Seasonal changes in photosynthesis of four understory herbs in deciduous forests

Seasonal patterns of photosynthesis and respiration of single leaves of four understory perennial herbs in deciduous forests were investigated in relation to their leaf growth and light conditions on the forest floor.Anemone flaccida shows rapid growth of leaf area and high rates of gross photosynthesis at light saturation (Psat) in its early stage of development. Its photosynthetic activity is restricted to a brief period of high light intensity before the closure of overstory canopies.Disporum smilacinum possesses light-photosynthesis curves of the shade-leaf type throughout its whole growing period. A shading experiment has shown that this plant is low-light adapted and can utilize weak light efficiently. The light-photosynthesis curve ofSyneilesis palmata shifts from the sun-leaf type to the shade-leaf type in response to the seasonal change of light regime on the forest floor. Evergreen leaves ofPyrola japonica have three year longevity, and light-photosynthesis curves of the shade-leaf type. They maintain some photosynthetic activity even in late autumn and winter.     

Leaf photosynthesis in pure swards of two grasses (Lolium perenne and Lolium multiflorum) subjected to contrasting intensities of defoliation

Photosynthetic rates were measured on light saturated, fully-expanded leaves in pure swards of Lolium perenne and Lolium multiflorum during late summer using 14-carbon dioxide. These swards were defoliated by cutting at three heights of 3, 6 and 9 cm above the ground. The photosynthetic rates of leaves on tillers in swards cut constantly 3 cm above ground level were higher than those of leaves in swards cut constantly 9 cm above ground level. Additional treatments with various sequences of cuts 3, 6 and 9 cm above ground level were designed to reduce damage to the growing points of tillers whilst ensuring satisfactory harvesting of the shoots. The photosynthetic rates of leaves on tillers cut to various heights above ground level at successive harvests were intermediate between those of similar leaves in the constantly low and high cut swards. The rates of photosynthesis of Italian ryegrass leaves were higher than those of perennial ryegrass leaves for a short time after defoliation of the constantly high cut swards. However, these perennial ryegrass leaves quickly adapted their rates of photosynthesis to the higher irradiances they received after cutting. Thus grass species differing in morphology adjust to management practices by the use of different photosynthetic strategies.     

Stand microclimate and physiological activity of tree leaves in an oak-hornbeam forest

Summary In an uneven-aged, multi-species oak-hornbeam forest at Báb, SW Slovakia (former IBP Forest Research Site), a series of micrometeorological and ecophysiological measurements started in 1985. The aims of the work are to improve understanding of physiological processes (photosynthesis, respiration, and transpiration) of adult trees and stand microclimate, to collect data for simulation of the canopy (stand) photosynthesis and for ecological synthesis of the functioning of the forest ecosystem. In this paper, photosynthetically active radiation (PAR), air temperature (AT) and relative humidity (RH), wind speed (WS), and CO₂ concentration ([CO₂]) in and above the forest are characterized for the fully leaved season, using diurnal courses, vertical profiles and isodiagrams (isopleths). Approximately 50% of incident PAR was absorbed by the upper 4–5 m layer of leaves and only approximately 5% or less penetrated to the forest floor. Vertical gradients of AT and RH were generally low, but large differences in diurnal ranges of AT and RH were observed between vertical levels. The upper leaf canopy greatly reduced WS, and at a height of about 14 m above the ground it was close to zero. The highest diurnal [CO₂] maximum and variations occurred at 1 m above the ground, and the lowest above the forest. In good light conditions in the forest, the entire leaf canopy (overstorey and understorey canopy) is a large sink of CO₂. At night the forest stand is a source of CO₂, the largest internal source being the soil and forest floor.     

盾叶莓的生物学性状及开发前景

盾叶莓为蔷薇科悬钩子属的多年生落叶灌木,主要分布区在我国苏、浙、皖等省,生长于海拔700 m以上到1 350 m的山区。其地上枝条为两年生,当年生枝条可生长到1～2 m高,秋季落叶经冬季休眠后,次年的5～7月份开花结果后迅速枯死。其果实大,且营养价值高,繁殖容易,是一种具有开发利用价值的野生水果资源。     

Phenology and wintering capacity of sporophytes and gametophytes of ferns native to northern Japan

Summary To investigate life history adaptations to cold climates, the leaf development, sporulation period, growing stage of gametophytes, and the frost and drought resistance of sporophytes and gametophytes of 67 fern species native to Kokkaido were studied. Most ferns common in Hokkaido are summer-green with leaves developing during late May to June and decaying during October. Most of the ferns in Hokkaido sporulate during August to early September. Spores dispersed from June to September germinate before winter begins, forming vegetative prothallia. Gametophytes mature only in the following summer. Thus in Hokkaido the gametophytes as well as perennial sporophytes are exposed to severe winter conditions. In order to correlate the life cycles of temperate ferns with winter cold stress, frost resistance of gametophytes, rhizomes, and leaves of sporophytes were determined. Maximal frost resistance of rhizomes reflects the stress conditions of their habitats: rhizomes of forest understory ferns are damaged at-5°to -17.5°C, epiphytic ferns and ferns of habitats exposed to severe frost sustained temperatures of -20° to-40°C. The leaves of winter-green and evergreen ferns resist frost ranging from -25° to -40°C. The leaves of summer-green ferns are killed by late frost below -5°C. With some exceptions, gametophytes of ferns growing on the forest floor resist frost to -40°C and are much hardier than sporophytes. These results suggest the possible restrictive effects of cold climate on the life span of leaves as well as on the sporulation period. If winter cold is one of the decisive factors for seasonality expression and habitat distribution of ferns, the sensitive generation must be the sporophyte rather than the gametophyte. The hardier gametophyte is therefore able to colonize habitats in which the sporophyte is excluded by frost if mechanisms of vegetative propagation are evolved.Contribution No. 2451 from The Institute of Low Temperature Science     

Coarse woody debris in Australian forest ecosystems: A review

Abstract Coarse woody debris (CWD) is the standing and fallen dead wood in a forest and serves an important role in ecosystem functioning. There have been several studies that include estimates of CWD in Australian forests but little synthesis of these results. This paper presents findings from a literature review of CWD and fine litter quantities. Estimates of forest‐floor CWD, snags and litter from the literature are presented for woodland, rainforest, open forest and tall open forest, pine plantation and native hardwood plantation. Mean mass of forest floor CWD in Australian native forests ranged from 19 t ha^?1 in woodland to 134 t ha^?1 in tall open forest. These values were generally within the range of those observed for similar ecosystems in other parts of the world. Quantities in tall open forests were found to be considerably higher than those observed for hardwood forests in North America, and more similar to the amounts reported for coniferous forests with large sized trees on the west coast of the USA and Canada. Mean proportion of total above‐ground biomass as forest floor CWD was approximately 18% in open forests, 16% in tall open forests, 13% in rainforests, and 4% in eucalypt plantations. CWD can be high in exotic pine plantations when there are considerable quantities of residue from previous native forest stands. Mean snag biomass in Australian forests was generally lower than the US mean for snags in conifer forests and higher than hardwood forest. These results are of value for studies of carbon and nutrient stocks and dynamics, habitat values and fire hazards.     

The Canopy Starts at 0.5 m: Predatory Mites (Acari: Mesostigmata) Differ between Rain Forest Floor Soil and Suspended Soil at any Height

Suspended soils in forest canopies are thought to harbor a substantial fraction of canopy biomass and many arboreal specialists, but do forest floor generalist predators with high vagility also use this habitat? We tested the hypothesis of no difference between forest floor and suspended-soil predatory mite faunas (Acari: Mesostigmata) in an Australian rain forest. Our results show that instead of being habitat generalists, many predatory mites partition soil into two main strata: soil suspended aboveground irrespective of height (0.5–20 m) and soil on the ground. Of 53 species of Mesostigmata in suspended soil, 53 percent (28 species) were absent from or rarely found on the ground. This increased to 60 percent (15/25 species) if only common species are considered. Among these 15 ‘suspended-soil specialists’, all but the three least abundant were found throughout the arboreal strata. Moreover, ten species also occurred in litter accumulated on the surface of decaying logs or boulders close to the forest floor. Thus, although the arboreal predatory mite fauna is distinct from that on the forest floor, it is not restricted to the high canopy: even slightly elevated substrate appears acceptable as habitat for these suspended-soil specialists. Our data suggest that a substantial portion of a rain forest's soil and litter fauna is held above the forest floor.     

Life-history monographs of Japanese plants. 12: Asarum caulescens Maxim. (Aristolochiaceae)

The life-history characteristics of Asarum caulescens Maxim. (Aristolochiaceae) are described here. This typical summer-green perennial of the Aristolochiaceae is a unique woodland element distributed in central Honshu, that is, south of Fukushima Prefecture to Shikoku and Kyushu, Japan. It grows in the somewhat shady understory of temperate montane deciduous broad-leaved or mixed forests associated with conifers, such as Cryptomeria japonica and Abies firma . One of the unique features of A. caulescens is its rhizome structures in combination with aerial shoots, which play an important role in vegetative propagation. The perennation strategy of A. caulescens is exceedingly complex, producing various combinations of linear or branched rhizomes bearing different numbers of leaves. Rhizomes consist of two or three segments, each 5 mm to 5 cm long as a unit, connected linearly, but also occasionally branched. At the tip of the newly formed rhizome segment, a single leaf or a pair of cordate leaves are formed. Exceedingly complex branching patterns of the rhizome segments were also recognized, forming ramets in late June to early July. Vegetative propagation by ramet formation obviously plays a very important role in the maintenance of local populations.     

Canopy gaps formed by mangrove trimming: an experimental test of impact on litter fall and standing litter stock in Southwest Florida (USA)

Mangroves in Florida (USA) are subject to horticultural pruning that may increase the size of canopy gaps and alter rates of litter production and accumulation. Mangrove canopy gap formation is a common phenomenon, known to alter abiotic conditions near the forest floor. Using a series of field experiments in Rookery Bay, Florida, the effects of mangrove trimming on canopy density, mangrove litter production, standing litter stocks, and the decomposition rate of Rhizophora mangle leaves on the forest floor were assessed. Litter trap collections over the year following mangrove trimming indicated that pruned mangrove stands (canopy coverage: 42.8±0.9%; mean±S.E.) produced approximately one-half of the litter of mangrove stands with relatively complete canopies (canopy coverage: 72.1±0.5%). However, there was no significant difference between the mass of standing litter on the forest floor beneath reduced canopy and intact canopy mangroves. Also, R. mangle leaves held on the forest floor in fiberglass litter bags at both reduced canopy and control sites did not decompose at different rates over 28 days. These results indicate that while system-wide mangrove litter production should be reduced by the formation of these gaps in mangrove forests, postproduction influences may obscure any site-specific declines in standing litter stocks.     

Leaf δ13C in Pinus resinosa trees and understory plants: variation associated with light and CO2 gradients

 Our objective was to evaluate the relative importance of gradients in light intensity and the isotopic composition of atmospheric CO₂ for variation in leaf carbon isotope ratios within a Pinus resinosa forest. In addition, we measured photosynthetic gas exchange and leaf carbon isotope ratios on four understory species (Dryopteris carthusiana, Epipactus helleborine, Hieracium floribundum, Rhamnus frangula), in order to estimate the consequence of the variation in the understory light microclimate for carbon gain in these plants. During midday, CO₂ concentration was relatively constant at vertical positions ranging from 15 m to 3 m above ground. Only at positions below 3 m was CO₂ concentration significantly elevated above that measured at 15 m. Based on the strong linear relationship between changes in CO₂ concentration and δ¹³C values for air samples collected during a diurnal cycle, we calculated the expected vertical profile for the carbon isotope ratio of atmospheric CO₂ within the forest. These calculations indicated that leaves at 3 m height and above were exposed to CO₂ of approximately the same isotopic composition during daylight periods. There was no significant difference between the daily mean δ¹³C values at 15 m (–7.77‰) and 3 m (–7.89‰), but atmospheric CO₂ was significantly depleted in ¹³C closer to the ground surface, with daily average δ¹³C values of –8.85‰ at 5 cm above ground. The light intensity gradient in the forest was substantial, with average photosynthetically active radiation (PAR) on the forest floor approximately 6% of that received at the top of the canopy. In contrast, there were only minor changes in air temperature, and so it is likely that the leaf-air vapour pressure difference was relatively constant from the top of the canopy to the forest floor. For red pine and elm tree samples, there was a significant correlation between leaf δ¹³C value and the height at which the leaf sample was collected. Leaf tissue sampled near the forest floor, on average, had lower δ¹³C values than samples collected near the top of the canopy. We suggest that the average light intensity gradient through the canopy was the major factor influencing vertical changes in tree leaf δ¹³C values. In addition, there was a wide range of variation (greater than 4‰) among the four understory plant species for average leaf δ¹³C values. Measurements of leaf gas exchange, under natural light conditions and with supplemental light, were used to estimate the influence of the light microclimate on the observed variation in leaf carbon isotope ratios in the understory plants. Our data suggest that one species, Epipactus helleborine, gained a substantial fraction of carbon during sunflecks. Received: 21 March 1996 / Accepted: 13 August 1996     

First direct landscape-scale measurement of tropical rain forest Leaf Area Index, a key driver of global primary productivity

Leaf Area Index (leaf area per unit ground area, LAI) is a key driver of forest productivity but has never previously been measured directly at the landscape scale in tropical rain forest (TRF). We used a modular tower and stratified random sampling to harvest all foliage from forest floor to canopy top in 55 vertical transects (4.6 m²) across 500 ha of old growth in Costa Rica. Landscape LAI was 6.00 ± 0.32 SEM. Trees, palms and lianas accounted for 89% of the total, and trees and lianas were 95% of the upper canopy. All vertical transects were organized into quantitatively defined strata, partially resolving the long-standing controversy over canopy stratification in TRF. Total LAI was strongly correlated with forest height up to 21 m, while the number of canopy strata increased with forest height across the full height range. These data are a benchmark for understanding the structure and functional composition of TRF canopies at landscape scales, and also provide insights for improving ecosystem models and remote sensing validation.     

Afforestation effects on SOC in former cropland: oak and spruce chronosequences resampled after 13 years

Chronosequences are commonly used to assess soil organic carbon (SOC) sequestration after land‐use change, but SOC dynamics predicted by this space‐for‐time substitution approach have rarely been validated by resampling. We conducted a combined chronosequence/resampling study in a former cropland area (Vestskoven) afforested with oak (Quercus robur) and Norway spruce (Picea abies) over the past 40 years. The aims of this study were (i) to compare present and previous chronosequence trends in forest floor and top mineral soil (0–25 cm) C stocks; (ii) to compare chronosequence estimates with current rates of C stock change based on resampling at the stand level; (iii) to estimate SOC changes in the subsoil (25–50 cm); and (iv) to assess the influence of two tree species on SOC dynamics. The two chronosequence trajectories for forest floor C stocks revealed consistently higher rates of C sequestration in spruce than oak. The chronosequence trajectory was validated by resampling and current rates of forest floor C sequestration decreased with stand age. Chronosequence trends in topsoil SOC in 2011 did not differ significantly from those reported in 1998, however, there was a shift from a negative rate (1998: ?0.3 Mg C ha^?1 yr^?1) to no change in 2011. In contrast SOC stocks in the subsoil increased with stand age, however, not significantly (P = 0.1), suggesting different C dynamics in and below the former plough layer. Current rates of C change estimated by repeated sampling decreased with stand age in forest floors but increased in the topsoil. The contrasting temporal change in forest floor and mineral soil C sequestration rates indicate a shift in C source‐sink strength after approximately 40 years. We conclude that afforestation of former cropland within the temperate region may induce soil C loss during the first decades followed by a recovery phase of yet unknown duration.     

The above-ground coarse wood productivity of 104 Neotropical forest plots

The net primary production of tropical forests and its partitioning between long‐lived carbon pools (wood) and shorter‐lived pools (leaves, fine roots) are of considerable importance in the global carbon cycle. However, these terms have only been studied at a handful of field sites, and with no consistent calculation methodology. Here we calculate above‐ground coarse wood carbon productivity for 104 forest plots in lowland New World humid tropical forests, using a consistent calculation methodology that incorporates corrections for spatial variations in tree‐size distributions and wood density, and for census interval length. Mean wood density is found to be lower in more productive forests. We estimate that above‐ground coarse wood productivity varies by more than a factor of three (between 1.5 and 5.5 Mg C ha^?1 a^?1) across the Neotropical plots, with a mean value of 3.1 Mg C ha^?1 a^?1. There appear to be no obvious relationships between wood productivity and rainfall, dry season length or sunshine, but there is some hint of increased productivity at lower temperatures. There is, however, also strong evidence for a positive relationship between wood productivity and soil fertility. Fertile soils tend to become more common towards the Andes and at slightly higher than average elevations, so the apparent temperature/productivity relationship is probably not a direct one. Coarse wood productivity accounts for only a fraction of overall tropical forest net primary productivity, but the available data indicate that it is approximately proportional to total above‐ground productivity. We speculate that the large variation in wood productivity is unlikely to directly imply an equivalent variation in gross primary production. Instead a shifting balance in carbon allocation between respiration, wood carbon and fine root production seems the more likely explanation.     

Cantharidin world in air: Spatiotemporal distributions of flying canthariphilous insects in the forest interior

The natural community in which the members interact using a toxic terpenoid cantharidin is named the “cantharidin world.” In previous studies, however, the members of this world have been surveyed only on the forest floor by setting pitfall traps with cantharidin as an attractant. In this study, we set cantharidin traps at various heights above the forest floor to investigate the structure and functional diversity of the canthariphilous flying insect community in the forest above‐ground space. A total of 3,168 arthropods were collected by the traps; among them, six species were more attracted to cantharidin than to control traps. Pseudopyrochroa brevitarsis and P. laticollis (Colecoptera: Pyrochroidae) both appeared for a short time during spring, but the latter species tended to use a lower layer of the forest. Clavicollis fugiens (Coleoptera: Anthicidae) also appeared in spring and flew near the ground. In these beetles, the attracted individuals were mostly males; they may use the obtained cantharidin for nuptial gifts to the female. Atrichopogon femoralis, A. insularis and Atrichopogon sp. (Diptera: Ceratopogonidae) were collected widely in the forest above‐ground space. These midges were almost females, probably because only females of these insects use chemical cues, including cantharidin, for searching for arthropods from which to suck hemolymph.     

Leaf temperatures and energy balance ofWelwitschia mirabilis in its natural habitat

Summary Welwitschia mirabilis is a perennial desert plant with extremely large leaves (0.5–1.0 m broad, 1–2 m long). Leaf temperatures were measured in the field and the energy budget was calculated. The portions of the leaf which were kept above the ground had leaf temperatures which were only 4–6°C above air temperature. In the leaf portions which were in contact with the ground leaf temperatures were 6–12°C above air temperature (absolute maximum 51°C). The important feature in the energy budget ofWelwitschia mirabilis is its high reflectivity (38% of the global radiation). Only about 56% of the global radiation is absorbed by the thick leathery leaves. The energy loss due to convection is of the same order of magnitude as the reflection and it is abouy the same in the portions of leaf on and above the ground. The difference in leaf temperatures found in these portions is due to the loss of thermal radiation from the section of leaf above the ground to the cooler ground which is shaded by the leaf. The provision of a heat sink due to the large area of shade cast by these large leaves is of significance to the existence ofWelwitschia mirabilis in its arid habitats.     

Forest Floor Leaf Cover as a Barrier for Dust Accumulation in Tai National Park: Implications for Primate Dental Wear Studies

Mastication causes distinct use wear scars on teeth that can aid in dietary reconstructions of fossils. However, the role played by exogenous grit in dental wear complicates the association between wear and diet. Dental wear analyses often assume that foods closer to the soil contain more exogenous grit than those in the forest canopy. Yet, a layer of leaf litter covering many forest floors may trap grit from the soil, keeping it from settling on surrounding vegetation or becoming part of atmospheric dust. Cercocebus atys is frequently referenced in dental wear studies because of its dependence on hard Sacoglottis gabonensis seeds collected from the forest floor. Here we examine quantities of dust deposition at different forest levels and assess its potential role in wear patterns observed in C. atys. We collected grit from S. gabonensis seeds (N?=?64) found under the leaf litter and compared them to grit samples taken from the surface of leaves at different forest strata (N?=?450) in Ivory Coast’s Taï National Park. Seeds underneath the leaf litter were coated with significantly more grit than leaves above the leaf material and we conclude that leaf litter is a significant barrier to grit particles originating from the soil. Given that evidence points to a significant difference in grit amount between foodstuffs on the ground and foods near the ground, the findings lead to a prediction of differences in dental wear patterns between purely arboreal foragers and those incorporating terrestrial food sources.     

Evaluating minirhizotron estimates of fine root longevity and production in the forest floor of a temperate broadleaf forest

The minirhizotron technique (MR) for in situ measurement of fine root dynamics offers the opportunity to obtain accurate and unbiased estimates of root production in perennial vegetation only if MR tubes do not affect the longevity of fine roots. Assuming fine root biomass is near steady-state, fine root production (g m^–2 yr^–1) can be estimated as the ratio of fine root biomass (g m^–2) to median fine root longevity (yr). This study evaluates the critical question of whether MR access tubes affect the longevity of fine roots, by comparing fine root survivorship obtained using MR with those from a non-intrusive in situ screen method in the forest floor horizons of a northern hardwood forest in New Hampshire, USA. Fine root survivorship was measured in 380 root screens during 1993–1997 and in six horizontal minirhizotron tubes during 1996–1997. No statistically significant difference was found between estimates of survivorship of fine roots (<1 mm dia.) at this site from MR versus from in situ screens, suggesting that MR tubes do not substantially affect fine root longevity in the forest floor of this northern hardwood forest and providing greater confidence in measurements of fine root production using the MR technique. Furthermore, the methodology for estimating fine root production from MR longevity data was evaluated by comparison of fine root longevity and production estimates made using single vs. multiple root cohorts, and using root-number, root-length, and root-mass weighted methods. Our results indicate that fine root-length longevity estimates based on multiple root cohorts throughout the year can be used to approximate fine root biomass production. Using this method, we estimated fine root longevity and production in the forest floor at this site to be 314 days (or 0.86 yr) and 303 g m^–2 yr^–1, respectively. Fine root production in this northern hardwood forest is approximately equivalent to standing biomass and was previously underestimated by root in-growth cores. We conclude that the use of MR to estimate fine root longevity and production as outlined here may result in improved estimates of fine root production in perennial vegetation.     

Recovery of pristine boreal forest floor community after selective removal of understorey, ground and humus layers

In boreal spruce forests that rarely experience extensive disturbances, fine-scale vegetation gaps are important for succession dynamics and species diversity. We examined the community implications of fine-scale gap disturbances by selective removal of vegetation layers in a pristine boreal spruce forest in Northern Finland. The aim was to investigate how the speed of recovery depends on the type of disturbance and the species growth form. We also wanted to know if there appeared changes in species composition after disturbance. Five different treatments were applied in the study: Control, removal of the ground layer (bryophytes and lichens), removal of the understorey layer (dwarf shrubs, herbs and graminoids), removal of both the ground and understorey layers, and complete removal of the vegetation and humus layers above the mineral soil. The vegetation recovery was monitored in terms of cover and species numbers over a 5-year period. Understorey layer cover, composed mainly of clonal dwarf shrubs, recovered completely in 4 years in treatments where the humus layer remained intact, whereas ground layer cover did not reach the control level in plots from where bryophytes and lichens were removed. Recovery was faster in terms of species number than species cover. Bryophytes, graminoids and dominant dwarf shrubs appeared in all disturbed plots quickly after disturbance. Seedlings of trees appeared exclusively in disturbed plots. Graminoids dominated after the removal of humus layer. The results indicate that the regeneration of forest floor after small gap disturbance occurs mainly by re-establishment of the dominant species. Although destruction of the humus layer leaves a long-lasting scar to the forest floor, exposing of mineral soil may enhance the sexual reproduction of dominant species and the colonization of weaker competitors.