The response of forest floor vegetation and tree regeneration to deer exclusion and disturbance in a riparian deciduous forest,central Japan

The response of forest floor vegetation and regeneration of major treespecies to deer exclusion in a riparian forest were studied for 3 years withtheinteractive effects of natural disturbances. At the start of this study, deerdensity had quickly increased to a fairly high level (29–31 individualskm^–2) during the last decade and had severely reduced theamount of forest floor vegetation in the study area. Dwarf bamboos, which weredominant before, had almost diminished from the forest floor. During the studyperiod, aboveground biomass increased steadily but species diversity did notchange much in the exclosures. Outside the exclosures, the seedlings of alltreespecies were damaged greatly by deer browsing, especially the taller ones. Deerbrowsing had little effect on the emergence of tree seedlings, but deertrampling might have accelerated emergence indirectly by disturbing the soilsurface for some species. Differences in plant responses were also observed fordeer browsing and the presence of dwarf bamboo that strongly inhibits therecruitment of tree seedlings. The nine major species were classified intothreegroups according to the response of seedlings to deer browsing and disturbance.Group 1 consists of the species whose seedling survival is affected bybrowsing,but seldom by disturbances (Phellodendron amurense,Kalopanax pictus, Quercus crispulaandMalus toringo). Groups 2 and 3 consist of species adaptedto tree-fall disturbance (Betula spp.) and ripariandisturbance (Alnus hirsuta var.sibirica, Ulmus davidiana var.japonica, Populus maximowiczii andSalix sachalinensis), respectively, and seedling survivalof these two groups is principally affected by light conditions. The effect ofdeer browsing on seedling survival and growth was greater for Group 2 than forGroup 3.     

Infestation of the coccid,Icerya seychellarum (Westw.), on the mangrove Avicennia marina (forsk.) vierh. on Aldabra Atoll,with special reference to tree age

Summary The level of infestation by the coccid Icerya seychellarum on the mangrove tree Avicennia marina was measured on a stand of ca. 670 trees on Aldabra Atoll in 1978. Trees varied in basal trunk diameter (btd) between 3 and 30 cm. Leaf infestation increased significantly with increasing btd-class and young trees (btd 3.0–4.5 cm) had a mean infestation of 0.3 mg coccids g^-1 leaf whilst the heaviest infested trees (btd 13.0–14.5 cm) supported 7.6 mg g^-1. Leaf and shoot growth were not correlated with coccid infestation but shoot vigour (leaf to green shoot ratio) declined significantly with increasing btd-class (or age). Leaves of older, heavily infested trees did not differ from those of young uninfested trees in the concentration of soluble or total nitrogen of in four other leaf elements, indicating that nutrient status was not an important factor in infestation. Field observations showed that many of the older trees were subjected to poor drainage which may have increased their susceptibility to attack and also that on the leaves of young trees salt secretion presented an effective barrier to the establishment of coccids. re19800212     

Differential seed and seedling predation by crabs: impacts on tropical coastal forest composition

Recently, the importance of seed predation by crabs on mangrove species distributions and densities has been established by several studies. In a tropical coastal terrestrial forest in Costa Rica, we investigated the relative importance of predation by land crabs, Gecarcinus quadratus, and hermit crabs, Coenobita compressus, on measured forest composition through a series of seed removal and seedling establishment experiments. We also used natural light-gaps and adjacent non-gap sites to test how canopy cover affects crab predation (seed removal) and seedling establishment. We found fewer tree species (S=18) and lower densities (seedlings, saplings, and adults) in the coastal zone within 100 m of coastline, than in the inland zone (S=59). Land crab densities were higher in the coastal zone (3.03±1.44 crabs m^–2) than in the inland zone (0.76±0.78 crabs m^–2), and hermit crabs were not present in the inland zone. Seed removal and seedling mortality also were higher in the coastal zone than in the inland zone, and in the open controls than in the crab exclosures. Mortality of seeds and seedlings was two to six times higher in the controls than exclosures for four of the five experiments. Crabs preferred seeds and younger seedlings over older seedlings but showed no species preferences in the seed (Anacardium excelsum, Enterolobium cyclocarpum, and Terminalia oblonga) and seedling (Pachira quinata and E. cyclocarpum) stages. We conclude that the observed differences in tree densities were caused by differential crab predation pressure along the coastal gradient, while the differences in species composition were due to predator escape (satiation) by seed quantity. Canopy cover did not affect seed removal rates, but did affect seedling survival with higher mortality in the non-gap versus gap environments. In summary, crab predation of seeds and seedlings, and secondarily canopy cover, are important factors affecting tree establishment in terrestrial coastal forests.     

Differential establishment and survival of species in deciduous and evergreen shrub patches and on bare ground,Mt. Koma,Hokkaido, Japan

The effects of patches formed by a deciduous shrub (Salix reinii) and an evergreen shrub (Gaultheria miqueliana) on cohabitants were examined on an active volcano Mt. Koma, northern Japan, using bareground as a comparative baseline (control). Species richness increased with increasing the areas of three habitats (Salix patches, Gaultheria patches and bareground), but significantly slowed in Gaultheria patches where the shoot densities of cohabitants also decreased. Calamagrostis hakonensis and Spiranthes sinensis were 2–3 times more likely to establish in Salix patches than in bareground, Anaphalis margaritacea var. angustior and Campanula lasiocarpa were 4–5 times more likely to establish on bareground than in Salix patches. Cohabitants was fewer with Gaultheria. Light intensities of the two patch types were ca 1/3 of that on bareground. Light intensity on Gaultheria was ca 88% of that on Salix patch and decreased with increasing patch size, because the differences in stem and leaf architectures and phenology between the two patch-forming species. Moisture and total nitrogen in volcanic deposits were 2–3 times higher in the patches than in bareground. In the patches, low light intensity restricted seed germination, in particular, for the plants characteristic of bareground, e.g., Anaphalis, while high soil moisture increased the survival rates of Calamagrostis. In Gaultheria patches, despite high moisture, the reduced light intensity influenced seed germination and inhibited the establishment of most species. Seedbanks estimated by a lab-experiment indicated that Salix patches trapped more seeds. As a consequence, Salix patches act as facilitator, in particular, for Calamagrostis, by improving moisture and nutrients status of volcanic deposits, moderating light intensity and seed trapping. Deciduous shrub patches, such as Salix reinii, have an improving role in accelerating rates of succession in harsh environments, while evergreen shrub patches slow plant establishment. Nomenclature: Ohwi and Kitagawa (1983)     

Altitudinal change in soil and foliar nutrient concentrations and in microclimate across the tree line on the subtropical island mountain Mt. Teide (Canary Islands)

Mt. Teide (Tenerife, Spain) is a high volcanic island mountain with an unusually low tree line elevation (2000–2100 m). While searching for the causes of this tree line depression, we analysed the concentrations of total N, available P, and salt-exchangeable Ca, K and Mg in three soil horizons, and the foliar N, P, Ca, K and Mg concentrations in six abundant plant species (trees, shrubs, forbs) along a transect from 1400 (1600) to 3100 m a.s.l. The objective of the study was to detect altitudinal trends in soil and plant nutrient status below and above the tree line. For characterising elevational changes in microclimate and hydrology, we also conducted measurements of air and soil temperatures (T_a and T_s), atmospheric water vapour saturation deficit (D), potential evaporation (E) and soil moisture (θ) on 3–4 occasions along the transect.T_a and T_s did not linearly decrease with elevation but were highest at or immediately below the tree line and were relatively low in the closed Pinus canariensis forest at 1600 m. T_s reached maxima at about 70 °C near the tree line. The absence of a linear temperature decrease with elevation was caused by a stable temperature inversion at about 2000 m and by canopy shading in the forest canopy below the tree line. In June, the topsoil (10 cm) dried out almost completely between 1800 and 3100 m, but remained moist in the pine forest at 1600 m. This gradient reflects the transition from the montane cloud belt to the dry alpine belt higher upslope. The subsoil (30 cm) contained >30 vol% of soil water at all elevations even in June except for the uppermost site (3100 m). Potential evaporation increased with elevation despite a decrease in D. We assume that this is mainly due to the air pressure-dependent increase with elevation in the diffusion coefficient for water vapour in air.The concentrations of N and ‘available’ P (after Olsen) in the mineral topsoil were by far smaller than in alpine soils of other humid mountains which is thought to be a consequence of a very dry and biologically inactive topsoil in the semi-arid alpine belt of Mt. Teide. In contrast to many other mountains, foliar N, P and cation concentrations in the plants did not increase with elevation but either remained unchanged, or decreased as in the tree line species P. canariensis. Nevertheless, P. canariensis probably is not limited by nutrient deficiency at the tree line despite rather low N and P needle concentrations. Rather, drought and heat stress effects on seedling establishment are thought to be the causes of the tree line depression.     

Tree regeneration patterns on Mount Usu, northern Japan, since the 1977–78 eruptions

Although the summit of Mount Usu was deforested by the 1977–78 eruptions, vegetative regeneration on the caldera rim was rapid due to the erosion of thick volcanic deposits by snow and rain. To obtain the mechanisms underlying regeneration patterns after the eruptions, we monitored the growth of permanently-marked stems from 1983 to 1990. Regeneration was from resprouting-branches buried in the volcanic deposits on the caldera rim, while on the crater basin, where thick volcanic deposits accumulated, regeneration was from seedlings. The seedling regeneration lagged approximately 3 years behind vegetative regeneration. Stem densities averaged 14,000 ha^-1 in the vegetatively-regenerated community on the caldera rim, and 28,000 ha^-1 in the seedling regeneration on the crater basin. Populus maximowiczii accounted for ca. 75% of total stems on the caldera rim, while P. maximowiczii accounted for ca. 30% on the crater basin where Salix integra and Betula platyphylla var. japonica were also common. In both stands, immigration and mortality rates were very low. The growth of vegetatively regenerated stems expressed as stem height and diameter was significantly faster than that of stems grown from seedlings. Herbivory damage on the terminal shoots of tall stems was restricted on the caldera rim and was restricted for B. platyphylla var. japonica on the crater basin, perhaps due to fast growth supporting herbivore avoidance or low palatability. Height growth was restricted when neighbors established in close proximity, especially in the seedling-regenerated forest. The results suggest that vegetative regeneration is rapid due to three mechanisms: 1) faster plant growth; 2) herbivore avoidance; and 3) decreased interference by neighboring.     

Eucalyptus recruitment in degraded woodlands: no benefit from elevated soil fertility

Grasses and forbs compete heavily with young tree seedlings for available resources, greatly reducing tree seedling establishment success. Soil nutrient enrichment associated with agricultural intensification can increase the growth of both herbaceous and woody lifeforms growing in isolation, but may change the balance of competitive advantage when growing together. The effects of nitrogen and phosphorus enrichment on pasture biomass and competition with two Australian grassy woodland trees (Eucalyptus albens and Eucalyptus microcarpa) was investigated in a field plot trial. Soil nutrients increased pasture biomass, but had no measurable effect on tree growth in our experiment. Competition from pasture species, even at low levels, led to high tree seedling mortality and greatly reduced tree seedling growth compared with pasture-free plots. However, when pasture-free plots were excluded from the analysis, tree seedling leaf area was not strongly correlated with herbaceous biomass. Tree seedling establishment was severely restricted even at the lowest levels of pasture biomass. We conclude that increased soil fertility resulted in a competitive advantage to the pasture, and does not improve tree seedling establishment when grown either with or without exotic herbaceous pasture (grassy understorey) species.     

Diurnal cycles of rhizosphere acidification byPinus contorta seedlings

The pH of the nutrient solution bathing the roots of four-month-oldPinus contorta var.latifolia Englm. seedlings was monitored continuously between additions of nutrients. Nitrogen was supplied in the form of NH₄NO₃, and was added three times per week in amounts relative to seedling fresh weight. No pH change was associated with the nutrient addition cycle; however, extinguishing of the lights at night resulted in a decrease in pH of almost half a pH unit in the first hour. The pH reverted to normal within a few hours. Re-illumination resulted in a pH increase of a smaller magnitude, but over a similar time span. Estimation of the proton extrusion rate gave values of about 17 µmol (g FW root)^–1 h^–1.     

Combined effects of seed and soil quality drive seedling performance of a late‐successional canopy tree in a tropical forest

Habitat loss and fragmentation affect the structure and functioning of forested ecosystems worldwide, yet we lack an understanding of how species respond to environmental changes. Here, we examined reproductive success and seedling performance of Poulsenia armata (Moraceae) in continuous and fragmented forests of Los Tuxtlas, southern Mexico. We further investigated how maternal habitat and soil conditions manifested in the seedling stage. We determined seed quality and seedling performance by combining isotopic analyses in seed quality with field observations of P. armata fruit production and a common‐‐garden experiment. Soil conditions in forest fragments negatively impacted P. armata reproductive success. Trees of P. armata in forest fragments were smaller in size and produced fewer fruits and smaller seeds with lower quality compared with trees from the continuous forest. The combined effects of maternal habitat and soil conditions determined seedling survival and growth of this tropical tree. Notably, seedlings had restricted plasticity for biomass allocation to roots, limiting the capacity of fragmented populations to compensate for the initial low N content in seeds. Trees in forest fragments at Los Tuxtlas produced offspring competitively inferior and potentially less resilient than counterparts in continuous forest, jeopardizing future persistence of this late‐successional tree species.     

Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan

To evaluate the effects of canopy gaps and forest floor microsites (soil, fallen logs, root-mounds, buttresses and stumps) on regeneration of subalpine forests, the gap regeneration and seedling occurrence of conifers (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis and Tsuga diversifolia) were studied in two stands of a subalpine old-growth forest, central Japan. The percentage of gap area to total surveyed area was 11.2–11.3% in the stands. Gap regeneration was not common for P. jezoensis var. hondoensis and T. diversifolia. In contrast, gap regeneration by advanced regeneration was relatively common for Abies. Seedling occurrence of P. jezoensis var. hondoensis and T. diversifolia was restricted on elevated surfaces such as stumps and root-mounds, while Abies seedlings could occur on soil as well as on elevated surfaces. Rotten stumps were the most favorable microsites for conifer seedling occurrence, which covered small area in the forest floor. Although canopy gaps were not always favorable for seedling occurrence, all conifer seedlings were larger under canopy gaps than under closed canopy. Canopy gaps and forest floor microsites clearly affected seedling occurrence and growth of conifers. This suggests that regeneration of conifers is related to the difference of growth advantage under canopy gaps and favorable microsites for seedling occurrence.     

Distribution of different mycorrhizal classes on Mount Koma, northern Japan

To investigate the role of mycorrhizae in nutrient-poor primary successional volcanic ecosystems, we surveyed mycorrhizal frequencies on the volcano Mount Koma (42°04N, 140°42E, 1,140 m elevation) in northern Japan. After the 1929 eruptions, plant community development started at the base of the volcano. Ammonia and nitrate levels, along with plant cover, decreased with increasing elevation, whereas phosphorus did not. In total, 305 individuals of 56 seed plant species were investigated in three elevational zones (550–600 m, 650–700 m, and 750–800 m). Five mycorrhizal classes were classified based on morphological traits: ecto- (ECM), arbuscular (AM), arbutoid, ericoid, and orchid mycorrhiza. All plant species were mycorrhizal to at least some extent, with most widespread tree species being heavily ectomycorrhizal. In addition, of 16 tree species collected in all three zones, 6 differed in the frequencies of ECM on roots between elevational zones, and 3 of these 6 species increased in frequency with increasing elevation. These results suggest that ECM colonization in some tree species is related to establishment in nutrient-poor habitats. All species of Ericaceae and Pyrolaceae had ericoid mycorrhizae, and an Orchidaceae species had orchid mycorrhizae. Herbaceous species, except for the low mycorrhizal frequency of Carex oxyandra and two Polygonaceae species, and ericoid and orchid mycorrhizal species, were generally AM. Of herbaceous species, Anaphalis margaritacea var. angustior increased AM frequency and decreased ECM frequency with increasing elevation, and Hieracium umbellatum increased ECM frequency. In total, the establishment of herbaceous species was not sufficiently explained by AM colonization on roots. Tree individuals developed 2–3 classes of mycorrhizae more than herbs at each elevational zone. We conclude that the symbiosis between seed plants and mycorrhizae, ECM in particular, greatly influences plant community structures on Mount Koma. Not only a single mycorrhizal class, but combinations of mycorrhizal classes should be studied to clarify effects on plant community dynamics.     

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.     

Relative contributions of sexual and asexual regeneration strategies in Populus nigra and Salix alba during the first years of establishment on a braided gravel bed river

Populus nigra L. var betulifolia and Salix alba L. var alba are early successional riparian tree species threatened throughout Continental Europe by significant changes to the natural physical processes governing their natural habitat – geomorphologically active floodplains. River management activities have dramatically altered natural patterns of river flow and rates of sediment delivery along rivers, with possible consequences for the balance between sexual and asexual regeneration strategies in these species. Conservation strategies will benefit from a greater understanding of the ways in which dynamic physical processes on the floodplain influence sexual and asexual recruitment. This paper describes a field survey investigating the relative abundance and spatial distributions of P. nigra and S. alba sexual and asexual recruits during the first years of establishment along a braided gravel bed river. Sexual and asexual recruits were identified by excavation along transects in a wet and a dry field season and distributional differences were described in terms of elevation on the floodplain, local sediment type and exposure to floodwaters. Regeneration was overwhelmingly from seed in the first 2–3 years following recruitment, but poor survival rates among sexual recruits saw a shift in the relative abundance of regeneration strategies over time. In relating hydrological data to recruitment, unseasonal flood disturbances had a negative effect on recruitment from seed and a positive effect on vegetative regeneration. Seedlings were associated with fine sediment deposits and were restricted primarily to low elevations on the flood plain, while asexual recruits had a wider spatial distribution. Certain microsite types were unique to either regeneration strategy.     

Survival,growth, and photosynthesis of tree seedlings competing with herbaceous vegetation along a water-light-nitrogen gradient

In herbaceous dominated patches and ecosystems, tree establishment is influenced partly by the ability of woody seedlings to survive and grow in direct competition with herbaceous vegetation. We studied the importance of season long wet and dry spells on the competitive interactions between herbaceous vegetation and oak seedlings along a light and nitrogen gradient in an infertile secondary successional grassland in central North America. We conducted a field experiment in which seedlings of bur oak (Quercus macrocarpa) and northern pin oak (Q. ellipsoidalis) were exposed to two levels of light (full sun and 80% shade), three levels of nitrogen input (0, 5, 15 g m^–1 yr^–1), and three levels of water input (low, medium and high). In addition, seedlings were grown with and without the presence of surrounding herbaceous vegetation under both light and all three water levels. Seedling survival, growth, and rate of photosynthesis were significantly affected by competition with herbaceous vegetation and these effects varied along the multiple resource gradient. Overall, seedling survival of both species was significantly greater in wetter and shaded plots and when surrounding herbaceous vegetation was removed and was lower in nitrogen enriched plots. We found that soil water was significantly affected by varying inputs of water, light, and the presence or absence of herbaceous vegetation, and that seedling survival and rate of photosynthesis were highly correlated with available soil water. Our findings show that the impact of season long wet and dry spells on tree seedling success in grasslands can be affected by light and soil nitrogen availability.     

Experimental Native Tree Seedling Establishment for the Restoration of a Mexican Cloud Forest

The cloud forests of Mexico have been degraded and severely fragmented, and urgently require restoration. However, progress with restoration has been constrained by a lack of information concerning the seedling ecology of native tree species. An experiment was therefore conducted to assess the influence of different environmental factors on the seedling survival and growth of four native tree species (Fagus grandifolia var. mexicana, Carpinus caroliniana, Symplocos coccinea, and Quercus acutifolia). The seedlings were established on three sites, in two contrasting environments: inside forest fragments and on adjacent agricultural land. Highly significant differences were recorded in seedling survival and growth among sites, environments, species, and interactions between these factors. Highest survival was recorded for Quercus, which uniquely among the four species displayed the same survival percentage inside and outside the forest. Survival of the other species was higher inside the forest. In contrast, growth rates of all four species were higher outside the forest. The most important cause of mortality outside the forest was desiccation, although significant seedling predation was also observed on two sites. Results indicate that all four species can be established successfully both within forest fragments and in neighboring agricultural areas, emphasizing the scope for forest restoration. However, the interactions observed between species, sites, and environments highlight the importance of accurate species–site matching if optimum rates of growth and survival are to be obtained. Quercus spp. have great potential for establishment on agricultural sites.     

Effects of arbuscular mycorrhizal fungi and soil developmental stages on herbaceous plants growing in the early stage of primary succession on Mount Fuji

A pot culture experiment was conducted to examine the effects of arbuscular mycorrhizal (AM) fungi and soil developmental stages on the growth and nutrient absorption of pioneer plants growing in the early stage of primary succession on Mt. Fuji. Four herbaceous plants, Polygonum cuspidatum (Polygonaceae), Miscanthus oligostachyus (Gramineae), Aster ageratoides var. ovatus (Compositae), and Hedysarum vicioides (Leguminosae), were grown from seed in soils collected from two different successional stages, bare ground and an herbaceous plant community. Spores of indigenous AM fungi collected from the herbaceous plant community were used as inoculum. The initial colonizer P. cuspidatum showed very low levels of AM colonization (<0.4%), whereas the average AM colonization levels of M. oligostachyus, A. ageratoides var. ovatus, and H. vicioides were within the range of 13–49%. AM fungi had positive effects on the growth and N acquisition of the leguminous species (H. vicioides) irrespective of soil developmental stages. In contrast, AM colonization did not increase the plant dry weight and N content of the non-leguminous species (P. cuspidatum, M. oligostachyus, and A. ageratoides var. ovatus) in both soil developmental stages. A positive effect of AM colonization on P content was observed in M. oligostachyus, A. ageratoides var. ovatus, and H. vicioides only in soil collected from the herbaceous plant community. P. cuspidatum showed no or a negative response to AM colonization in all cases. These results suggest that the effect of AM fungi on plant growth depends more on the plant species than soil developmental stages in the early stage of primary succession in this volcanic area.     

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.     

Lead tolerance of Betula and Salix in the mining area of Mechernich/Germany

Natural populations of woody perennials on lead-mining sites in the Mechernich area of the Eifel Mountains were investigated with respect to soil factors determining the degree and type of heavy metal tolerance. Salix caprea L. (Goat Willow) grew on soils with up to 17000 mg kg^–1 total lead (ca. 4000 mg kg^–1 ammonium acetate-exchangeable Pb). Betula pendula Roth (Silver Birch) was found on soils containing as much as 29000 mg kg^–1 total lead (7000 mg kg^–1 ammonium acetate-exchangeable Pb). Other woody perennials, with the exception of the dwarf shrub Calluna vulgaris, were not found in the contaminated area even though they did occur in the immediate vicinity. The two lead-tolerant tree species did not form mixed populations.Because of a significantly lower Pb/Ca ratio in Salix soils (2.2) compared with Betula soils (7.4), a calcium-dependent mechanism of lead tolerance is suggested for Salix, but not for Betula.The Betula population could be divided into two groups, each showing a highly significant correlation between root-lead content and exchangeable lead amounts in the soil, but with different levels of lead uptake. The only soil factor distinguishing the two groups was found to be the level of soluble phosphate. A distinctly low level of soil phosphate correlated with a high lead concentration in roots of the one group (30000 mg Pb kg^–1 DW), whereas high phosphate amounts corresponded with a much lower lead concentration in roots of the other (12000 mg Pb kg^–1 DW^–1). Since the correlation between lead in the soil and in plants was similar for the two groups, it is concluded that the type of lead tolerance in Betula is determined by the status of plant phosphate nutrition, rather than by simple phosphate precipitation in the soil.A comparison of growth between different populations of Betula seedlings on homogenized soils from the mining area revealed the Mechaernich population to be a distinct ecotype with respect to lead tolerance. The control population obtained from a non-contaminated area exhibited a lower degree of lead tolerance coupled with a two-step strategy of adaptation to lead.     

Forest structure of a subtropical mangrove along a river inferred from potential tree height and biomass

The hypothesis was tested that potential tree height and biomass in mangroves decrease downstream with the tidal gradient along the Okukubi River in Okinawa Island, Japan. The mangrove stands consisted of Bruguiera gymnorrhiza (L.) Lamk. and Kandelia obovata (S., L.) Yong (Rhizophoraceae). Four sites were selected considering the distance from the mouth of the river. Soil salinity increased downstream, while soil total nitrogen content decreased. The soil redox potential did not vary along the river. Maximum gross photosynthesis and tree height for each species decreased downstream. The potential tree height (H_max) inferred from the stem diameter (D_0.1)–tree height (H) relationship ( a, h, coefficient) in each species decreased downstream. The tree density (ρ)–mean tree size () relationships ( K, α, coefficient) determined for four sites revealed that the mean tree size at any given tree density decreased downstream, which indicates the decrease of potential biomass. Furthermore, an index for biomass () was homogeneous within a site regardless of tree density, i.e. the value of α at each site did not differ significantly from 1.0 (p > 0.05). The decreases in potential tree height and biomass may be partially ascribed to the stressful environments at the downstream sites characterized by high salinity (>2.6%) and nitrogen-poor soils (<0.25 ppt) in our study area.     

Response of pondcypress growth rates to sewage effluent application

Cores were collected from dominant pondcypress trees growing in a swamp that had received sewage effluent for 7 yr and a nearby control swamp to determine the combined effects of changes in nutrient supply and hydrologic regime on tree growth. The cores were used to measure two indices of tree growth: basal area increment (BAI) and relative basal area increment (RBAI, which accounts for differences in growth due to the size of teh tree) between 1970–1983 while one swamp remained untreated and the other received weekly additions of sewage effluent from 1974–1981. Throughout the whole period, the mean BAI and RBAI of pond-cypress trees in the untreated swamp remained unchanged, ranging between 5.55–6.38 cm² yr^–1 and 1.09–1.27% yr^–1, respectively. In contrast, trees in the treated swamp increased their BAI approximately two-fold from 7.40 cm² yr^–1 prior to treatment to 14.83 cm² yr^–1 after the onset of treatment and maintained this rate of growth in the 2 yr period after cessation of treatment. Relative basal area increment showed a similar response, but the proportional increase due to treatment was less (1.5-fold factor) than for BAI. The response of pondcypress trees to the sewage effluent differed depending upon whether the trees were located in the deep or shallow water zones. Trees in the deep zone of the treated swamp had lower BAIs and RBAIs than those in the shallow zone during the treatment period, whereas in pre- and post-treatment periods growth indices were equal in both zones. No significant differences in growth between deep and shallow zones were observed during all three time periods in the control swamp.