Recovery of floristic diversity and basal area in natural forest regeneration and planted plots in a Costa Rican wet forest

When compared to planted reforestation, natural unassisted regeneration is often reported to result in slow recovery of biomass and biodiversity, especially early in succession. In some cases, naturally regenerating forests are not comparable to the community structure of primary forests after many decades. However, direct comparison of the outcomes of tropical forest restoration and natural regeneration is hindered by differences in metrics of forest recovery, inconsistency in land use histories, and dissimilarities in experimental design. We present the results of a replicated reforestation experiment comparing natural regeneration and polyculture tree planting at multiple diversity levels (3, 6, 9, or 12 native tree species), with uniform land use history and initial edaphic conditions. We compare the recovery of basal area and floristic diversity in these treatments after 5 yr of succession. Total basal area was higher in planted plots than in naturally regenerating plots, but it but did not vary among the different planted diversity levels. The basal area of woody recruits did not differ among treatments. The diversity of woody recruits increased substantially over time but did not vary among planting treatments. Species composition trajectories showed directional turnover over time, with no consistent differences among treatments. The convergence of restoration trajectories and similarity of floristic community diversity and composition across all treatments, after only 5 yr, provides evidence of the viability of natural regeneration for rapid restoration of forest biodiversity.     

Enrichment planting does not improve tree restoration when compared with natural regeneration in a former pine plantation in Kibale National Park,Uganda

Given the high rates of deforestation and subsequent land abandonment, there are increasing calls to reforest degraded lands; however, many areas are in a state of arrested succession. Plantations can break arrested succession and the sale of timber can pay for restoration efforts. However, if the harvest damages native regeneration, it may be necessary to intervene with enrichment planting. Unfortunately, it is not clear when intervention is necessary. Here, we document the rate of biomass accumulation of planted seedlings relative to natural regeneration in a harvested plantation in Kibale National Park, Uganda. We established two 2‐ha plots and in one, we planted 100 seedlings of each of four native species, and we monitored all tree regeneration in this area and the control plot. After 4 years, naturally regenerating trees were much taller, larger and more common than the planted seedlings. Species richness and two nonparametric estimators of richness were comparable between the plots. The cumulative biomass of planted seedlings accounted for 0.04% of the total above‐ground tree biomass. The use of plantations facilitated the growth of indigenous trees, and enrichment planting subsequent to harvesting was not necessary to obtain a rich tree community with a large number of new recruits.     

Non-native grass invasion suppresses forest succession

Multiple factors can affect the process of forest succession including seed dispersal patterns, seedling survival, and environmental heterogeneity. A relatively understudied factor affecting the process of succession is invasions by non-native plants. Invasions can increase competition, alter abiotic conditions, and provide refuge for consumers. Functional traits of trees such as seed size and life history stage may mediate the effects of invasions on succession. We tested the effects of the forest invader Microstegium vimineum on planted and naturally regenerating trees in a multi-year field experiment. We established plots containing nine species of small- and large-seeded tree species planted as seeds or saplings, and experimentally added Microstegium to half of all plots. Over 3 years, Microstegium invasion had an overall negative effect on small-seeded species driven primarily by the effect on sweetgum, the most abundant small-seeded species, but did not affect large-seeded species such as hickory and oak species, which have more stored seed resources. Natural regeneration was over 400% greater in control than invaded plots for box elder, red maple, and spicebush, and box elder seedlings were 58% smaller in invaded plots. In contrast to the effects on tree seedlings, invasion did not affect tree sapling survival or growth. Microstegium may be directly reducing tree regeneration through competition. Invaded plots had greater overall herbaceous biomass in 2006 and 2008 and reduced light availability late in the growing season. Indirect effects may also be important. Invaded plots had 120% more thatch biomass, a physical barrier to seedling establishment, and significantly greater vole damage to tree saplings during 2006 and 2007. Our results show that two tree functional traits, seed size and life history stage, determined the effects of Microstegium on tree regeneration. Suppression of tree regeneration by Microstegium invasions may slow the rate of forest succession and alter tree species composition.     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

Strategic light manipulation as a restoration strategy to reduce alien grasses and encourage native regeneration in Hawaiian mesic forests

Question: Is there a light level at which alien grass biomass is reduced while still supporting growth and survival of native woody species, allowing for native species regeneration in abandoned pastures? Location: Island of Hawaii, USA. Methods: In a two‐part study we examined the effect of light availability on common native woody and alien grass species found in secondary forests in Hawaii. A field survey was conducted to examine the relationship between light availability and canopy type (open pasture, planted canopy and secondary forest) on understory grass biomass and litter accumulation. We then experimentally manipulated light levels to determine the effect of light availability on growth and survival of six native woody species and three alien grasses. Low‐light (5%), medium‐light (10%) and high‐light (20‐30%) treatments were created using shade structures erected beneath the existing secondary koa canopy. Results: In the field survey, alien grass biomass was greatest under the open pasture and lowest in the secondary forest. There was a positive correlation between understory light availability and alien grass biomass. In the experimental study, large reductions in relative growth rates were documented for all of the grass species and four of the six woody species under the lowest light level. Although growth at 5% light is substantially reduced, survival is still high (84‐100%), indicating that these species may persist under closed canopy. Conclusion: Low‐light conditions result in the greatest reduction in alien grass biomass while creating an environment in which native woody species can grow and survive.     

Demographic costs and benefits of natural regeneration during tropical forest restoration

For tropical forest restoration to result in long‐term biodiversity gains, native trees must establish self‐sustaining populations in degraded sites. While many have asked how seedling recruitment varies between restoration treatments, the long‐term fate of these recruits remains unknown. We address this research gap by tracking natural recruits of 27 species during the first 7 years of a tropical forest restoration experiment that included both planted and naturally regenerating plots. We used an individual‐based model to estimate the probability that a seedling achieves reproductive maturity after several years of growth and survival. We found an advantage for recruits in naturally regenerating plots, with up to 40% increased probability of reproduction in this treatment, relative to planted plots. The demographic advantage of natural regeneration was highest for mid‐successional species, with relatively minor differences between treatments for early‐successional species. Our research demonstrates the consequences of restoration decision making across the life cycle of tropical tree species.     

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments. __________ Translated from Acta Ecologica Sinica, 2006, 26(12): 4019–4026 [译自: 生态学报]     

Responses of Photosynthesis and Biomass Allocation of an Understorey Herb, Pteridophyllum racemosum, to Gradual Increases in Irradiance

In wave-regenerating Abies forests in central Japan, light availabilityon the forest floor of a mature-tree stand increases graduallyover 4 to 5 years during the senescence of trees. The understoreyevergreen perennialPteridophyllum racemosum Sieb. et Zucc. (Papaveraceae)is commonly found in such forests. To determine the acclimationcharacteristics of this species to gradual changes in lightavailability, leaf photosynthetic traits and biomass allocationpatterns were determined along a transect running from fullyshaded understorey to a canopy gap. Along the transect, thephoton flux density on the forest floor relative to full sun(RPFD) varied from 2.3–36.7%. The light-saturated netphotosynthetic rate of field-grown plants increased as the lightenvironment increased from 2.3–14.1% RPFD. However, aslight conditions increased from moderate (14.1%) to high (36.7%),the light-saturated net photosynthetic rate decreased to a levelcomparable to that exhibited by low-light grown plants. On theother hand, root mass per unit leaf area (RMA) increased withincreasing light conditions. A higher RMA contributes to a higherwater uptake capacity per unit leaf area which in turn supportshigher rates of photosynthesis and transpiration in high light.We conclude that in wave-regenerating forests P. racemosum cansurvive in high-light environments by avoiding a severe waterdeficit and heat load by increasing RMA. Copyright 2001 Annalsof Botany Company Biomass allocation, photosynthesis, Pteridophyllum racemosum, root mass per unit leaf area, wave regeneration     

Environmental factors influencing herb layer productivity in Central European oak forests: insights from soil and biomass analyses and a phytometer experiment

Habitat productivity and vegetation biomass are important factors affecting species diversity and ecosystem function, but factors determining productivity are still insufficiently known, especially in the forest herb layer. These factors are difficult to identify because different methods often yield different results. We sampled the herb layer biomass and assessed soil nutrients, moisture and light availability in 100 m² plots in Czech oak forests. Habitat productivity was estimated independently from nutrient content in the soil, herb layer biomass and using a bioassay experiment (growing phytometer plants of Raphanus sativus under standardised conditions in soil samples taken from forest plots). The generalised linear model for herb layer biomass showed it to increase with light, soil phosphorus and moisture availability, but only 10.7% of its variation was explained by these factors. The phytometer biomass increased mainly with soil pH and phosphorus availability; together with soil C/N ratio these factors explained 56.1% of the phytometer biomass variability. Combined evidence based on different approaches indicates that canopy shading and soil phosphorus tend to be the most important factors influencing the herb layer productivity of the studied oak forests.     

Impacts of non-native plant and animal invaders on gap regeneration in a protected boreal forest

In balsam fir (Abies balsamea)-dominated boreal forests of Gros Morne National Park, Newfoundland (Canada), non-native Cirsium arvense (Canada thistle) has invaded forest gaps. Its management is complicated by the lack of viable control techniques and an overarching issue of gap regeneration failure attributed to browsing by non-native moose (Alces alces). This study identifies the impacts of thistle invasion on balsam fir regeneration and explores protocols to re-establish fir in gaps invaded by thistle and moose. Fir seeds were planted into ten gaps (five natural; five anthropogenic) and the emergence, growth, herbivory damage, and survival of fir was determined for 2 years amongst five treatments (n = 50 plots; 32 seeds/plot): (1) thistle monocultures in gaps; (2) where aboveground thistle biomass was removed; (3) where above- and below-ground thistle biomass was removed; (4) non-invaded areas in gaps; and, (5) adjacent uninvaded forest edges. In addition, 432 fir seedlings (aged 15 months) were transplanted into four forest gaps within the above treatments and followed for 1 year. Results indicate that invasion of C. arvense negatively affects fir emergence and early survival, and may further contribute to continued balsam fir regeneration failure independent of future moose densities. However, older fir seedlings transplanted into thistle monocultures experienced a positive facilitative effect due to the protection thistle provided against small mammal herbivory. Restoration actions that combine moose density reductions with the planting of fir seedlings offers the most viable long-term strategy to re-establish the native forest canopy in thistle-invaded gaps and would likely lead to the eventual decline of shade-intolerant C. arvense.     

Ecological restoration, carbon sequestration and biodiversity conservation: The experience of the Society for Wildlife Research and Environmental Education (SPVS) in the Atlantic Rain Forest of Southern Brazil

Since 1999, SPVS has been involved in three projects that combine two fundamental goals over the course of 40 years: the conservation of one of Brazil's most important remnants of Atlantic Forest and the implementation of projects for carbon sequestration. In addition, there is an interest in replicating these projects in order to restore other degraded areas, protect the Brazilian biomes, and help to diminish deforestation and forest fire, therefore reducing carbon emissions. The acquisition of 19,000 ha of degraded areas of high biological importance in southern Brazil was the first step towards the implementation of the projects. These areas are owned by SPVS, a Brazilian NGO, and are being restored, conserved and transformed into Private Natural Reserves, in partnership with the NGO – The Nature Conservancy, and financed by the companies – American Electric Power, General Motors and Chevron Texaco. The process of forest restoration involves several stages: soil studies, surveying the region's native plants, planning for restoration by means of a Geographical Information System, production of seedlings, application of different techniques for planting (such as manual or mechanised planting with seedlings and stakes), and biomass and biodiversity monitoring. To guarantee the survival of the seedlings on the planted areas, during the first three years, there is a continuous and systematic maintenance programme including weeding of undergrowth, crowing and organic fertilisation. The three projects already planted around 500,000 seedlings of native species until September 2004, and aim to plant a further 300,000 until 2008.     

Understory Colonization of Eucalyptus Plantations in Hawaii in Relation to Light and Nutrient Levels

Exotic tree plantations may serve as catalysts for native forest regeneration in agriculturally degraded landscapes. In 2001, we evaluated plant species regeneration in the understory of a 7‐year‐old experimental Eucalyptus saligna forest in Hawaii approximately 1 year after the cessation of 5 years of herbicide. These forests were organized in a 2 × 2–factorial design of planting density (1 × 1– or 3 × 3–m spacing) and fertilization (unfertilized control and regular fertilization), which resulted in varying resource availabilities. We found that understory biomass was highest under high light conditions, regardless of fertilization treatment, whereas species richness was lowest under fertilized 1 × 1–m plots. The understory was dominated by species exotic to Hawaii. The most common tree species, the noxious weed Citharexylum caudatum, was particularly successful because high light–saturated photosynthesis rates and a low light compensation point allowed for high growth and survival under both light conditions. To assess longer‐term recruitment patterns, we resurveyed a portion of this site in 2006 and also surveyed five Eucalyptus plantations in this region of Hawaii that differed in age (5–23 years), species (E. saligna, E. grandis, E. cloeziana, E. microcorys), and management (experimental, industrial, nonindustrial stewardship); all were established on previous agricultural sites within approximately 3 km of native‐dominated forest. Again, very few native species were present in any of the stands, indicating that within certain landscapes and for native species with certain life history traits, exotic plantations may be ineffective nursery ecosystems for the regeneration of native species.     

Structure and Function of Oak Forests in Central Himalaya. I. Dry Matter Dynamics

The present study deals with structure and functioning of threeareas of Himalayan oak forest. Low- and mid-altitude oaks, namelyQuercus leucotrichophora, and Quercus floribunda, form predominantevergreen forests in Central and Western Himalaya. The totaltree basal cover ranged between 33·89 m² ha^–1 (Q.floribunda site) to 36·83 m² ha^–1 (Q. leucotrichophorasite). The density ranged between 570 and 760 individuals ha^–1.Allometric equations relating biomass of different tree componentsto GBH (girth at breast height) were significant with the exceptionof leaf biomass in Q. leucotrichophora and Rhododendron arboreum.Total vegetation biomass (29·40–467·0 tha^–1) was distributed as 377·1 t ha^–1 intrees, 5·40 t ha^–1 in shrubs and 1·23 tha^–1 in herbs. Total forest floor biomass ranged between4·6 and 6·2 t ha^–1. Of the total annuallitter fall (4·7–4·8 t ha^–1), 77·5% was contributed by leaf litter, 17·8 % by wood litterand 4·7 % by miscellaneous litter. Turnover rate of treelitter varied from 0·66 to 0·70. Net primary productionof total vegetation ranged between 15·9 and 20·6t ha^–1 yr^–1, of which the contribution of trees,shrubs and herbs was 81·2 %, 8·6 % and 10·2%, respectively. A compartment model of dry matter on the basisof mean data across sites was developed to show dry matter storageand flow of dry matter within the system. Quercus leucotrichophora forest, Q. floribunda forest, Q. lanuginosa forest, biomass, litter fall, net primary production, compartmental transfer     

Invasion of glossy privet (<Emphasis Type="Italic">Ligustrum lucidum</Emphasis>) and native forest loss in the Sierras Chicas of Córdoba,Argentina

Glossy privet (Ligustrum lucidum) is a tree native to China that successfully invades forests of central Argentina. To fully understand glossy privet’s ecological effects on native forest, it is necessary to accurately map the distribution of glossy privet stands and the changes in biodiversity and forest structure of the invaded areas. The objectives of this paper were (1) to map the distribution of glossy privet stands in an area representative of the Sierras Chicas (Córdoba, Argentina) and (2) compare composition, structure and regeneration between glossy privet invaded stands and native forest stands. Using four Landsat TM images (October 2005, March, May and July 2006) we mapped the distribution of a glossy privet-dominated stand using a support vector machine, a non-parametric classifier. We recorded forest structure variables and tree diversity on 105 field plots. Glossy privet-dominated stands occupied 3,407 ha of the total forested land in the study area (27,758 ha), had an average of 33 glossy privet trees (dbh > 2.5 cm) per plot and the cover of their shrub and herb strata was substantially reduced compared with native forest. Forest regeneration was dominated by glossy privet in native forest stands adjacent to glossy privet-dominated stands. We conclude that in the Sierras Chicas glossy privet has become a widespread invader, changing the patterns of vertical structure, diversity, and regeneration in native forests.     

Shading by invasive shrub reduces seed production and pollinator services in a native herb

Plant invasions disrupt native plant reproduction directly via competition for light and other resources and indirectly via competition for pollination. Furthermore, shading by an invasive plant may reduce pollinator visitation and therefore reproduction in native plants. Our study quantifies and identifies mechanisms of these direct and indirect effects of an invasive shrub on pollination and reproductive success of a native herb. We measured pollinator visitation rate, pollen deposition, and female reproductive success in potted arrays of native Geranium maculatum in deciduous forest plots invaded by the non-native shrub Lonicera maackii and in two removal treatments: removal of aboveground L. maackii biomass and removal of flowers. We compared fruit and seed production between open-pollinated and pollen-supplemented plants to test for pollen and light limitation of reproduction. Plots with L. maackii had significantly lower light, pollinator visitation rate, and conspecific pollen deposition to G. maculatum than biomass removal plots. Lonicera maackii flower removal did not increase pollinator visitation or pollen deposition compared to unmanipulated invaded plots, refuting the hypothesis of competition for pollinators. Thus, pollinator-mediated impacts of invasive plants are not limited to periods of co-flowering or pollinator sharing between potential competitors. Geranium maculatum plants produced significantly fewer seeds in plots containing L. maackii than in plant removal plots. Seed set was similar between pollen-supplemented and open-pollinated plants, but pollen-supplemented plants exhibited higher seed set in plant removal plots compared to invaded plots. Therefore, we conclude that the mechanism of impact of L. maackii on G. maculatum reproduction was increased understory shade.     

Heterotrophic Soil Respiration in Relation to Environmental Factors and Microbial Biomass in Two Wet Tropical Forests

We examined the correlation between fungal and bacterial biomass, abiotic factors such as soil moisture, carbon in the light soil fraction and soil nitrogen to a depth of 0–25 cm and heterotrophic soil respiration using a trenching technique – in a secondary forest (Myrcia splendens, Miconia prasina and Casearia arborea) and a pine (Pinus caribeae) plantation in the Luquillo Experimental Forest in Puerto Rico. Soil respiration was significantly reduced where roots were excluded for 7 years in both the secondary forest and the pine plantation. Microbial biomass was also significantly reduced in the root exclusion plots. In root exclusion treatment, total fungal biomass was on average 31 and 65% lower than the control plots in the pine plantation and the secondary forest, respectively, but the total bacterial biomass was 24 and 8.3% lower than the control plots in the pine plantation and the secondary forest, respectively. Heterotrophic soil respiration was positively correlated with fungal biomass (R²=0.63, R²=0.39), bacterial biomass (R²=0.16, R²=0.45), soil moisture (R²=0.41, R²=0.56), carbon in light fraction (R²=0.45, R²=0.39) and total nitrogen (R²=0.69, R²=0.67) in the pine plantation and the secondary forest, respectively. The regression analysis suggested that fungal biomass might have a greater influence on heterotrophic soil respiration in the pine plantation, while the bacterial biomass might have a greater influence in the secondary forest. Heterotrophic soil respiration was more sensitive to total N than to carbon in the light fraction, and soil moisture was a major factor influencing heterotrophic soil respiration in these forests where temperature is high and relatively invariable.     

Light, nutrients and the growth of herbaceous forest species

The herb layer of forests planted on former agricultural land often differs from that of old-growth forest. This study investigates if the expected increased nutrient availability in the shaded conditions of newly planted forests and the plasticity of the species to adjust their biomass allocation to different levels of light and nutrients help to explain these differences in the herb layers of the two forest types. In a greenhouse experiment biomass distribution and production of two species characteristic for the highly shaded forest floor, Circaea lutetiana and Mercurialis perennis, and two species more common in the forest-edge, Aegopodium podagraria and Impatiens parviflora were studied at different levels of light (2%, 8% and 66% of the full light level) and nutrients (30 and 300 kg N ha^–1 per year). The main factor affecting allocation and biomass production was light availability. Nutrient supply only had a significant effect at the higher light levels. Species were mainly plastic to changes in light and the two species from the forest floor showed to be more rigid in allocation pattern than the species from the forest-edge. So, although the species from the forest-edge were more plastic, they did not profit from the increased nutrient supply because the main factor affecting biomass distribution and production was light availability.     

The Effect of Shading on the Photosynthetic Rate and Longevity of Grass Leaves

The rate at which the net photosynthesis of grass leaves grownin bright light (119 W m^–2) decreased as they aged wasincreased by severe shading (to 21 W m^–2 or less). However,less severe shading (light intensities of 36 W m^–2 ormore) had no effect. The decrease in photosynthesis was unaffectedby whether the whole plant was shaded or only the leaf whosephotosynthesis was measured. In both shaded and unshaded leaves, photosynthesis measuredin bright light fell faster as the leaf aged than did photosynthesisin dim light. Both mesophyll and stomatal diffusion resistancesrose as the leaf aged but the former rose faster. The chlorophyllcontent fell only towards the end of the life of the leaves.     

Scaling of Pelagic Metabolism to Size,Trophy and Forest Cover in Small Danish Lakes

ABSTRACT We tested whether pelagic light and nutrient availability, metabolism, organic pools and CO₂-supersaturation were related to lake size and surrounding forest cover in late summer–autumn measurements among 64 small (0.02–20 ha), shallow seepage lakes located in nutrient-rich, calcareous moraine soils in North Zealand, Denmark. We found a strong implicit scaling to lake size as light availability increased significantly with lake size while nutrient availability, phytoplankton biomass and dissolved organic matter declined. Forest lakes had significantly stronger net heterotrophic traits than open lakes as higher values were observed for light attenuation above and in the water, dissolved organic matter, pelagic community respiration (R) relative to maximum gross primary production (R/GPP) and CO₂-supersaturation. Total-phosphorus was the main predictor of phytoplankton biomass (Chl) despite a much weaker relationship than observed in previous studies of larger lakes. Maximum gross primary production increased with algal biomass and decreased with dissolved organic matter, whereas community respiration increased with dissolved organic matter and particularly with gross primary production. These results suggest that exogenous organic matter supplements primary production as an energy source to heterotrophs in these small lakes, and particularly so in forest lakes experiencing substantial shading from the forest and dissolved humic material. This suggestion is supported by 20–30-fold CO₂ supersaturation in the surface water of the smallest forest lakes and more than sixfold supersaturation in 75% of all measurements making these lakes among the most supersaturated temperate lakes examined so far.