly discussed. Continuing adjustments in the range of slow-dispersing Nothofagus spp. are strongly suggested, and the climatic suitability of extensive rhyolitic basins in the central North Island, from which these species are largely absent, is confirmed.     

Effect of volcanic ash deposition on length and radial growths of a deciduous montane tree (Nothofagus pumilio)

Extreme environmental events such as volcanic eruptions can trigger plant responses that largely exceed those recorded for moderate‐intensity disturbances. We assessed the effects of the June 2011 eruption of the Puyehue – Cordón Caulle volcano on the length and radial growths of juvenile Nothofagus pumilio trees at two sites located 20 (with >40 cm ash accretion) and 75 (without ash) km from the volcano. Variations in length and radial growth were evaluated for the periods 1999–2013 and 1993–2013 respectively; pre‐ and post‐eruption growth rates were computed. The length growth of the N. pumilio trees located close to the volcano increased significantly after the eruption: shoot extensions during the growing season after the eruption were, on average, two to three times longer than average according to ontogenetic growth trends. Variations in radial growth after the eruption were comparatively less noticeable than those in length growth. No significant effects of the eruption were observed in those trees located 75 km from the volcano. In order to explain the exceptionally positive response of N. pumilio's length growth to the volcanic eruption, three non‐exclusive explanations were proposed: (i) thick ash layers increase water retention in the soil; (ii) volcanic ash facilitates the access of plants to nutrients; and (iii) volcanic ashes decrease herbivory and competition. The comparatively lower sensitivity of radial growth to this extreme volcanic event is also noteworthy. These findings highlight the need to further examine how large‐scale volcanic events influence structure and/or functioning of ecosystem in the Patagonian forest.     

Coevolution and the adaptive value of autumn tree colours: colour preference and growth rates of a southern beech aphid

The evolutionary explanation for the change in leaf colour during autumn is still debated. Autumn colours could be a signal of defensive commitment towards insects (coevolution) or an adaptation against physical damage because of light at low temperatures (photoprotection). These two hypotheses have different predictions: (1) under the coevolution hypothesis, insects should not prefer red leaves in autumn and grow better in spring on trees with green autumn leaves; and (2) under the photoprotection hypothesis, insects should prefer and grow better on trees with red leaves because they provide better nutrition. Studying colour preference in autumn and growth rates in spring of a southern beech aphid species (Neuquenaphis staryi) on Nothofagus alessandrii, we found preference for green leaves in autumn but no differential performance of aphids in spring. We suggest that aphid preference for green might have evolved to exploit better their host during the autumn rather than to improve their performance in spring.     

Structure and tree-fall gap dynamics of old-growth Nothofagus forests in Tierra del Fuego,Argentina

Abstract. Tree size and age structure, tree-fall and gap characteristics, and regeneration in gaps were studied in Nothofagus-dominated old-growth forests in Tierra del Fuego, Argentina. Gap-phase regeneration has resulted in all-aged populations for N. pumilio, N. betuloides, and Drimys winteri, and regeneration in gaps appears to be maintaining coexistence between species in mixed stands. N. betuloides fills many gaps via advance regeneration and some individuals persist for > 150 yr in the understory. Multiple periods of release and suppression indicate that N. betuloides may take advantage of several gap events to reach the main canopy. Likewise, Drimys grows well under closed canopy and can rapidly respond to gap formation, sometimes impeding the regeneration of N. betuloides. In contrast, N. pumilio regenerates in gaps mainly from seed or from advance regeneration of small, ephemeral seedlings. Gap turnover times in Fuegian forests were estimated at 300 - 500 yr, although gap formation was highly episodic and possibly associated with regionally extensive windstorms, earthquakes, and stand-level dieback. 92 % of gaps involved multiple tree-falls, and at least 53 % involved secondary expansion. Gap and tree-fall characteristics in Tierra del Fuego were similar to results from northern Patagonia, Chile, and New Zealand; however, we emphasize that regeneration of Nothofagus spp. and Drimys winteri in gaps depends on associated vegetation and varies along both local and regional environmental gradients.     

Environmental controls and patterns of cumulative radial increment of evergreen tree species in montane,temperate rainforests of Chiloé Island,southern Chile

We investigated the local environmental controls on daily fluctuations of cumulative radial increment and cambial hydration of three dominant, evergreen tree species from montane, Coastal rainforests of Chiloé Island, Chile (42° 22′ S). During 2 years (1997–1998 and 1998–1999) we recorded hourly cumulative radial increments using electronic band dendrometers in the long‐lived conifer Fitzroya cupressoides (Cupressaceae), the evergreen broad‐leaved Nothofagus nitida (Nothofagaceae), and the narrow‐leaved conifer Podocarpus nubigena (Podocarpaceae). We also measured soil and cambial tissue hydration using capacitance sensors, together with air and soil temperature and rainfall during the period of the study. In addition, we collected cores of these tree species to evaluate how dendrometer measurements reflect annual tree ring width. One‐year long daily time series of cumulative radial increments suggests that radial growth of Fitzroya cupressoides was initiated slowly in early spring, with a maximum in early summer. Multiple regressions showed positive relations between daily precipitation and radial index (i.e. the difference in cumulative radial increment of two consecutive days) in the three species. According to path analysis there was a significant direct effect of changes in tree hydration on radial index of the three focal species. In emergent, pioneer species such as Nothofagus and Fitzroya, radial index was negatively affected by changes in maximum air temperature and photosynthetically active radiation, probably because of high evapotranspiration demand on warm sunny days. The shade‐tolerant species Podocarpus nubigena was positively affected by photosynthetically active radiation. Our diel scale findings support the use of tree ring widths for reconstructing past climate in these southern temperate forests and provide evidence that rainforest trees may be highly sensitive to future declines in rainfall and temperature increases during summer.     

Vertical stratification in the spatial distribution of the beech scale insect (Ultracoelostoma assimile) in Nothofagus tree canopies in New Zealand

Abstract. 1. The degree of infestation by New Zealand sooty beech scale insects (Ultracoelostoma assimile, Homoptera: Margarodidae) varies dramatically among adjacent southern beech trees (Nothofagus spp., Fagaceae), but has previously been assumed to be uniformly or randomly distributed within individual host trees. In this study, a full‐census survey was conducted from ground level to canopy level on 14 naturally occurring, canopy‐dominant red beech (Nothofagus fusca) trees (size range 38.7–107.6 cm diameter at breast height) to determine the degree of within‐tree heterogeneity in herbivore density. 2. The within‐tree distribution of the sooty beech scale was vertically stratified and highly heterogeneous, with the greatest densities occurring on bark surfaces in the canopy rather than on the trunk, and on the lower rather than upper sides of the branches. The spatial distribution was strongly negatively correlated with trunk and branch diameter, and increasing bark thickness (as a function of diameter) provides a plausible explanation for differences in the establishment and population density of sooty beech scale insects with trunk and branch size. Furthermore, there was a significant change in the spatial distribution of scale insect populations on trunks and branches of trees of increasing diameter at breast height. This indicates a strong temporal component to the spatial dynamics of the sooty beech scale insect driven by changing host phenology. Future studies on phytophagous insects infesting large host trees need to consider more explicitly changes in population dynamics through space and time. 3. Because of the high degree of within‐tree heterogeneity in population density, the total population size of scale insects on an individual tree could not be predicted from any measure of population density low on the trunk. However, the dry weight biomass of sooty mould fungi growing on the ground beneath infested trees was a remarkably accurate predictor of the total population size of scale insects. The use of sooty mould fungi as a relative measure of population size could be incorporated into studies of other honeydew‐producing hemipterans, since the growth of sooty mould is a distinctive feature synonymous with high concentrations of honeydew production worldwide.     

Elevation‐dependent responses of tree mast seeding to climate change over 45 years

We use seed count data from a New Zealand mono‐specific mountain beech forest to test for decadal trends in seed production along an elevation gradient in relation to changes in climate. Seedfall was collected (1965 to 2009) from seed trays located on transect lines at fixed elevations along an elevation gradient (1020 to 1370 m). We counted the number of seeds in the catch of each tray, for each year, and determined the number of viable seeds. Climate variables were obtained from a nearby (<2 km) climate station (914‐m elevation). Variables were the sum or mean of daily measurements, using periods within each year known to correlate with subsequent interannual variation in seed production. To determine trends in mean seed production, at each elevation, and climate variables, we used generalized least squares (GLS) regression. We demonstrate a trend of increasing total and viable seed production, particularly at higher elevations, which emerged from marked interannual variation. Significant changes in four seasonal climate variables had GLS regression coefficients consistent with predictions of increased seed production. These variables subsumed the effect of year in GLS regressions with a greater influence on seed production with increasing elevation. Regression models enforce a view that the sequence of climate variables was additive in their influence on seed production throughout a reproductive cycle spanning more than 2 years and including three summers. Models with the most support always included summer precipitation as the earliest variable in the sequence followed by summer maximum daily temperatures. We interpret this as reflecting precipitation driven increases in soil nutrient availability enhancing seed production at higher elevations rather than the direct effects of climate, stand development or rising atmospheric CO₂ partial pressures. Greater sensitivity of tree seeding at higher elevations to changes in climate reveals how ecosystem responses to climate change will be spatially variable.