The responses to shade of seedlings of very small-seeded tree and shrub species from tropical rain forest in Singapore

1. Newly germinated seedlings of six tree and shrub species with very small seeds (31–460 μg dry mass), one light-demanding and five shade-tolerant at the stage of establishment in the wild, were grown for 5 months in neutral shade houses with 0·5, 1, 3·5 and 7·5% daylight.
2. The ratio of yield in 7·5% to that in 1% was 8:1 for the light-demanding Melastoma malabathricum but only 2:1 for the confamilial shade-tolerator Pternandra echinata. The Pternandra, Urophyllum hirsutum, Ficus chartacea, Ficus grossularioides and Pellacalyx saccardianus showed a graded series of responses to irradiance, generally consistent with their apparent demands for light in the wild. In contrast, survival of very deep shade was not clearly related to light demand in nature.
3. The results support the conclusion drawn from observational studies that large seed size is not primarily adaptive in resisting shade but in resisting the associated risks of burial by litter, desiccation during dry spells, uprooting by birds and other kinds of damage by animals or falling debris.     

Amazonian caatinga and related communities at La Esmeralda, Venezuela: forest structure, physiognomy and floristics, and control by soil factors

Tall and short Amazonian caatinga at La Esmeralda (rainfall 2600 mm yr^–1) are described and compared with those at San Carlos de Río Negro (3600 mm yr^-1). The tall forests have much in common: thin-boled trees, many with scleromorphic mesophylls, lack of thick-stemmed climbers and of herbs, species paucity in all life-forms, domination by Eperua, and a high proportion of Bombacaceae, Clusiaceae and Euphorbiaceae. The short caatingas are also structurally and floristically similar but at La Esmeralda the notophyll scrubland species are lacking. At La Esmeralda, the common tree species are autochorous, but the majority of plants (particularly understorey species) are zoochorous. Relatively high transmission of light to the floor of the tall caatinga (1.5% PAR) may explain how trees with light-demanding architecture regenerate in the understorey. The presence of Cecropia and weedy species in the caatinga only at San Carlos may be the result of greater human interference there. The soils at San Carlos and La Esmeralda are similar: thick humus overlying bleached sand, waterlogged most of the year. The water table at La Esmeralda drops to >1 m depth during the drier months, but tensiometers at 30 cm depth did not indicate significant water shortage. Soils are consistently wetter in the short caatinga than the tall. Shortage of nitrogen appears to determine the major structural and physiognomic features of the caatinga, and depth of aerated soil appears to determine its stature. Comparisons are made between tall caatinga, wallaba and swamp forest in the Guianas and heath forests of Malaysia.Abbreviations PRF Palm-Rich Forest - SC Short Caatinga - TC Tall Caatinga     

Tree diversity in primary forest and different land use systems in Central Sulawesi,Indonesia

We studied the tree communities in primary forest and three different land use systems (forest gardens, ca. 5-year-old secondary forests, cacao plantations) at 900–1200 m elevation in the environs of Lore Lindu National Park, Central Sulawesi. The primary forests had ca. 150 tree species 10 cm diameter at breast height (dbh) per hectare, which is unusually high for forests at this elevation in southeast Asia. Basal area in the primary forest was 140 m² ha^–1, one of the highest values ever recorded in tropical forests worldwide. Tree species richness declined gradually from primary forest to forest gardens, secondary forests, and cacao plantations. This decline was paralleled by shifts in tree family composition, with Lauraceae, Meliaceae, and Euphorbiaceae being predominant in primary forests, Euphorbiaceae, Rubiaceae and Myristicaeae dominating in the forest gardens and Euphorbiaceae, Urticaceae, and Ulmaceae in the secondary forests. Cacao plantations were composed almost exclusively of cacao trees and two species of legume shade trees. Forest gardens further differed from primary forests by a much lower density of understorey trees, while secondary forests had fewer species of commercial interest. Comparative studies of birds and butterflies demonstrated parallel declines of species richness, showing the importance of trees in structuring tropical forest habitats and in providing resources.     

Ontogeny, understorey light interception and simulated carbon gain of juvenile rainforest evergreens differing in shade tolerance

Background and Aims

A long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried.

Methods

The biomass distribution of juveniles (17–740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain.

Results

An index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LAR_d = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey.

Conclusions

The results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.     

Fern richness,tree species surrogacy,and fragment complementarity in a Mexican tropical montane cloud forest

We related pteridophytes versus tree species composition to identify surrogate measures of diversity, and complementarity of seven cloud forest fragments. Forest structure, and fern and tree composition were determined in 70 (2 × 50 m) transects. Fern density (10,150–25,080 individuals/ha) differed among sites. We recorded 83 fern species in the transects. Nonparametric richness estimators indicated that more sampling effort was needed to complete fern inventories (14 more species). However, ferns recorded outside of the transects increased richness to 103 species (six more species than predicted). Twenty-eight species were unique and rare due to special habitat requirements (Diplazium expansum, Hymenophyllum hirsutum, Melpomene leptostoma, Terpsichore asplenifolia), or were at a geographical distribution edge (Diplazium plantaginifolium, Lycopodium thyoides, Pecluma consimilis, Polypodium puberulum). Correlations between fern richness and tree richness and density were not significant, but were significant between fern richness and fern density, between epiphytic fern density and tree richness and density. Tree richness is not a good surrogate for fern diversity. Only three species were recorded in all fragments (Polypodium lepidotrichum, P. longepinnulatum, P. plebeium); thus fragments pteridophytes compositions are highly complementary, but more similar for ferns than for trees. A regional conservation approach which includes many small reserves needs to focus supplementarity on patterns of tree and fern species richness.     

Carbohydrate storage,survival, and growth of two evergreen <Emphasis Type="Italic">Nothofagus</Emphasis> species in two contrasting light environments

A number of traits have been attributed important roles in tolerance of shade by plants. Some explanations emphasize traits enhancing net carbon gain; others emphasize energy conservation traits such as storage of non-structural carbohydrates (NSC). To date, cross-species studies have provided mixed support for the role of NSC storage in low-light survival. We examined NSC status, survival, biomass, and growth of large seedlings of two evergreen species of differing shade tolerance (Nothofagus nitida and N. dombeyi) grown in deep shade and 50% light for two growing seasons. We expected to find higher NSC concentration in the more shade-tolerant N. nitida and since allocation to storage involves sacrificing growth, higher growth rate in the shade-intolerant N. dombeyi. NSC concentration of both species was >twofold higher in 50% light than in deep shade, and in roots and stems did not differ significantly between species in either environment. NSC contents per plant were also similar between dead and living plants in deep shade. N. dombeyi outgrew N. nitida in 50% light, while this pattern was reversed in deep shade. Survival in deep shade was not correlated with NSC concentration. Leaf mass fraction was similar between species in 50% light, but lower in N. dombeyi in deep shade. Results provide little evidence of a link between carbohydrate storage and low-light survival in Nothofagus species, and support the view that understorey survival is primarily a function of net carbon gain. Patterns of variation in NSC concentration of the temperate species we studied are likely dominated by more important influences than adaptation to shade, such as limitation of growth or adaptation to cold stress.     

Altitudinal distribution of evergreen broad-leaved trees and their leaf-size pattern on a humid subtropical mountain,Mt. Emei,Sichuan, China

Altitudinal distribution of evergreen broad-leaved trees and changes in their leaf sizes were studied on a humid subtropical mountain, Mt. Emei (3099 m a.s.l., 29°34.5 N, 103°21.5 E), Sichuan, China. Among the total woody flora of ca. 540 species, evergreen broad-leaved trees account for 88 species in 39 genera and 23 families, corresponding to the northern limit of subtropical evergreen broad-leaved trees. The number of evergreen broad-leaved tree species greatly decreased from the low-altitudinal, evergreen broad-leaved forest zone (600–1500 m) to the mid-altitudinal, mixed forest zone (1500–2500 m), and to the high-altitudinal, coniferous forest zone (2500–3099 m). The overall trend of reduced leaf size toward upper zones was analyzed and documented in detail. The 88 species were assigned to three leaf-size classes: notophylls (48%), microphylls (36%), and mesophylls (16%). The leaf size was relatively small and the specific leaf weight (SLW, mg cm^–2) was much larger in high altitude as compared to low altitude. No overall correlation was found between leaf size and SLW, but leaf size decreased as SLW increased toward high altitude for certain species having relatively wide altitudinal ranges. Moreover, leaf size varied with forest stratification: canopy trees were predominantly notophyllous species, while subcanopy and understorey trees were mainly microphyllous species. The tendency is compatible with the trend found in other mountains of East Asia.     

Seasonal use by birds of stream-side riparian habitat in coniferous forest of northcentral British Columbia

To determine use of riparian habitats by birds in the northern coniferous forest of British Columbia, we censused birds and vegetation along 500 m transects placed parallel and perpendicular to three second-order streams. Censuses were conducted during spring, summer, autumn, and winter to investigate how use of riparian habitat changed seasonally. Stream-side riparian zones were characterized by a dense understorey of deciduous vegetation not found in the upslope forest. Nine bird species preferred the riparian understorey for breeding, six preferred it only during migration. Neotropical migrants (16 of 46 species) were more closely associated with stream-sides than year-round residents (11 species). Some breeding birds (five species) were significantly negatively associated with riparian habitats. The density of riparian birds declined with distance upstream but did not decline up to 250 m away from the stream. The more extensive riparian areas downstream supported a greater density of birds in all seasons compared to upstream areas, but more species only in spring and autumn. Species that nested in non-riparian areas in summer used riparian habitat in autumn, making riparian corridors in the northern coniferous forest important during migration. Maintaining both riparian and upslope habitats is necessary to preserve species diversity al the landscape level.     

Understorey gaps influence regeneration dynamics in subtropical coastal dune forest

Dominant understorey species influence forest dynamics by preventing tree regeneration at the seedling stage. We examined factors driving the spatial distribution of the monocarpic species Isoglossa woodii, a dominant understorey herb in coastal dune forests, and the effect that its cover has on forest regeneration. We used line transects to quantify the area of the forest understorey with I. woodii cover and with gaps in the cover. Paired experimental plots were established in semi-permanent understorey gaps with I. woodii naturally absent and in adjacent areas with I. woodii present to compare plant community composition, soil, and light availability between the two habitats. Isoglossa woodii was widespread, covering 65–95% of the understorey, while gaps covered the remaining 5–35% of the area. The spatial distribution of this species was strongly related to tree canopy structure, with I.␣woodii excluded from sites with dense tree cover. Seedling establishment was inhibited by low light availability (<1% of PAR) beneath I.␣woodii. When present, I. woodii reduced the density and species richness of tree seedlings. The tree seedling community beneath I. woodii represented a subset of the seedling community in gaps. Some species that were found in gaps did not occur beneath I. woodii at all. There were no significant differences between the sapling and canopy tree communities in areas with I. woodii gaps and cover. In the coastal dune forest system, seedling survival under I. woodii is dependent on a species’ shade tolerance, its ability to grow quickly during I. woodii dieback, and/or the capacity to regenerate by re-sprouting and multi-stemming. We propose a general conceptual model of forest regeneration dynamics in which the abundant understorey species, I. woodii, limits local tree seedling establishment and survival but gaps in the understorey maintain tree species diversity on a landscape scale.     

Loss of quantum yield in extremely low light

It has generally been assumed that the photosynthetic quantum yield of all C₃ plants is essentially the same for all unstressed leaves at the same temperature and CO₂ and O₂ concentrations. However, some recent work by H.C. Timm et al. (2002, Trees 16:47–62) has shown that quantum yield can be reduced for some time after leaves have been exposed to darkness. To investigate under what light conditions quantum yield can be reduced, we carried out a number of experiments on leaves of a partial-shade (unlit greenhouse)-grown Coleus blumei Benth. hybrid. We found that after leaves had been exposed to complete darkness, quantum yield was reduced by about 60%. Only very low light levels were needed for quantum yield to be fully restored, with 5 mol quanta m^–2 s^–1 being sufficient for 85% of the quantum yield of fully induced leaves to be achieved. Leaves regained higher quantum yields upon exposure to higher light levels with an estimated time constant of 130 s. It was concluded that the loss of quantum yield would be quantitatively important only for leaves growing in very dense understoreys where maximum light levels might not exceed 5 mol quanta m^–2 s^–1 even in the middle of the day. Most leaves, even in understorey conditions, do, however, experience light levels in excess of 5 mol quanta m^–2 s^–1 over periods where they obtain most of their carbon so that the loss of quantum yield would affect total carbon gain of those leaves only marginally.Abbreviations FBPase Fructose-1,6-bisphosphatase - RuBP Ribulose-1,5-bisphosphate - Rubisco RuBP carboxylase/oxygenase     

The functional morphology of juvenile plants tolerant of strong summer drought in shaded forest understories in southern Spain

It has been hypothesized that plants cannot tolerate combined shade and drought, as a result of morphological trade-offs. However, numerous plant species are reportedly widespread in shaded forest understories that face drought, whether seasonal or occasional. We studied juveniles of six plant species that cope with strong summer drought in the understoreys of mixed Quercus forests in southern Spain: the tall-shrubs Phillyrea latifolia and Viburnum tinus, the perennial herb Rubia peregrina, the small shrub Ruscus aculeatus, and climbers Hedera helix and Smilax aspera. All of these species persist in evergreen shade (c. 3% daylight). Two other species were studied as comparators, Ruscus hypoglossum, less tolerant of drought, and Ceratonia siliqua, less tolerant of shade. Morphological and chemical variables relevant to shade and drought tolerance were measured for juveniles in a range of sizes, and also for the leaves of mature plants. The species converge in features that confer tolerance of shade plus drought by reducing demand for resources. Demand for water is reduced through a moderate to high below-ground mass fraction and low to moderate specific leaf area (respectively 0.22–0.52 and 112–172 cm² g^–1 at 1.00 g total dry mass). Demand for both irradiance and water is reduced through a low to moderate foliar nitrogen concentration and long-lived, physically protected leaves (2 yr). The species also converge in features that confer tolerance of either low irradiance or drought through specialized capture of resource, without precluding the other tolerance. These features include deep roots relative to shoot size, moderately higher specific leaf area in shade (1.2–2.0 × that in sun) and higher chlorophyll:nitrogen ratio in shade. Foliar chlorophyll per unit mass was higher in shade, but chlorophyll was not necessarily synthesized in greater amounts; rather, it was higher apparently due to shade effects on structural features linked with specific leaf area. In contrast, N per unit mass was higher in sun leaves independently of specific leaf area. Despite these convergences, the species diverge considerably in their root mass allocation and architecture, leaf saturated water content, density of stomata and guard cell size. No single narrowly defined functional type is needed for tolerance of shade plus drought.     

The influence of shading on associative N2 fixation

Summary The effect of reduced solar radiation on associative N₂-fixation and plant parameters was studied in three field experiments (1978–80). Gahi-3 pearl millet (Pennisetum americanum (L.) K. Monch.) field plots were shaded with saran shade cloth that reduced solar radiation by 50% and 75%. Acetylene reduction activity (ARA) was reduced by shading in one of the three experiments. The two non-responding experiments were conducted on a wall-drained, low-activity site (ARA means ranging 17–68 n moles ethylene core^–1 h^–1), the responding experiment was conducted on a poorly drained, high-ARA site.Shading affected the plants drastically, reducing fresh weight and dry matter yields up to 46% (50% shade) and 57% (75% shade). Shading also reduced dry matter percentage from 19.6 (no shade) to 15.3 (75% shade) and increased nitrogen content from 0.6% (no shade) to 1.53% (75% shading). However, shading did not affect protein yield. Inoculation withAzospirillum brasilense had no measurable effect on yield or acetylene reduction in the first two experiments.In the third experiment, shading reduced mean ARA of inoculated plots over 100% but had no significant effect on control plots. Inoculation significantly increased ARA in the nonshaded plots but not in shaded plots. Acetylene reduction activity was high, with means ranging between 208 and 465 n moles ethylene evolved core^–1 h^–1. Soil moisture and millet growth stage also affected acetylene reduction activity.     

Establishment of a plant regeneration system for wheatgrasses (Thinopyrum,Agropyron and Pascopyrum)

Wheatgrasses (Thinopyrum, Agropyron and Pascopyrum spp.) are a ubiquitous group of cool-season grasses used throughout much of the semiarid temperate world. In order to explore the potential of biotechnology to accelerate conventional breeding efforts, we developed an efficient plant regeneration system for different wheatgrass species: tall wheatgrass [Thinopyrum ponticum (Podp.) Liu and Wang], intermediate and pubescent wheatgrass [Thinopyrum intermedium (Host) Barkw. and D.R. Dewey], crested wheatgrass [Agropyron cristatum (L.) Gaertner], and western wheatgrass [Pascopyrum smithii (Rydb.) Löve]. Embryogenic callus was induced from mature and immature embryos and immature inflorescence with an induction frequency in the range of 0.5–8.3% for the different wheatgrass species tested. Individual embryogenic calluses were then used to establish single genotype-derived suspension cultures. Efficient plant regeneration was achieved from the established em-bryogenic suspensions with regeneration frequencies in the range of 20–65% for tall wheatgrass, 21–40% for intermediate and pubescent wheatgrasses, 32–51% for crested wheatgrass, and 25–48% for western wheatgrass. The cell suspension-derived wheatgrass plants were fertile and phenotypically normal in the field. The efficient plant regeneration system provides a solid basis for genetic transformation of wheatgrasses.     

Photosynthetic Response to Irradiance in Valeriana jatamansi Jones,a Threatened Understorey Medicinal Herb of Western Himalaya

Net photosynthetic rate (P _N) of Valeriana jatamansi plants, grown under nylon net shade or under different tree canopies, was saturated with photons at 1 000 mol m^–2 s^–1 photosynthetic photon-flux-density (PPFD), whereas open-grown plants were able to photosynthesise even at higher PPFD, e.g. of 2 000 mol m^–2 s^–1. Plants grown under net shade had higher total chlorophyll (Chl) content per unit area of leaf surface. However, Chl a/b ratio was maximal in open-grown plants, but remained unchanged in plants grown in nylon net shade and under different tree canopies. Sun-grown plants had thicker leaves (higher leaf mass per leaf area unit), higher wax content, and higher P _N than shade grown plants. Thus V. jatamansi is able to acclimate to high PPFD and therefore this Himalayan species may be cultivated in open habitat to meet the ever-increasing industrial demand.     

Vascular plant species count in a wet forest in the Chocó area on the Pacific coast of Colombia

The total number of vascular plant species was counted and growth form distribution was studied in the Chocó area on the Pacific coast of Colombia, in two transects 400×10m and ten transects 2×50m, for a total sampled area of 0.9ha. The species count of the ten transects (442 species in 0.1ha) appears to be the highest number of species recorded with this methodology. There were 970 species for the total area (0.9ha). Ninety to ninety-five percent of the species were under 10cm dbh and 70–86% under 2.5cm dbh, epiphytes and small trees and treelets 10cm dbh being the most diverse growth forms. The most species-rich families and genera were those represented by herbaceous plants and treelets. Individuals were counted only in the ten 2 × 50 m transects (0.1 ha), where 4459 individuals were found. Palms and ferns were the most abundant growth forms. Arguments are presented against the way diversity is usually measured. Recommendations are made to include other growth forms besides trees when assessing alpha diversity for conservation purposes.     

Experimental defaunation of terrestrial mammalian herbivores alters tropical rainforest understorey diversity

It has been suggested that tropical defaunation may unleash community-wide cascading effects, leading to reductions in plant diversity. However, experimental evidence establishing cause–effect relationships thereof is poor. Through a 5 year exclosure experiment, we tested the hypothesis that mammalian defaunation affects tree seedling/sapling community dynamics leading to reductions in understorey plant diversity. We established plot triplets (n = 25) representing three defaunation contexts: terrestrial-mammal exclosure (TE), medium/large mammal exclosure (PE) and open access controls (C). Seedlings/saplings 30–100 cm tall were marked and identified within each of these plots and re-censused three times to record survival and recruitment. In the periods 2010–2011 and 2011–2013, survival was greater in PE than in C plots and recruitment was higher in TE plots than in C plots. Overall, seedling density increased by 61% in TE plots and 23% in PE plots, whereas it decreased by 5% in C plots. Common species highly consumed by mammals (e.g. Brosimum alicastrum and Ampelocera hottlei) increased in their abundance in TE plots. Rarefaction curves showed that species diversity decreased in TE plots from 2008 to 2013, whereas it remained similar for C plots. Given the prevalence of tropical defaunation, we posit this is an anthropogenic effect threatening the maintenance of tropical forest diversity.     

In situ responses to elevated CO2 in tropical forest understorey plants

1. Plants growing in deep shade and high temperature, such as in the understorey of humid tropical forests, have been predicted to be particularly sensitive to rising atmospheric CO₂. We tested this hypothesis in five species whose microhabitat quantum flux density (QFD) was documented as a covariable. After 7 (tree seedlings of Tachigalia versicolor and Beilschmiedia pendula ) and 18 months (shrubs Piper cordulatum and Psychotria limonensis, and grass Pharus latifolius ) of elevated CO₂ treatment ( c. 700 μl litre^–1) under mean QFD of less than 11 μmol m^–2 s^–1, all species produced more biomass (25–76%) under elevated CO₂.
2. Total plant biomass tended to increase with microhabitat QFD (daytime means varying from 5 to 11μmol m^–2 s^–1) but the relative stimulation by elevated CO₂ was higher at low QFD except in Pharus .
3. Non-structural carbohydrate concentrations in leaves increased significantly in Pharus (+ 27%) and Tachigalia (+ 40%).
4. The data support the hypothesis that tropical plants growing near the photosynthetic light compensation point are responsive to elevated CO₂. An improved plant carbon balance in deep shade is likely to influence understorey plant recruitment and competition as atmospheric CO₂ continues to rise.     

Nitrogen mineralisation in podzol soils under boreal Scots pine and Norway spruce stands

N mineralisation was investigated in the mor humus layer of a podzol at a forested catchment area of Saarejärve Lake in Eastern Estonia. The investigated areas were pine (Rhodococcumunderstorey) and spruce (Vaccinium understorey) stands, which are permanent sample plots of an integrated monitoring network. The seasonal pattern of net N mineralisation was studied by incubating undisturbed cores of mor humus (0–8 cm) in buried polyethylene bags in situ. Samples were collected and incubated between July 1996 and April 1998. The period of incubation was approximately 1 month, except for wintertime when incubation lasted till thawing of ground (5 months). The amounts of mineral nitrogen formed during monthly incubations in vegetation period vary considerably (0.4–8.7 kg ha^–1). About 70% of the variation of net ammonification could be explained by environmental factors - temperature, initial moisture and pH. Ammonium was the dominant form of mineral nitrogen, which is typical for mor humus. The rate of nitrification was very low, and most of the annual net nitrification occurred during just one or two months (May–June, October) depending on site and year. Measured annual net N mineralisation was 29.2 kg ha^–1 for the spruce stand and 23.6 kg ha^–1 for the pine stand. These measures were found to be in good accordance with other N-fluxes in the ecosystem.     

Overstorey Control of Understorey Species Composition in a Near-natural Temperate Broadleaved Forest in Denmark

Little is known about the importance of the forest overstorey relative to other factors in controlling the spatial variability in understorey species composition in near-natural temperate broadleaved forests. We addressed this question for the 19 ha ancient forest Suserup Skov (55°22′ N, 11°34′ E) in Denmark, one of the few old-growth temperate broadleaved forest remnants in north-western Europe, by inventorying understorey species composition and environmental conditions in 163 100 m² plots. We use unconstrained and constrained ordinations, variation partitioning, and Indicator Species Analysis to provide a quantitative assessment of the importance of the forest overstorey in controlling understorey species composition. Comparison of the gradients extracted by unconstrained and constrained ordinations showed that the main gradients in understorey species composition in our old-growth temperate broadleaved forest remnant are not caused by variability in the forest overstorey, but are related to topography and soil, edge effects, and unknown broad-scale factors. Nevertheless, overstorey-related variables uniquely accounted for 15% of the total explained variation in understorey species composition, with the pure overstorey-related (R_po), topography and soil (R_pt), edge and anthropogenic disturbance effects (R_pa), and spatial (R_ps) variation fractions being of equal magnitude. The forward variable selection showed that among the overstorey-related variables understorey light availability and to a lesser extent vertical forest structure were most important for understorey species composition. No unique influence of overstorey tree species identity could be documented. There were many indicator species for high understorey light levels and canopy gap centres, but none for medium or low light or closed canopy. Hence, no understorey species behaved as obligate shade plants. Our study shows that, the forest overstorey has a weak control of understorey species composition in near-natural broadleaved forest, in contrast to results from natural and managed forests comprising both conifer and broadleaved species. Nevertheless, >20% of the understorey species found were indicators of high light conditions or canopy openings. Hence, variability in canopy structure and understorey light availability is important for maintaining understorey species diversity.     

Run-on contribution to a Sahelian two-phase mosaic system: Soil water regime and vegetation life cycles

An experiment was carried out from 1992 to 1995, in south-western Niger on a banded vegetation pattern which dominates on a laterite-capped plateau in the region. We quantified the changes in infiltration and vegetation in a thicket from which run-on from the upslope bare soil zone was artificially divested. A concrete wall (40 m long, 60 cm high, 20 cm thick, with a foundation 25 cm deep) was constructed at its upslope boundary. Infiltration was measured to a depth of 5.4 m by a neutron probe, and densities of annual plants were monitored along transects crossing perpendicularly a control thicket and the thicket deprived of run-on. Phenological phases and leaf water potential of the two dominant shrub species were recorded from stratified sampling according to their preferred location along the water resource gradient. Results indicated that run-on contributed the most to infiltration in the central zone, but the water content available to the annual plants (layer 0–10 cm) was not affected by run-on deprivation. Significant differences were found in the water content available to the shrubs (layer 0–100 cm) both between zones (upslope and central), and between thickets after the wall was built. However, in the thicket deprived of run-on, life cycle and physiology of the shrubs were severely disturbed upslope, while much smaller effects were observed in the centre. Surprisingly, within thestudy interval, run-on contribution was not found to be as essential to shrubs' life cycle at the location where it contributed the most to the infiltration.