Multi‐taxon and forest structure sampling for identification of indicators and monitoring of old‐growth forest

Abstract

The most commonly used old‐growth forest indicators are structural attributes; nevertheless, they do not necessarily represent the biodiversity value of old‐growth forests. The aim of this study is to analyse the relationships between species richness data of different taxa and structural indicators of old‐growth and to identify taxonomic/functional groups, species and structural attributes that may be used as indicators of old‐growth. To achieve this goal we sampled forest structure, vascular plants, lichens, bryophytes, fungi, saproxylic beetles and birds in mature and old‐growth stands in southern Italy. We calculated Spearman’s correlation coefficients between species richness data and structural attributes. Analyses of indicator species, co‐occurrences and two‐way clusters were performed on the multi‐taxonomic list. The group of vascular plants most significantly correlated with other groups in terms of species richness; furthermore, it displays the highest proportion of between‐group co‐occurrences. The resulting multi‐taxonomic list of potential indicators may serve as an effective means of detecting and monitoring forest ecosystems; however, for this goal, structure‐based indicators, such as forest structural attributes and vascular plant species composition, are of primary importance.     

What controls the distribution of tropical forest and savanna?

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

Comparison of structure,regeneration and dead wood in virgin forest remnant and managed forest on Grmeč Mountain in Western Bosnia

This paper compares the forest structure, regeneration and distribution of dead wood in a virgin forest remnant and a close-to-nature managed beech–conifer mixture situated on Grme? Mountain in Western Bosnia. The investigations were carried out in a 1 ha permanent sample plot and 35 circular plots (20 m radius) in the virgin forest and in 17 circular plots (25 m radius) in managed forests. The number of trees in the managed forest was significantly (p = 0.05) higher than that in virgin forest and the distribution of the number of trees per diameter classes had a decreasing trend, but with a different shape in the virgin forest compared to the managed stands. In the lower diameter classes, the stock volume recorded in virgin forest was half of that in the managed forest, whilst for higher diameter classes the cumulated volume of the growing stock was almost double in virgin forest. The young crops had a significantly lower presence in the virgin forest and a larger volume of dead wood was identified in the virgin forest than in managed stands. The study results are important in assessing the consequences of close-to-nature management on the forest structure and regeneration when compared to the condition in virgin forests.     

Male competition and paternity in wild chimpanzees of the Taï forest

In social animals, competition among males for mates affects individual reproductive success. The priority‐of‐access model attempts to account for the influence of demographic conditions within groups upon male reproductive success, but empirical data for testing this model are scarce. Our long‐term study of chimpanzees in the Taï National Park, Côte d'Ivoire, encompasses a period of steady decrease in community size and fluctuating numbers of competing males and sexually receptive females. These demographic changes, in combination with genetic assessment of paternity for 48 offspring from three communities, allowed us to quantify the effects of varying levels of competition upon male reproductive success. On average, the highest‐ranking male sired 50% of all analyzed offspring during a 14‐year period from 1987–2000. Competition among males strongly decreased the relative reproductive success of the alpha male, such that the alpha male's rate of success decreased from 67% with few competitors to only 38% with four or more competitors. The increasing number of synchronously receptive females in large groups also reduced the proportion of paternities by the alpha male. Thus, patterns of paternity in Taï chimpanzees fit well the predictions of the priority‐of‐access model. We also found that despite the inability of dominants to monopolize reproduction, they achieved a higher reproductive rate in large multimale groups, because these have more females and a higher infant survival rate. Varied levels of male competition within communities seem to explain differences in the reproductive success of alpha males observed in different chimpanzee populations, and in other primate species. Am J Phys Anthropol, 2006. © 2005 Wiley‐Liss, Inc.     

Attitudes toward forest diversity and forest ecosystem services—a cross-cultural comparison between China and Switzerland

Aims Despite the current interest in services provided by ecosystems and the role of biodiversity, the relationship among human attitudes, biodiversity and ecosystem services has hardly been investigated. Moreover, few studies have examined attitudes toward nature in cross-cultural comparisons. This study investigates the attitudes of Chinese and Swiss people, both environmental experts and laypersons, toward forest biodiversity and ecosystem services.Methods Overall, 640 people in China and Switzerland were interviewed with the help of a standardized questionnaire. In each country, the study population was equally divided into an urban (80 city dwellers and 80 environmental science students) and a rural (80 forest visitors and 80 farmers) study group. The 15-minute interviews took place in the cities of Beijing and Zurich and in the rural forested areas of Dujiangyan, Sichuan Province and Lake Sempach, canton Lucerne. Attitudes toward forest biodiversity were investigated with the help of color photographs that depicted both monocultures and species-rich forests typical for China and Switzerland. Attitudes toward ecosystem services were investigated with the help of 13 statements on provisioning, regulating, cultural and supporting services of forests.Important findings On average, Chinese participants showed no strong preferences for biodiversity, whereas the Swiss clearly preferred species-rich forests over monocultures. However, Chinese environmental science students did prefer species-rich forests and attributed to them a higher conservation value because of their higher biodiversity. Although there were no strong preferences for Chinese versus Swiss forests, all participants correctly answered that Chinese forests are more species rich in terms of plants and animals and thus found them less boring and more interesting, but also less managed, than Swiss forests. All participants highly valued the ecosystem services provided by forests; especially the regulating and supporting ones. Environmental science students and farmers placed more importance on the provisioning services, whereas city dwellers and forest visitors emphasized more on the regulating services. The disjuncture between the high ecological quality of species-rich forests and their low attractiveness to Chinese study participants points to a potential conflict between conservation policies and the public's preferences. A better communication of ecosystem services provided by forest biodiversity to the public might change these preferences in favor of ecological quality, as already observed among Chinese environmental science students.     

Impacts of El Niño related drought and forest fires on sun bear fruit resources in lowland dipterocarp forest of East Borneo

Droughts and forest fires, induced by the El Niño/Southern Oscillation (ENSO) event, have increased considerably over the last decades affecting millions of hectares of rainforest. We investigated the effects of the 1997–1998 forest fires and drought, associated with an exceptionally severe ENSO event, on fruit species important in the diet of Malayan sun bears (Helarctos malayanus) in lowland dipterocarp forest, East Kalimantan, Indonesian Borneo. Densities of sun bear fruit trees (≥10 cm DBH) were reduced by ～80%, from 167±41 (SD) fruit trees ha^?1 in unburned forest to 37±18 fruit trees ha^?1 in burned forest. Densities of hemi-epiphytic figs, one of the main fallback resources for sun bears during periods of food scarcity, declined by 95% in burned forest. Species diversity of sun bear food trees decreased by 44% in burned forest. Drought also affected sun bear fruit trees in unburned primary forest, with elevated mortality rates for the duration of 2 years, returning to levels reported as normal in region in the third year after the ENSO event. Mortality in unburned forest near the burn-edge was higher (25±5% of trees ≥10 cm DBH dead) than in the forest interior (14±5% of trees), indicating possible edge effects. Combined effects of fire and drought in burned primary forest resulted in an overall tree mortality of 78±11% (≥10 cm DBH) 33 months after the fire event. Disturbance due to fires has resulted in a serious decline of fruit resources for sun bears and, due to the scale of fire damage, in a serious decline of prime sun bear habitat. Recovery of sun bear populations in these burned-over forests will depend on regeneration of the forest, its future species composition, and efforts to prevent subsequent fire events.     

Resilience of New?Zealand indigenous forest fragments to impacts of livestock and pest mammals

A number of factors have combined to diminish ecosystem integrity in New?Zealand indigenous lowland forest fragments surrounded by intensively grazed pasture. Livestock grazing, mammalian pests, adventive weeds and altered nutrient input regimes are important drivers compounding the changes in fragment structure and function due to historical deforestation and fragmentation. We used qualitative systems modelling and empirical data from Beilschmiedia tawa dominated lowland forest fragments in the Waikato Region to explore the relevance of two common resilience paradigms ? engineering resilience and ecological resilience ? for addressing the conservation management of forest fragments into the future. Grazing by livestock and foraging/predation by introduced mammalian pests both have direct detrimental impacts on key structural and functional attributes of forest fragments. Release from these perturbations through fencing and pest control leads to partial or full recovery of some key indicators (i.e. increased indigenous plant regeneration and cover, increased invertebrate populations and litter mass, decreased soil fertility and increased nesting success) relative to levels seen in larger forest systems over a range of timescales. These changes indicate that forest fragments do show resilience consistent with adopting an engineering resilience paradigm for conservation management, in the landscape context studied. The relevance of the ecological resilience paradigm in these ecosystems is obscured by limited data. We characterise forest fragment dynamics in terms of changes in indigenous species occupancy and functional dominance, and present a conceptual model for the management of forest fragment ecosystems.     

Formation of forest gaps accelerates C,N and P release from foliar litter during 4 years of decomposition in an alpine forest

High-latitude boreal and arctic surface/inland waters contain sizeable reservoirs of dissolved organic matter (DOM) and trace elements (TE), which are subject to seasonal freezing. Specifically, shallow ponds and lakes in the permafrost zone often freeze solid, which can lead to transformations in the colloidal and dissolved fractions of DOM and TE. Here, we present results from experimental freeze-thaw cycles using iron (Fe)- and DOM-rich water from thaw ponds situated in Stordalen and Storflaket palsa mires in northern Sweden. After ten cycles of freezing, 85% of Fe and 25% of dissolved organic carbon (DOC) were removed from solution in circumneutral fen water (pH 6.9) but a much smaller removal of Fe and DOC (< 7%) was found in acidic bog water (pH 3.6). This removal pattern was consistent with initial supersaturation of fen water with respect to Fe hydroxide and a lack of supersaturation with any secondary mineral phase in the bog water. There was a nearly two- to threefold increase in the low-molecular-weight (LMW) fraction of organic carbon (OC) and several TEs caused by the repeated freeze-thaw cycles. Future increases in the freeze-thaw frequency of surface waters with climate warming may remove up to 25% of DOC in circumneutral organic-rich waters. Furthermore, an increase of LMW OC may result in enhanced carbon dioxide losses from aquatic ecosystems since this fraction is potentially more susceptible to biodegradation.     

Restoration of beech forest for saproxylic beetles—effects of habitat fragmentation and substrate density on species diversity and distribution

The influence of spatial location and density of beech snags on species diversity and distribution patterns of saproxylic beetles was studied in a 2,400 ha forest landscape in southern Sweden. Complete snag surveys were combined with a beetle survey using small window traps directly attached to the beech snags. The density of beech snags ≥30 cm dbh varied between one and seven snags per ha within the study area, corresponding to 1.1–5.1 m³/ha. A total of 2,610 specimens of 180 saproxylic beetles species were trapped, of which 19 species were red-listed. Within the study area, the number of red-listed and formerly red-listed species was highest around traps in old-growth stands, intermediate in managed stands contiguous with old-growth and lowest in managed stands isolated from old-growth by a two km-wide zone without beech forest. Logistic regressions revealed negative relationships between distance to old-growth forest and occurrence of eleven species, among them six red-listed or formerly red-listed species. The number of non red-listed species was not correlated with isolation from old-growth forest. The number of red-listed species also increased with snag density within 200–300 m around the traps. Our results suggest that red-listed species generally have a lower dispersal capacity than other saproxylic beetles. We conclude that retention of dead wood close to existing populations is more beneficial for red-listed species than an even distribution of snags across the forest landscape.     

Monkeys and matrices: On the coexistence of “omnivorous” forest primates

Summary Members of competing species may influence their share of food sources either by finding new feeding sites more frequently or by more effectively monopolizing those sites once found. We describe a model that separates the effects of these two forms of competition; in a set of African forest primates, only the first has a major effect on species' relative population densities. The model identifies factors which might lead to a greater role of interference at food sources in other communities.     

Composition and diversity of bird communities in Swedish farmland–forest mosaic landscapes

CapsuleThe amount of forest (at local and landscape scales) and occurrence of residual habitats at the local scale are shown to be the major factors influencing bird community composition in farmland–forest landscapes in central Sweden.

Aims To investigate the importance of local habitat and landscape structure for breeding birds in farmland–forest landscapes in central Sweden.

Methods Breeding birds were censused at 292 points. A detailed habitat mapping was made within 300 m of the points. Within a 300–600 m radius only two major habitats (forests and arable fields) were identified.

Results Cluster analyses of bird communities identified three site types that also differed in habitat composition: (i) partially forested sites in forested landscapes; (ii) heterogeneous sites with residual habitats in mosaic landscapes; and (iii) field-dominated farmland sites in open landscapes. A total of 19 of 25 farmland bird species (restricted to farmland or using both farmland and forest) had the highest abundance in farmland sites with mosaics of forest and farmland, while only six farmland species had the highest abundance in field-dominated sites. The bird community changed from being dominated by farmland species to being dominated by forest species (common in forest landscapes without farmland) at small proportions (10–20%) of forest at the local scale. A major difference in habitat composition between heterogeneous and field-dominated sites was the occurrence of different residual habitats (e.g. shrubby areas and seminatural grasslands). These habitats seemed to influence bird community composition more than land-use, despite covering <10% of the area. Seminatural grasslands were important for bird community composition and species-richness, but grazing seemed to be less important. Among different land-use types, cereal crops were the least preferred fields. Set-asides with tall vegetation and short rotation coppices were positively associated with species-richness of farmland birds.

Conclusion In general, the composition of the landscape was important for bird community composition, although amount and distribution of forests, occurrence of residual habitats and land-use of fields at the local scale had the strongest influence on bird community composition. The possible implications of these patterns for managing farmland–forest landscapes are discussed.     

Twenty-five years of change in forest structure and nesting behavior of Hawaiʻi ʻelepaio

Long-term ecological studies are invaluable for detecting changes over time. Forest restoration has been a conservation priority in Hawaiʻi, where invasive species have negatively impacted native bird habitat. During 1993–1994, a study was conducted of Hawaiʻi ʻelepaio (Chasiempis sandwichensis) nest site selection and forest composition in mesic montane forest along Mauna Loa Road in Hawaiʻi Volcanoes National Park. We returned to the site and repeated the methods used in the earlier study to record Hawaiʻi ʻelepaio nest site selection (tree species, tree and nest height, and tree girth [diameter at breast height, DBH]) from 2015 to 2017, and measured forest composition (tree density, relative abundance, height, and DBH) in 2019. Three times as many nests were found in ʻaʻaliʻi (Dodonaea viscosa) as in koa (Acacia koa) in both time periods. Heights of koa and ʻaʻaliʻi did not change, but their DBH increased over time. The relative height of nests in ʻaʻaliʻi trees did not change, but nests in koa were placed higher in the crown during the later study. Overall tree density increased from 1,619 (95% confidence interval [CI] = 1,499–1,740) trees/ha to 2,583 (95% CI = 2,176–3,096) trees/ha. Koa relative abundance was 53% of total trees earlier and 45% later, ʻaʻaliʻi remained at 47%, and māmane (Sophora chrysophylla) increased from 0 to 8% over time. Our results indicate that changes in forest composition affect nesting behavior, but in ways that are not necessarily simple or consistent.     

Quantitative classification and carbon density of the forest vegetation in Lüliang Mountains of China

Forests play a major role in global carbon (C) cycle, and the carbon density (CD) could reflect its ecological function of C sequestration. Study on the CD of different forest types on a community scale is crucial to characterize in depth the capacity of forest C sequestration. In this study, based on the forest inventory data of 168 field plots in the study area (E 111°30′–113°50′, N 37°30′–39°40′), the forest vegetation was classified by using quantitative method (TWINSPAN); the living biomass of trees was estimated using the volume-derived method; the CD of different forest types was estimated from the biomass of their tree species; and the effects of biotic and abiotic factors on CD were studied using a multiple linear regression analysis. The results show that the forest vegetation in this region could be classified into 9 forest formations. The average CD of the 9 forest formations was 32.09 Mg ha⁻¹ in 2000 and 33.86 Mg ha⁻¹ in 2005. Form. Picea meyeri had the highest CD (56.48 Mg ha⁻¹), and Form. Quercus liaotungensis + Acer mono had the lowest CD (16.14 Mg ha⁻¹). Pre-mature forests and mature forests were very important stages in C sequestration among four age classes in these formations. Forest densities, average age of forest stand, and elevation had positive relationships with forest CD, while slope location had negative correlation with forest CD.     

An interplay between forest and non‐forest South American avifaunas suggested by a phylogeny of lepidocolaptes woodcreepers (dendrocolaptinae)

Very few South American avian superspecies or species groups are composed of both forest and non‐forest taxa. The genus Lepidocolaptes comprises 8–9 species of woodcreepers, most of which are forest birds, but two species, L. angustirostris and L. souleyetii, inhabit open vegetations. Therefore, this genus should play an important role in the discussion about the relationships between forest and non‐forest South American avifaunas. Nucleotide sequences from two mitochondrial genes, cytochrome b and ND2, suggest that: (i) L. fuscus should be removed from the genus since its association with other members of this genus is poorly supported. This view has been pointed out also by morphological and behavioural data; (ii) the phylogenetic position of the open‐vegetation species within the Lepidocolaptes radiation indicate that the split between forest and non‐forest elements within this genus took place as recently as two million years ago. This result suggests that the evolutionary relationships between forest and non‐forest biotas in South America may have been more dynamic than previously thought.     

Changes in stable isotopic signatures of soil nitrogen and carbon during 40 years of forest development

Understanding what governs patterns of soil δ¹⁵N and δ¹³C is limited by the absence of these data assembled throughout the development of individual ecosystems. These patterns are important because stable isotopes of soil organic N and C are integrative indicators of biogeochemical processing of soil organic matter. We examined δ¹⁵N of soil organic matter (δ¹⁵N_SOM) and δ¹³C_SOM of archived soil samples across four decades from four depths of an aggrading forest in southeastern USA. The site supports an old-field pine forest in which the N cycle is affected by former agricultural fertilization, massive accumulation of soil N by aggrading trees over four decades, and small to insignificant fluxes of N via NH₃ volatilization, nitrification, and denitrification. We examine isotopic data and the N and C dynamics of this ecosystem to evaluate mechanisms driving isotopic shifts over time. With forest development, δ¹³C_SOM became depth-dependent. This trend resulted from a decline of ~2‰ in the surficial 15 cm of mineral soil to −26.0‰, due to organic matter inputs from forest vegetation. Deeper layers exhibited relatively little trend in δ¹³C_SOM with time. In contrast, δ¹⁵N_SOM was most dynamic in deeper layers. During the four decades of forest development, the deepest layer (35–60 cm) reached a maximum δ¹⁵N value of 9.1‰, increasing by 7.6‰. The transfer of >800 kg ha⁻¹ of soil organic N into aggrading vegetation and the forest floor and the apparent large proportion of ectomycorrhizal (ECM) fungi in these soils suggest that fractionation via microbial transformations must be the major process changing δ¹⁵N in these soils. Accretion of isotopically enriched compounds derived from microbial cells (i.e., ECM fungi) likely promote isotopic enrichment of soils over time. The work indicates the rapid rate at which ecosystem development can impart δ¹⁵N_SOM and δ¹³C_SOM signatures associated with undisturbed soil profiles.     

The mid-Holocene extinction of silver fir (Abies alba) in the Southern Alps: a consequence of forest fires? Palaeobotanical records and forest simulations

Pollen records suggest that Abies alba played a dominating role in both the montane and lowland forests at the border of the Southern Alps between ca. 8500 and 5700 years ago. Two major declines in fir, at about 7300–7000 cal b.p. and at ca. 6000 cal b.p., followed by the local extinction of the species are characteristic of the area below ca. 1000 m a.s.l. In order to test the impact of fire on the population dynamics of silver fir, a dynamic model (DisCForm) with a fire module was applied to simulate the early- and mid-Holocene forest development. Simulation outputs based on different fire scenarios were compared with the pollen record from Lago di Annone (226 m a.s.l.). The marked Abies decreases shown in the pollen record can be simulated with very intensive fire scenarios, but they do not result in an extinction of silver fir in the model. Low charcoal influx values related to the Abies declines in the palaeobotanical record suggest that fire was not the only reason for the extinction of silver fir. Human impact, as well as Holocene climatic changes leading to temporary moisture deficits and reduced adaptability due to low genetic variation may have had a significant impact on the Abies forests.     

Do food protein and carbohydrate content influence the pattern of feeding and the tendency to explore of forest tent caterpillars?

This study examines whether the ratio of protein to carbohydrate affects the timing of meals and the propensity to explore of forest tent caterpillars (Malacosoma disstria). The behavior of fourth instar caterpillars was observed on three semi-defined artificial diets varying in protein (p)-carbohydrate (c) ratio. These diets were (a) p14:c28, (b) p28:c14, and (c) p35:c7. The probability of initiating feeding at first contact with the food and the duration of the first feeding event did not vary across diets, suggesting not much difference in phagostimulatory power. There was also no difference in the total time spent eating, at rest and in motion between diets. However, the timing and duration of meals varied significantly; more short meals were observed on the carbohydrate-biased diet. The duration of pauses between meals also increased with food protein content. Furthermore, caterpillars on the carbohydrate-biased diet were more likely to leave the trail leading to the known food source and to discover a second food source, suggesting that protein deprivation promotes exploration. These findings shed insight into the physiological responses to protein and carbohydrate ingestion and demonstrate how post-ingestive effects can favor consumption of foods containing protein without invoking an explicit mechanism of independent nutrient regulation, but simply by influencing the pattern of feeding and the propensity to explore.     

Leaf defoliation of tropical dry forest tree seedlings – implications for survival and growth

 The effect of herbivory on survival and growth of seedlings of four species, Cedrela odorata, Hymenaea courbaril, Manilkara chicle and Swietenia macrophylla, was studied in secondary dry forests in Guanacaste, Costa Rica. Potted seedlings were planted at two sites in a 2×2 factorial design, combining thinning to increase light levels at the forest floor, and trenching to reduce root competition around the planted seedlings. C. odorata and S. macrophylla were repeatedly severely defoliated by insects, while H. courbaril became less affected. M. chicle did not show any significant signs of defoliation and was not further analysed. Defoliation levels were generally higher in both thinned and trenched treatments, and also positively correlated with larger initial seedlings sizes. Decreased growth rates caused by defoliation were seen in S. macrophylla and C. odorata in the growing season. Defoliation of more than 50% in combination with abiotic factors, particularly drought, were sufficient to contribute to retarded seedling development and increased seedling mortality of C. odorata and S. macrophylla. Received: 10 April 1997 / Accepted: 26 June 1998     

Enantioselective kinetics of α-hexachlorocyclohexane in earthworm (Eisenia fedtia) and forest soil

The enantioselective bioaccumulation and elimination behaviors of α-hexachlorocyclohexane (α-HCH) enantiomers in earthworm and soil were investigated by chiral gas chromatography. Enantiomer fraction values were calculated as indicators of the enantioselectivity. The mature earthworms were exposed to 0.10 μg g(-1)(wwt) (0.14 μg g(-1)(dwt)) spiked soil continuously for the bioaccumulation, and the elimination was conducted after an enrichment period in the soil. The results showed that both the bioaccumulation and elimination processes followed monophasic kinetics, body residues of α-HCH in earthworm increased to high level at the fifth day, and enantioselectivity was found in the bioaccumulation process with the rate constant (k) of 0.80 d(-1) for (+)-α-HCH and 0.74 d(-1) for (-)-α-HCH. The half life (t(1/2)) of the enantiomers obtained in the elimination process was within one day. The bioaccumulation factors of steady state of α-HCH enantiomers were 2.82 for (+)-α-HCH and 2.75 for (-)-α-HCH. The enantiomer fractions of earthworm and soil obviously below 0.5 during uptake and elimination processes indicate significant enantioselectivity and preferential depuration of (+)-α-HCH in earthworm. However, earthworms do not have a great capacity for getting rid of α-HCH in polluted soil shown by a contradistinctive experiment.