Inter-annual dynamics and persistence of small mammal communities in a selectively logged tropical forest in Borneo

Understanding temporal change and long-term persistence of species and communities is vital if we are to accurately assess the relative values of human-modified habitats for biodiversity. Despite a large literature and emerging consensus demonstrating a high conservation value of selectively logged tropical rainforests, few studies have taken a long-term perspective. We resampled small mammals (≤?1 kg) in a heavily logged landscape in Sabah, Borneo between 2011 and 2016 to investigate temporal patterns of species-level changes in population density. We found that small mammal population density in heavily logged forest was highly variable among years, consistent with patterns previously observed in unlogged forest, and uncovered evidence suggesting that one species is potentially declining towards local extinction. Across nine species, population densities varied almost sevenfold during our 6-year study period, highlighting the extremely dynamic nature of small mammal communities in this ecosystem. Strictly terrestrial murid species tended to exhibit strong temporal dynamics, whereas semi-arboreal foraging species such as treeshrews had more stable dynamics. We found no relationships between population density and fruit/seed mass, and therefore no evidence that our patterns represent responses to inter-annual mast fruiting of the dominant canopy dipterocarp trees. This may be due to the removal of most of the canopy during logging, and hence the dipterocarp seed resource, although it possibly also reflects spatiotemporal limitations of our data. Our results underline the importance of understanding long-term variability in animal communities before developing conservation and management recommendations for human-altered ecosystems.     

Biomass and carbon dynamics of a tropical mountain rain forest in China

Biometric inventories for 25 years, from 1983 to 2005, indicated that the Jianfengling tropical mountain rain forest in Hainan, China, was either a source or a modest sink of carbon. Overall, this forest was a small carbon sink with an accumulation rate of (0.56±0.22) Mg C ha⁻¹yr⁻¹, integrated from the long-term measurement data of two plots (P9201 and P8302). These findings were similar to those for African and American rain forests ((0.62±0.23) Mg C ha⁻¹yr⁻¹). The carbon density varied between (201.43±29.38) Mg C ha⁻¹ and (229.16±39.2) Mg C ha⁻¹, and averaged (214.17±32.42) Mg C ha⁻¹ for plot P9201. Plot P8302, however, varied between (223.95±45.92) Mg C ha⁻¹ and (254.85±48.86) Mg C ha⁻¹, and averaged (243.35±47.64) Mg C ha⁻¹. Quadratic relationships were found between the strength of carbon sequestration and heavy rainstorms and dry months. Precipitation and evapotranspiration are two major factors controlling carbon sequestration in the tropical mountain rain forest.     

Variation in wood density determines spatial patterns inAmazonian forest biomass

Uncertainty in biomass estimates is one of the greatest limitations to models of carbon flux in tropical forests. Previous comparisons of field‐based estimates of the aboveground biomass (AGB) of trees greater than 10 cm diameter within Amazonia have been limited by the paucity of data for western Amazon forests, and the use of site‐specific methods to estimate biomass from inventory data. In addition, the role of regional variation in stand‐level wood specific gravity has not previously been considered. Using data from 56 mature forest plots across Amazonia, we consider the relative roles of species composition (wood specific gravity) and forest structure (basal area) in determining variation in AGB. Mean stand‐level wood specific gravity, on a per stem basis, is 15.8% higher in forests in central and eastern, compared with northwestern Amazonia. This pattern is due to the higher diversity and abundance of taxa with high specific gravity values in central and eastern Amazonia, and the greater diversity and abundance of taxa with low specific gravity values in western Amazonia. For two estimates of AGB derived using different allometric equations, basal area explains 51.7% and 63.4%, and stand‐level specific gravity 45.4% and 29.7%, of the total variation in AGB. The variation in specific gravity is important because it determines the regional scale, spatial pattern of AGB. When weighting by specific gravity is included, central and eastern Amazon forests have significantly higher AGB than stands in northwest or southwest Amazonia. The regional‐scale pattern of species composition therefore defines a broad gradient of AGB across Amazonia.     

湖北省主要森林类型生态系统生物量与碳密度比较

利用野外调查数据对湖北省封山育林下的次生林、次生林、人工林森林生态系统碳密度进行了分析,结果表明:封山育林下的次生林、次生林和人工林生态系统乔木层平均碳密度分别为133.87、73.42和111.62t·hm-2,灌木层平均碳密度分别为1.65、1.40和1.52t·hm-2,草本层平均碳密度分别为0.13、0.09和0.13t·hm-2,枯落物层平均碳密度分别为0.47、1.34和0.93t·hm-2,乔木层碳密度作为生态系统碳储量的主要贡献者占总生物碳密度的98.35%、96.29%和97.74%,林下植被(灌木层和草本层)碳密度分别占1.31%、1.95%和1.44%,凋落物层碳密度分别占0.34%、1.76%和0.82%。土壤(0～100cm)碳密度平均值分别为57.04、66.92和54.12t·hm-2,土壤碳密度的60%储存在0～40cm土壤中,并随土层深度增加,各层次土壤碳密度逐渐减少。森林生态系统的乔木层、灌木层、草本层、凋落物层生物量和土壤层碳密度均表现出:封山育林下的次生林、次生林大于人工林。封山育林下的次生林、次生林和人工林碳密度分布序列为土壤(0～100cm)>乔木层>灌木层>草本层>枯落物层。可见,封山育林下的次生林更有助于提高森林碳汇,实施近自然林经营是提升该区域森林碳汇能力的重要途径。     

The relationship between wood density and mortality in a global tropical forest data set

Wood density is thought to be an important indicator of plant life history because it is coupled to many aspects of whole-plant form and function. We used a hierarchical Bayesian approach to explain variation in mortality rates with wood density, drawing on data for 765,500 trees from 1639 species at 10 sites located across the Old and New World tropics. Mortality rates declined with increasing wood density at five of 10 sites. Similar negative trends were detected at four additional sites, while one site showed no relationship. Our model explained 40% of variation in mortality on average. Both wood density and mortality rates show a high degree of phylogenetic conservatism. Grouping species by family across sites in a second analysis, we found considerable variation in the relationship between wood density and mortality, with 10 of 27 families demonstrating a strong negative relationship. Our results highlight the importance of wood density as a functional trait in tropical forests, as it is strongly linked to variation in survival. However, the relationship varied among families, plots, and even census intervals within sites, indicating that the factors responsible for the relationship between wood density and mortality vary spatially, taxonomically and temporally.     

Spatial dynamics of forest decline: the role of neighbouring trees

Abstract. Changes in horizontal structure and the influence of neighbours on tree vitality were studied in a spruce forest under air pollution stress. Five permanent plots along an altitudinal gradient in the Krkono?e Mts., Czech Republic, were monitored for 18 years. Digitized maps of each plot were used for the analysis: the health of each tree, expressed by the defoliation degree was recorded each year, the biometrical characteristics were measured at five-year intervals. Various indices of neighbourhood competition were used to evaluate the interference with neighbours. The results show that the suppressed trees are most susceptible to other environmental stresses, particularly to the air pollution stress. Similarly, tree damage is more severe in plots near the natural timberline, where the trees are close to their natural environmental limits. The spatial pattern of surviving trees changes towards regularity at a scale of 2–5 m (K-function analysis) when the suppressed trees, usually those with close neighbours, die.     

Survey of Hylobates agilis albibarbis in a logged peat-swamp forest: Sabangau catchment, Central Kalimantan

Few data are available on gibbon populations in peat-swamp forest. In order to assess the importance of this habitat for gibbon conservation, a population of Hylobates agilis albibarbis was surveyed in the Sabangau peat-swamp forest, Central Kalimantan, Indonesia. This is an area of about 5,500 km² of selectively logged peat-swamp forest, which was formally gazetted as a national park during 2005. The study was conducted during June and July 2004 using auditory sampling methods. Five sample areas were selected and each was surveyed for four consecutive days by three teams of researchers at designated listening posts. Researchers recorded compass bearings of, and estimated distances to, singing groups. Nineteen groups were located. Population density is estimated to be 2.16 (±0.46) groups/km². Sightings occurring either at the listening posts or that were obtained by tracking in on calling groups yielded a mean group size of 3.4 individuals, hence individual gibbon density is estimated to be 7.4 (±1.59) individuals/km². The density estimates fall at the mid-range of those calculated for other gibbon populations, thus suggesting that peat-swamp forest is an important habitat for gibbon conservation in Borneo. A tentative extrapolation of results suggests a potential gibbon population size of 19,000 individuals within the mixed-swamp forest habitat sub-type in the Sabangau. This represents one of the largest remaining continuous populations of Bornean agile gibbons. The designation of the Sabangau forest as a national park will hopefully address the problem of illegal logging and hunting in the region. Further studies should note any difference in gibbon density post protection.     

Unfavorable successional pathways and the conservation value of logged tropical forest

Conservation agencies are increasingly advocating that the survival of many tropical forest species may depend on disturbed forests and are directing scarce conservation resources to managing logged forests. This requires critical evaluation. In this 10-year study, we quantify tree community dynamics in three selectively logged areas harvested at different intensities and compare their recovery to two unlogged areas in Kibale National Park, Uganda. Over the 10 years, 527 of the original 4840 tagged trees (10cm DBH) died; while the mortality rate was highest in the heavily logged area, it was only significantly greater than one of the unlogged sites. The density of new trees varied significantly among areas being highest in the lightly logged area, but the density of new recruits in the heavily logged area did not differ from the unlogged areas. Overall, the more heavily logged areas had higher growth rates than the unlogged or lightly logged areas. However, there was no difference among areas in the magnitude of the change in basal area; and in both 1990 (20+ years post-harvest) and 2000 the basal area in the heavily logged area was significantly less than the unlogged areas. In general, findings from the heavily logged area suggest that the expectation that in logged areas a number of processes of forest regeneration will be accelerated relative to unlogged forest was not met. Thus, 30+ years after logging evidence suggests that forest recovery will be slower than expected. We also present evidence to suggest that the climate in the region is changing, which may account for changes in the population dynamics of certain species in the unlogged forests.     

Biomass and litter dynamics in a Melaleuca forest on a seasonally inundated floodplain in tropical,northern Australia

Litterfall from a Melaleuca forest was investigated as part of chemical cycling studies on the Magela Creek floodplain in tropical, northern Australia. The forest contained two species of tree, Melaleuca cajaputi and Melaleuca viridiflora, with a combined average density of 294 trees ha^–1. The M. viridiflora trees had diameter breast height measurements ranging from 11.8 to 62.0 cm, median class 25.1–30.0cm and a mean value of 29.2±1.0 cm, compared to 13.0 to 66.3 cm, 30.1–35.0cm and 33.5±1.0cm for M. cajaputi trees. A regression model between tree height, diameter breast height and fresh weight was determined and used to calculate average tree weights of 775±1.6kg for M. viridiflora and 1009±1.6kg for M. cajaputi, and a total above-ground fresh weight of 263±0.3t ha^–1. The weight of litter recorded each month on the ground beneath the tree canopy ranged from 582±103 to 2176±376 g m^–2 with a monthly mean value of 1105±51 g m^–2. The coefficient of variation of 52% on this mean indicates the large spatial and temporal variability in litter distribution over the study site. This variability was greatly affected by the pattern of water flow and litter transport during the Wet season. Litterfall from the trees was evaluated using two techniques - nets and trays. The results from these techniques were not significantly different with annual litterfall collected in the nets being 705 ± 25 g m^–2 and in the trays 716±49 g m^–2. The maximum monthly amount of litterfall, 108 ±55g m^–2, occurred during the Dry season months of June–July. Leaf material comprised 70% of the total annual weight of litter, 480±29 g m^–2 in the nets and 495 ± 21 g m^–2 in the trays. The tree density and weight of litter suggest that the Melaleuca forests are highly productive and contribute a large amount of material to the detrital/debris turnover cycle on the floodplain.     

Above-ground forest biomass is not consistently related to wood density in tropical forests

Aim  It is increasingly accepted that the mean wood density of trees within a forest is tightly coupled to above-ground forest biomass. It is unknown, however, if a positive relationship between forest biomass and mean community wood density is a general phenomenon across forests. Understanding spatial variation in biomass as a function of wood density both within and among forests is important for predicting changes in stored carbon in response to global change, and here we evaluated the generality of a positive biomass–wood density relationship within and among six tropical forests.
Location  Costa Rica, Panama, Puerto Rico and Ecuador.
Methods  Individual stem data, including diameter at breast height and spatial position, for six forest dynamics plots were merged with an extensive wood density database. Individual stem biomass values were calculated from these data using published statistical models. Total above ground biomass, total basal area and mean community wood density were also quantified across a range of subcommunity plot sizes within each forest.
Results  Among forests, biomass did not vary with mean community wood density. The relationship between subcommunity biomass and mean wood density within a forest varied from negative to null to positive depending on the size of subcommunities and forest identity. The direction of correlation was determined by the associated total basal area–mean wood density correlation, the slope of which increased strongly with whole forest mean wood density.
Main conclusions  There is no general relationship between forest biomass and wood density, and in some forests, stored carbon is highest where wood density is lowest. Our results suggest that declining wood density, due to global change, will result in decreased or increased stored carbon in forests with high or low mean wood density, respectively.     

Butterfly species and traits associated with selectively logged forest in Borneo

Logging can significantly change the structure of rainforest communities. To better understand how logging drives this change, butterflies and environmental variables were assessed within both unlogged and logged forest in Indonesian Borneo. In the whole dataset, we found local environmental variables and geographic distance combined captured 53.1% of the variation in butterfly community composition; 29.6% was associated with measured local environmental variables, 13.6% with geographic distance between sites, and 9.9% with covariation between geographic distance and environmental variables. The primary axis of variation in butterfly community composition represented a disturbance gradient from unlogged to logged forest. Subsequent axes represented gradients influenced by variables such as canopy cover and total tree density. There were significant associations between environmental variables and geographic range and larval host plant use of species. Specifically, butterflies using trees as larval host plants and those with distributions limited to Borneo were more likely to be present in unlogged forest. By contrast, species that tended to be more abundant in logged forest were those with widespread distributions and those using lianas and grasses as larval host plants. The results of this study highlight the importance of environmental variables and disturbance, e.g., selective logging, in structuring rainforest community diversity. Moreover, they confirm how species traits, such as larval food use and geographic distributions can determine patterns of species abundance following environmental change.     

Soil organic matter dynamics under decaying wood in a subtropical wet forest: effect of tree species and decay stage

Decaying wood is an important structural and functional component of forests: it contributes to generate habitat diversity, acts as either sink or source of nutrients, and plays a preponderant role in soil formation. Thus, decaying wood might likely have measurable effects on chemical properties of the underlying soil. We hypothesized that decaying wood would have a stronger effect on soil as decomposition advances and that such effect would vary according to wood quality. Twenty logs from two species with contrasting wood properties (Dacryodes excelsa Vahl. and Swietenia macrophylla King) and at two different decay stages (6 and 15 years after falling) were selected, and soil under and 50 cm away from decaying logs was sampled for soil organic matter (SOM) fractions [NaOH-extractable and water-extractable organic matter -(WEOM)] and properties (WEOM aromaticity). NaOH-extractable C and WEOM were higher in the soil influenced by 15-year-old logs, while the degree of aromaticity of WEOM was higher in the soil influenced by the 6-year-old logs. Decaying logs did influence properties of the underlying soil with differing effects according to the species since there was more NaOH-extractable C in the soil associated to D. excelsa logs and more WEOM in the soil associated to S. macrophylla older logs. It is proposed that such effects occurred through changes in the relative quantity and quality of different SOM fractions, as influenced by species and advancement in decomposition. Through its effect on SOM and nutrient dynamics, decaying wood can contribute to the spatial heterogeneity of soil properties, and can affect process of soil formation and nutrient cycling. Responsible Editor: Barbara Wick.     

Co-dominance and succession in forest dynamics: the role of interspecific differences in crown transmissivity

Forests that are composed of two or more tree species with similar ecological strategies appear to contradict the competitive exclusion principle. Beech-maple communities are a well-known example of such a system. On a local scale, a number of mechanisms have been proposed to explain the coexistence of these two species. These are reciprocal replacement, external factors that favour alternatively one or the other species and demographic stochasticity. This paper presents and analyses a simple mathematical model that shows that external factors are not an essential requirement for coexistence. Rather, coexistence requires interspecific differences in light transmissivity through the crowns of adult trees. However, all the three mechanisms mentioned above can be interpreted within the framework of the model. Furthermore, many models of forest dynamics make use of shade tolerance as a key feature in describing successional dynamics. Despite its importance, however, shade tolerance does not have a commonly accepted quantitative definition. Here, a simple scheme is proposed where the relationship between shade tolerance, individual traits (growth and survival) and successional status is defined. This might have important implications in understanding the overall dynamics. Theoretical results have been compared with a number of studies carried out in North American forests. In particular, coexistence in beech-maple communities and the relation between shade tolerance and successional status in a beech-hemlock-birch community have been discussed.     

Hydraulic failure and tree dieback are associated with high wood density in a temperate forest under extreme drought

Catastrophic hydraulic failure will likely be an important mechanism contributing to large‐scale tree dieback caused by increased frequency and intensity of droughts under global climate change. To compare the susceptibility of 22 temperate deciduous tree and shrub species to hydraulic failure during a record drought in the southeastern USA, we quantified leaf desiccation, native embolism, wood density, stomatal conductance and predawn and midday leaf water potential at four sites with varying drought intensities. At the two driest sites, there was widespread leaf wilting and desiccation, and most species exhibited predawn leaf water potentials of ≤3 MPa and >60% loss of xylem conductivity in branches. Although species with high wood density were more resistant to cavitation, they had higher levels of native embolism and greater canopy dieback than species with low wood density. This unexpected result can be explained by the failure of species with dense wood to avert a decline in water potential to dangerous levels during the drought. Leaf water potential was negatively correlated with wood density, and the relationship was strongest under conditions of severe water deficit. Species with low wood density avoided catastrophic embolism by relying on an avoidance strategy that involves partial drought deciduousness, higher sensitivity of stomata to leaf water potential and perhaps greater rooting depth. These species therefore maintained water potential at levels that ensured a greater margin of safety against embolism. These differences among species may mediate rapid shifts in species composition of temperate forests if droughts intensify due to climate change.     

Tree growth traits and social status affect the wood density of pioneer species in secondary subtropical forest

Wood density (WD) is not only an important parameter to estimate aboveground biomass but also an indicator of timber quality and plant adaptation strategies to stressful conditions (i.e., windthrow, pests, and pathogens). This study had three objectives: (1) to compare WD among seven subtropical tree species; (2) to determine how tree growth traits may influence possible differences in WD between the pioneer and shade‐tolerant species; and (3) to examine whether or not WD differs by tree social status (dominant vs. suppressed trees) within species. To do this, 70 trees were destructively harvested. From each tree, disks at different stem heights were obtained and subjected to a method of stem analysis to measure whole tree level WD. The results showed that WD differed significantly among the seven species (p < .001). Their average WD was 0.537 g/cm³, ranging from 0.409 g/cm³ for Choerospondias axillaris to 0.691 g/cm³ for Cyclobalanopsis glauca. The average WD of the four pioneer species (0.497 ± 0.13 g/cm³) was significantly lower (p < .01) than that of the three shade‐tolerant species (0.589 ± 0.12 g/cm³). The WD of the pioneers had a significant positive correlation with their stem diameter at breast height (DBH), tree height (H), and tree age, but WD had a significant negative correlation with relative growth rate (RGR). In contrast, the WD of the shade‐tolerant tree species had no significant relationships with DBH, H, tree age, or RGR. The dominant trees of the pioneer species had a higher WD than the suppressed trees, whereas the shade‐tolerant species had a lower WD for dominant trees than the suppressed trees. However, the differences in WD between dominant and suppressed trees were not significant. Taken together, the results suggest that classifying species into pioneer and shade‐tolerant groups to examine the effects of tree growth traits and social status could improve our understanding of intra‐ and interspecific variation in WD among subtropical tree species.     

The limited role of canopy gaps in the successional dynamics of a mature mixed Quercus forest remnant

Question: Are canopy gap dynamics responsible for driving the structural and compositional changes that have occurred over a 26‐year period in a mature Quercus forest remnant? Location: Dobbs Natural Area, an unlogged 3.6‐ha forest preserve in west‐central Indiana, USA. Methods: We analyzed mapped permanent plot data for a site that illustrates a trend common in Quercus‐dominated forests in eastern North America, where recruitment of new stems is dominated by mesophytic, shade‐tolerant species such as Acer saccharum, rather than Quercus. We developed a GIS database from stand census measurements taken in 1974 and 2000, employing it to conduct tree‐by‐tree comparisons that allow direct determination of ingrowth, mortality and survivorship, and to relate the spatial patterns of subcanopy dynamics to canopy gap occurrence. Results: The re‐census shows modest changes in canopy composition, but much greater turnover in the subcanopy. Nearly half of all individuals originally present died; much of this mortality resulted from a major decline in subcanopy Ulmus americana. While overall density remained fairly constant, the subcanopy experienced substantial ingrowth of shade‐tolerant Acer saccharum, Fagus grandifolia, and Tilia americana. Canopy gaps, although forming at rates in the upper range of regional averages, did not significantly benefit subcanopy populations of Quercus spp. or most other taxa with limited shade tolerance. Conclusions: Canopy gaps play a minor role in driving the recent demographic trends of this stand. The spatial and temporal scales of light availability in gaps do not support regeneration of most shade‐intolerant species. Compositional change parallels a historical shift in light regimes.