Out of the Qinghai-Tibet Plateau: evidence for the origin and dispersal of Eurasian temperate plants from a phylogeographic study of Hippophaë rhamnoides (Elaeagnaceae)

Numerous temperate plants now distributed across Eurasia are hypothesized to have originated and migrated from the Qinghai-Tibet Plateau (QTP) and adjacent regions. However, this hypothesis has never been tested through a phylogeographic analysis of a widely distributed species. Here, we use Hippopha? rhamnoides as a model to test this hypothesis. We collected 635 individuals from 63 populations of the nine subspecies of H. rhamnoides. We sequenced two maternally inherited chloroplast (cp) DNA fragments and also the bi-paternally inherited nuclear ribosomal ITS. We recovered five major clades in phylogenetic trees constructed from cpDNA and internal transcribed spacer (ITS) sequence variation. Most sampled individuals of six subspecies that are distributed in northern China, central Asia and Asia Minor/Europe, respectively, comprised monophyletic clades (or subclades) nested within those found in the QTP. Two subspecies in the QTP were paraphyletic, while the placement of another subspecies from the Mongolian Plateau differed between the ITS and cpDNA phylogenetic trees. Our phylogeographic analyses supported an 'out-of-QTP' hypothesis for H. rhamnoides followed by allopatric divergence, hybridization and introgression. These findings highlight the complexity of intraspecific evolutions and the importance of the QTP as a center of origin for many temperate plants.     

Heterogeneity in soil water and light environments and dispersal limitation: what facilitates tree species coexistence in a temperate forest?

In the present study, we analysed the habitat association of tree species in an old‐growth temperate forest across all life stages to test theories on the coexistence of tree species in forest communities. An inventory for trees was implemented at a 6‐ha plot in Ogawa Forest Reserve for adults, juveniles, saplings and seedlings. Volumetric soil water content (SMC) and light levels were measured in 10‐m grids. Relationships between the actual number of stems and environmental variables were determined for 35 major tree species, and the spatial correlations within and among species were analysed. The light level had no statistically significant effect on distribution of saplings and seedlings of any species. In contrast, most species had specific optimal values along the SMC gradient. The optimal values were almost identical in earlier life stages, but were more variable in later life stages among species. However, no effective niche partitioning among the species was apparent even at the adult stage. Furthermore, results of spatial analyses suggest that dispersal limitation was not sufficient to mitigate competition between species. This might result from well‐scattered seed distribution via wind and bird dispersal, as well as conspecific density‐dependent mortality of seeds and seedlings. Thus, both niche partitioning and dispersal limitation appeared less important for facilitating coexistence of species within this forest than expected in tropical forests. The tree species assembly in this temperate forest might be controlled through a neutral process at the spatial scale tested in this study.     

Phylogeography of Liquidambar styraciflua (Altingiaceae) in Mesoamerica: survivors of a Neogene widespread temperate forest (or cloud forest) in North America?

We investigate the genetic variation between populations of the American sweetgum (Liquidambar styraciflua), a tree species with a disjunct distribution between northeastern Texas and Mexico, by analyzing sequences of two chloroplast DNA plastid regions in Mesoamerica. Our results revealed phylogeographical structure, with private haplotypes distributed in unique environmental space at either side of the Trans‐Mexican Volcanic Belt, and a split in the absence of gene flow dating back ca. 4.2–1.4 million years ago (MYA). Species distribution modeling results fit a model of refugia along the Gulf and Atlantic coasts but the present ranges of US and Mesoamerican populations persisted disjunct during glacial/interglacial cycles. Divergence between the US and Mesoamerican (ca. 8.4–2.8 MYA) populations of L. styraciflua and asymmetrical gene flow patterns support the hypothesis of a long‐distance dispersal during the Pliocene, with fragmentation since the most recent glacial advance (120,000 years BP) according to coalescent simulations and high effective migration rates from Mesoamerica to the USA and close to zero in the opposite direction. Our findings implicate the Trans‐Mexican Volcanic Belt as a porous barrier driving genetic divergence of L. styraciflua, corresponding with environmental niche differences, during the Pliocene to Quaternary volcanic arc episode 3.6 MYA, and a Mesoamerican origin of populations in the USA.     

Phosphorus release from resuspended sediment in the shallow and wind-exposed Lake Arresø, Denmark

Wind-induced sediment resuspension occurs frequently in the shallow and eutrophic Lake Arresø, Denmark. The impact of resuspension on internal phosphorus loading was investigated by laboratory experiments studying P-release from the undisturbed sediment surface and by experiments simulating resuspension events.Phosphorus release from undisturbed sediment sampled in May and August was 12 mg and 4 mg m^–2 d^–1, respectively. During experimental simulation of resuspension, soluble reactive phosphate (SRP) increased by 20–80 µg l^–1, which indicates that a typical resuspension event in the lake would be accompanied by the release of 150 mg SRP m^–2. The internal P loading induced by resuspension is estimated to be 60–70 mg m^–2 d^–1, or 20–30 times greater than the release from undisturbed sediment.SRP release during simulation of resuspension was mainly dependent on the equilibrium conditions in the water column and was basically independent of the increase in suspended solids and the duration of resuspension. A second simulation of resuspension conducted 26 hours later, did not result in any further release of SRP from sediment sampled in May. In contrast, there was an additional SRP release from sediment sampled in August, indicating that an exchangable P pool, capable of altering equilibrium conditions, is built up between resuspension events.It is concluded that resuspension, by increasing the P flux between sediment and water, plays a major role in the maintenance of the high nutrient level in Lake Arresø. A relatively high release rate is maintained during resuspension because of the low Fe:P ratio and the high concentration of NH₄Cl-extractable P in the sediment.     

What controls the distribution of tropical forest and savanna?

Forest and savanna biomes dominate the tropics, yet factors controlling their distribution remain poorly understood. Climate is clearly important, but extensive savannas in some high rainfall areas suggest a decoupling of climate and vegetation. In some situations edaphic factors are important, with forest often associated with high nutrient availability. Fire also plays a key role in limiting forest, with fire exclusion often causing a switch from savanna to forest. These observations can be captured by a broad conceptual model with two components: (1) forest and savanna are alternative stable states, maintained by tree cover-fire feedbacks, (2) the interaction between tree growth rates and fire frequency limits forest development; any factor that increases growth (e.g. elevated availability of water, nutrients, CO(2)), or decreases fire frequency, will favour canopy closure. This model is consistent with the range of environmental variables correlated with forest distribution, and with the current trend of forest expansion, likely driven by increasing CO(2) concentrations. Resolving the drivers of forest and savanna distribution has moved beyond simple correlative studies that are unlikely to establish ultimate causation. Experiments using Dynamic Global Vegetation Models, parameterised with measurements from each continent, provide an important tool for understanding the controls of these systems.     

Abundance of climbing plants in a southern temperate rain forest: host tree characteristics or light availability?

Question: In a southern temperate rain forest, we addressed three questions: (1) Does the abundance of climbing plants increase with light availability? (2) Do host tree species differ in their susceptibility to vine infestation? (3) How does the relationship between host tree trunk diameter and relative abundance of vines vary with their climbing mechanism? Location: Two sites in the temperate evergreen rain forest of southern Chile: Puyehue (40°39′S, 72°09′W; 350 m a.s.l.) and Pastahue (42°22′S, 73°49′W; 285 m a.s.l.). Methods: We sampled vines in 60 25‐m² plots, with 20 plots in each of three light environments: mature forest, forest edges and canopy gaps. In each plot, for every tree ≥1.50‐m tall of any diameter we counted and identified all climbing plant individuals at a height of 1.30 m. We also counted, measured (trunk diameter at 1.30 m) and identified all these trees, and determined prevalence of vine infestation for each tree species. Results: Light availability in forest plots did not affect vine abundance when the number and size of host trees was taken into account. Overall, vine abundance increased with host tree trunk diameter. Tree species did not differ in the prevalence of vine infestation. The relative abundance of stem twiners and adhesive climbers decreased and increased with trunk diameter, respectively. The densities of stem twiners and adhesive climbers were negatively correlated across the forest. Conclusion: We provide further evidence that the pattern of vine abundance is independent of light availability in southern temperate rain forests, in contrast to results commonly reported for tropical rain forests. We also show that support suitability across the forest varies with the mechanism by which vines climb, probably due in part to biomechanical constraints and in part to vine interspecific competition, a virtually unexplored ecological factor.     

Climbing plants in a temperate rainforest understorey: searching for high light or coping with deep shade?

Background and Aims

While the climbing habit allows vines to reach well-lit canopy areas with a minimum investment in support biomass, many of them have to survive under the dim understorey light during certain stages of their life cycle. But, if the growth/survival trade-off widely reported for trees hold for climbing plants, they cannot maximize both light-interception efficiency and shade avoidance (i.e. escaping from the understorey). The seven most important woody climbers occurring in a Chilean temperate evergreen rainforest were studied with the hypothesis that light-capture efficiency of climbers would be positively associated with their abundance in the understorey.

Methods

Species abundance in the understorey was quantified from their relative frequency and density in field plots, the light environment was quantified by hemispherical photography, the photosynthetic response to light was measured with portable gas-exchange analyser, and the whole shoot light-interception efficiency and carbon gain was estimated with the 3-D computer model Y-plant.

Key Results

Species differed in specific leaf area, leaf mass fraction, above ground leaf area ratio, light-interception efficiency and potential carbon gain. Abundance of species in the understorey was related to whole shoot features but not to leaf level features such as specific leaf area. Potential carbon gain was inversely related to light-interception efficiency. Mutual shading among leaves within a shoot was very low (<20 %).

Conclusions

The abundance of climbing plants in this southern rainforest understorey was directly related to their capacity to intercept light efficiently but not to their potential carbon gain. The most abundant climbers in this ecosystem match well with a shade-tolerance syndrome in contrast to the pioneer-like nature of climbers observed in tropical studies. The climbers studied seem to sacrifice high-light searching for coping with the dim understorey light.     

Long-term changes of the δ15N natural abundance of plants and soil in a temperate grassland

Tracing back the N use efficiency of long-term fertilizer trials is important for future management recommendations. Here we tested the changes in natural N-isotope composition as an indicator for N- management within a long-term fertilization lysimeter experiment in a low mountain range pasture ecosystem at Rengen (Eifel Mountains), Germany. Cattle slurry (δ¹⁵N?=?8.9?±?0.5‰) and mineral fertilizers (calcium ammonium nitrate; δ¹⁵N?=??1.0?±?0.2‰) were applied at a rate between 0 and 480 kg N ha^?1?yr^?1 throughout 20 years from 1985 onwards. In 2006, samples were taken from different grass species, coarse and fine particulate soil organic matter, bulk soil and leachates. Total soil N content hardly changed during fertilization experiment. As also N leaching has been small within the stagnant water regime, most N was lost through the gaseous phase beside plant uptake and cutting. Unlike N uptake by plants, the process of N volatilization resulted in strong discrimination against the ¹⁵N isotope. As a consequence, the δ¹⁵N values of top soil samples increased from 1.8?±?0.4‰ to 6.0?±?0.4‰ and that of the plants from ?1.2?±?1.3‰ to 4.8?±?1.2‰ with increasing N fertilizer rate. Samples receiving organic fertilizer were most enriched in δ¹⁵N. The results suggest that parts of the fertilizer N signal was preserved in soils and even discovered in soil organic matter pools with slow N turnover. However, a ¹⁵N/¹⁴N isotope fractionation of up to 1.5‰ added to the δ¹⁵N values recovered in soils and plants, rendering the increase in δ¹⁵N value a powerful indicator to long-term inefficient N usage and past N management in the terrestrial environment.     

Is the Ca:Al ratio superior to pH,Ca or Al concentrations of soils in accounting for the distribution of plants in deciduous forest?

The distributions of vascular plants in south Swedish deciduous forests were related to exchangeable (exc) and soil solution concentrations of H⁺ (pH), Ca, Al and the Ca:Al ratios within these fractions. Topsoils (0–5 cm) of 172 sites with a pH_KCl of 3.2–3.9 (corresponding to 3.7–4.4 in soil solution) were used. In the soil solution both total Al_t and quickly reacting Al_r were determined. Exchangeable concentrations were generally well related to plant distributions, the highest correlation coefficients usually being given by pH_KCl>Ca_exc>Al_exc.>(Ca:Al)_exc. The (Ca:Al)_exc ratio was clearly inferior. Out of the soil solution variables studied, Ca concentration, followed by pH, was best correlated with plant distributions, Al_t, Al_r, and the Ca:Al ratios having similar and lower coefficients. It is concluded that the use of Ca:Al ratios as a general measure of Al toxicity in controlling plant distributions is rather problematic. It seems difficult to apply evidence for Ca-Al interactions from solution culture experiments to field conditions when measured as exchangeable or soil solution concentrations of the soil.     

δ15N of forest soil and understorey vegetation reflect the former agricultural land use

Since the middle of the 19th century, the area covered by forests in France has doubled. These new forests grow on previous agricultural lands. We have studied the influence of this agricultural history on the ¹⁵N abundance of present-day forests planted on farmlands in the Vosges mountains (north-eastern France) between 1898 and 1930. Different types of land use were identified from old cadastres (1814–1836) of 16 farms. Ancient forests adjacent to farmlands were used as controls. Former pastures, meadows, croplands, gardens and ancient forests were compared for soil δ¹⁵N (fraction <50 μm and total soil), C/N, P and N content and fern (Dryopteris carthusiana) δ¹⁵N. The mean δ¹⁵N of soil increased in the order ancient forests (+0.0‰)<pastures (+1.4‰)<croplands (+1.6‰)<meadows (+2.5‰)<gardens (+3.8‰). This increase in soil δ¹⁵N with the intensity of former land use was related to the former input of ¹⁵N-enriched manure, and to an activation of soil nitrification leading to ¹⁵N-depleted nitrate export on previously manured parcels. Fern δ¹⁵N increased in the same order as soil δ¹⁵N in relation to past land use. The mean δ¹⁵N of fern in ancient forests (–4.4‰) and former pastures (–3.4‰) was 5‰ lower than soil δ¹⁵N and the two variables were strongly correlated. The δ¹⁵N of fern in formerly manured parcels varied little (cropland: –2.7‰, meadows: –2.6‰ and gardens: –2.2‰) and independently of soil δ¹⁵N, suggesting that the soil sources of fern N differed between unmanured and manured parcels. Understorey plant δ¹⁵N and soil δ¹⁵N appear to be excellent tracers of previous land use in forests, and could be used in historical studies. The persistence of high isotopic ratios in previously manured parcels, almost a century after afforestation, suggests a long-term influence of former land use on the N cycle in forest soils. Received: 22 January 1999 / Accepted: 22 July 1999     

Seed dispersal mechanisms and the vegetation of forest islands in a West African forest-savanna mosaic (Comoé National Park,Ivory Coast)

The transition zone between forest and savanna is typically characterized by a dynamic patchwork of forest and savanna. We studied the woody plant species composition of 49 forest islands, 18 savanna, and 3 gallery forest plots in the Comoé National Park (Ivory Coast), West Africa's largest savanna reserve. TWINSPAN makes a clear distinction in vegetational composition between these three major habitat types but, nevertheless, more than 50% of the 292 species occur in at least 2 of them. The gallery forest is dominated by Cynometra megalophylla (Caesalpiniaceae), a genus known to dominate humid forests in other afrotropical regions. Ordination reveals four distinct categories of forest islands, (1) humid forests comparable to the gallery forest in their species composition, (2) dry disturbed and (3) dry undisturbed forests and (4) forests formerly inhabited by humans. Disturbed forests harbor more savanna species but also a distinct group of disturbance-tolerant forest species. Compared to other forest species, in this latter group we found an exceptionally high fraction of animal dispersed species (80% vs. 58%), while wind dispersed species or species lacking long distance seed dispersal mechanisms were correspondingly rare. This pattern occurs in spite of the fact that the frequency of wind dispersed species in general increases from dense humid forests to open dry forests and savanna. Species lacking long distance dispersal mechanisms are most abundant among those specialized on humid forests. These observations suggest that the species composition of forest islands is to some extent determined by the seed dispersal abilities of the different species.     

Are cloud forest tree structure and environment related in the Venezuelan Andes?

Cloud forest vegetation structure and composition were studied in the Venezuelan Andes at three sites in Mérida State. Although the sites are within 10 to 30 km of each other, climatic, geologic and topographic differences are remarkable. The main purpose of the study was to determine the relationship of specific environmental variables to forest vegetation characteristics, including basal area, tree height, density and diversity, and leaf area index (LAI). At 51 plots, all trees' diameter at breast height >10 cm were recorded and identified. Although the environment at the three sites is distinctive, the tree species composition of the most abundant species was very similar. None of the measured environment variables were significantly correlated with the measured vegetation structure variables, except LAI, which was correlated with slope orientation; LAI showed higher values at south‐facing plots. Tree height was relatively uniform, while basal area was highly variable and reached very high values. Stem densities were in the range reported elsewhere in cloud forests. Multivariate analysis using structure or composition data shows segregation of the plots by site. Principal component analyses by site indicate a minor impact of environmental factors on forest variables. At each site, a particular group of species are correlated with the ordination axes. We conclude that species pools and forest dynamics add to the complexity of the structure of the studied cloud forests.     

Regional distribution of genetically modified organisms (GMOs)—Up-scaling the dispersal and persistence potential of herbicide resistant oilseed rape (Brassisca napus)

Most genetically modified (GM) crop plants are designed to be grown on large areas. However, empirical investigations for risk assessment are limited in their temporal and spatial extent. In the case of GM crop plants it is difficult to test the relevance of anticipated risks on the same spatial scale as the intended use. Processes which are difficult to assess experimentally include combinatory effects, interactions between different integration levels, persistence, long distance dispersal and occurrence of rare events. To a limited extent, it is possible to combine results of investigations on small spatial scales in a way that large-scale and long-term implications on the regional scale can be analysed by using modelling and extrapolation approaches. It is thus possible to indicate some of the involved risks which are not accessible otherwise.In this paper we present the results of an extrapolation methodology comprising several scales from the field size up to the landscape level. This methodology aimed at analysing the implications of a large-scale release of genetically modified oilseed rape (GM OSR). The approach consisted of an extrapolation scheme beginning with a landscape analysis which generated representative scenarios considering climate and OSR cultivation characteristics. For the spatial extent of several fields this information was applied in an individual-based model representing ontogeny, dispersal and persistence of cultivated, volunteers and feral oilseed rape. In a final step, simulation results were extrapolated to the region of Northern Germany.Here we focus on the model results which were extrapolated to the regional level by applying a set of ecological indicators which allowed to assess potential implications on this level. These indicators included the number and distribution of flowering GM plants and the dynamics of GM OSR seeds in the soil seedbank. Specific results related to the long-term dynamics in the seedbank and volunteer development. Model results emphasise the long-term consequences of GM OSR cultivation and the explicit necessity to regard high variability in potential GMO admixture. This has to be considered when developing landscape management schemes for co-existence.The extrapolation approach presented here, integrates different traits to assess effects of GMOs on large spatial scales with respect to persistence and dispersal. The developed methodology is equally applicable for other crops, regions and different agricultural conditions.     

Ubiquitous presence of β-glucuronidase (GUS) in plants and its regulation in some model plants

The enzyme β-glucuronidase (GUS) is well characterized in animals and microbes. However, this enzyme is not well studied in plants and is widely assumed to be absent in them. In this study we document the ubiquitous presence of GUS in the model plants Arabidopsis thaliana, Oryza sativa, Nicotiana tabacum and Zea mays and record its expression pattern. The pH of the assay buffer was found to be critical with pH 4.0 being optimum for detection in all the species. GUS in plants appears to be associated with growth. In general, younger regions of the organs showed more GUS activity than the older and more mature tissues. In Brassica juncea roots stained for GUS, intense blue color could be seen in the trichoblast cells and the growing root hair cells as compared to the non-root hair forming epidermal cells or the fully elongated root hairs. Cotton fibers showed high GUS activity during the initial phase of elongation while the seed coat, from which the fibers formed, did not stain for GUS activity. The activity in the fibers disappeared after they were fully elongated. The level of GUS activity increased 2.58 folds in leaf tissues of N. tabacum when cultured in MS medium supplemented with 6-benzylaminopurine, while gibberellic acid enhanced GUS activity 2.9 folds in the inter-nodal regions of rice in 12-h treatment. In addition, elongation of stem, root and root hairs in tobacco seedlings was strongly inhibited by the specific inhibitor of GUS, saccharo-1-4-lactone in a reversible manner. Taken together, these evidences suggest a probable association of plant GUS in cell growth.Charu Sudan and Shiva Prakash, the first two authors, have contributed equally.     

Diversity and distribution of oribatid mites (Acari:Oribatida) in a lowland rain forest in Peru and in several environments of the Brazilians states of Amazonas, Rond?nia, Roraima and Pará.

We are summarizing the current state of knowledge of the diversity and distribution of oribatid mites in 26 environments in northern Brazil and of a rain forest in Peru. The published studies were mostly concentrated in Central Amazon. Only one report is a result from an agricultural polyculture. We are providing the first lists of species for savannas and for the Brazilian states of Roraima and Pará. Up to date, 146 species are definitively identified from a total of 444 taxa with 188 known genera, reinforcing the notion of a rich biodiverse area. The high number of 298 non-described species (morphospecies) clearly shows the inadequacy of the current taxonomic knowledge for the region. Most of the registers are from forest environments. In the soil from primary forests, we registered the highest diversity (54-155 species/morphospecies). Eighty-nine species were unique to primary forests, followed by 34 for savannas, 32 in trees, 10 in "igapó", 4 in caatinga, 3 in secondary forests, two in "várzea" and one in polyculture. Twenty genera were the most speciose. The species with the largest home ranges were Rostrozetes foveolatus, Scheloribates sp. A, and Galumna sp. A. Our numbers reflect the lack of taxonomists and show that the taxonomic knowledge must be improved for the region or we will continue to work with taxonomic resolution of Order or Family and a high percentage of morphospecies, which will probably be appropriate to the question being asked in each study, but not for a comparison among environments.     

Invasion success of alien plants: do habitat affinities in the native distribution range matter?

Aim To assess how habitat affinities in the native distribution range influence the invasion success of 282 central European neophytes (alien plants introduced after ad 1500). Location Czech Republic. Methods Classification trees were used to determine which native habitats donate the most alien species, the correspondence between habitats occupied by species in their native and invaded distribution ranges, and invasion success of species originating from different habitats. Results The species most likely to naturalize in Central Europe are those associated with thermophile woodland fringes in their native range (81%), cultivated areas of gardens and parks (75%) and broad‐leaved deciduous woodlands (72%). The largest proportions of invasive species recruit from those that occur on riverine terraces and eroded slopes, or grow in both deciduous woodland and riverine scrub. When the relative role of habitats in the native range is assessed as a determinant of the probability that a species will become invasive in concert with other factors (the species’ residence time, life history, region of origin), the direct effect of habitat is negligible. However, the effect of native habitats on patterns of invasions observed in central Europe is manifested by large differences in the numbers of species they supply to the invaded region. More than 50 neophytes were recruited from each of the following habitats: dry grasslands, ruderal habitats, deciduous woodland, inland cliffs, rock pavements and outcrops, and tall‐herb fringes and meadows. Main conclusions Casual species recruit from a wider range of habitats in their native range than they occupy in the invaded range; naturalized but not invasive species inhabit a comparable spectrum of habitats in both ranges, and successful invaders occupy a wider range of habitats in the invaded than in the native range. This supports the idea that the invasive phase of the process is associated with changes in biological features that allow for extension of the spectrum of habitats invaded.