Effects of forest management on ground-dwelling beetles (Coleoptera; Carabidae, Staphylinidae) in Central Europe are mainly mediated by changes in forest structure

Forest management is known to influence species diversity of various taxa but inconsistent or even contrasting effects are reported for arthropods. Regional differences in management as well as differences in regional species pools might be responsible for these inconsistencies, but, inter-regional replicated studies that account for regional variability are rare. We investigated the effect of forest type on the abundance, diversity, community structure and composition of two important ground-dwelling beetle families, Carabidae and Staphylinidae, in 149 forest stands distributed over three regions in Germany. In particular we focused on recent forestry history, stand age and dominant tree species, in addition to a number of environmental descriptors. Overall management effects on beetle communities were small and mainly mediated by structural habitat parameters such as the cover of forest canopy or the plant diversity on forest stands. The general response of both beetle taxa to forest management was similar in all regions: abundance and species richness of beetles was higher in older than in younger stands and species richness was lower in unmanaged than in managed stands. The abundance ratio of forest species-to-open habitat species differed between regions, but generally increased from young to old stands, from coniferous to deciduous stands and from managed to unmanaged stands. The response of both beetle families to dominant tree species was variable among regions and staphylinid richness varied in the response to recent forestry history. Our results suggest that current forest management practices change the composition of ground-dwelling beetle communities mainly by favoring generalists and open habitat species. To protect important forest beetle communities and thus the ecosystem functions and services provided by them, we suggest to shelter remaining ancient forests and to develop near-to-nature management strategies by prolonging rotation periods and increasing structural diversity of managed forests. Possible geographic variations in the response of beetle communities need to be considered in conservation-orientated forest management strategies.

Forest structure and composition of previously selectively logged and non-logged montane forests at Mt. Kilimanjaro

The montane forests of Mount Kilimanjaro in Tanzania have been subjected to a long history of selective logging. However, since 1984 logging of indigenous trees is prohibited. Today, these forests allow us to evaluate the long-term effects of selective logging. We mapped the height and diameter at breast height (DBH) of all trees >10 cm DBH on 10 sites of 0.25 ha. Five sites represent non-logged forests, another five selectively logged forests. We tested whether forests were still visibly affected 30–40 years after selective logging in terms of their forest structure and tree diversity. Additionally we compared tree densities of different species guilds, including disturbance-indicator species, late-successional species and main timber species. Furthermore, we specifically compared the community size distributions of selectively logged and non-logged forests, first across all species and then for the most important timber species, Ocotea usambarensis, alone. 30–40 years after selective logging forests still showed a higher overall stem density, mainly due to higher relative abundances of small trees (<50 cm DBH) in general, and higher densities of small size class stems of late-successional species specifically. For O. usambarensis, the selectively logged sites harboured higher relative abundances of small trees and lower relative abundances of harvestable trees. The higher relative abundance of small O. usambarensis-stems in selectively logged forests appears promising for future forest recovery. Thus, outside protected areas, selective logging may be a sustainable management option if logging cycles are considerably longer than 40 years, enough large source trees remain, and the recruiting O. usambarensis individuals find open space for their establishment.

Improved methods for measuring forest landscape structure: LiDAR complements field-based habitat assessment

Conservation and monitoring of forest biodiversity requires reliable information about forest structure and composition at multiple spatial scales. However, detailed data about forest habitat characteristics across large areas are often incomplete due to difficulties associated with field sampling methods. To overcome this limitation we employed a nationally available light detection and ranging (LiDAR) remote sensing dataset to develop variables describing forest landscape structure across a large environmental gradient in Switzerland. Using a model species indicative of structurally rich mountain forests (hazel grouse Bonasa bonasia), we tested the potential of such variables to predict species occurrence and evaluated the additional benefit of LiDAR data when used in combination with traditional, sample plot-based field variables. We calibrated boosted regression trees (BRT) models for both variable sets separately and in combination, and compared the models’ accuracies. While both field-based and LiDAR models performed well, combining the two data sources improved the accuracy of the species’ habitat model. The variables retained from the two datasets held different types of information: field variables mostly quantified food resources and cover in the field and shrub layer, LiDAR variables characterized heterogeneity of vegetation structure which correlated with field variables describing the understory and ground vegetation. When combined with data on forest vegetation composition from field surveys, LiDAR provides valuable complementary information for encompassing species niches more comprehensively. Thus, LiDAR bridges the gap between precise, locally restricted field-data and coarse digital land cover information by reliably identifying habitat structure and quality across large areas.

Forest dynamics during the transition from the Oldest Dryas to the Bølling–Allerød at Gerzensee—a simulation study

The aim of this study was to explore potential causes and mechanisms for the sequence and temporal pattern of tree taxa, specifically for the shift from shrub-tundra to birch–juniper woodland during and after the transition from the Oldest Dryas to the Bølling–Allerød in the region surrounding the lake Gerzensee in southern Central Europe. We tested the influence of climate, forest dynamics, community dynamics compared to other causes for delays. For this aim temperature reconstructed from a δ18O-record was used as input driving the multi-species forest-landscape model TreeMig. In a stepwise scenario analysis, population dynamics along with pollen production and transport were simulated and compared with pollen-influx data, according to scenarios of different δ18O/temperature sensitivities, different precipitation levels, with/without inter-specific competition, and with/without prescribed arrival of species. In the best-fitting scenarios, the effects on competitive relationships, pollen production, spatial forest structure, albedo, and surface roughness were examined in more detail. The appearance of most taxa in the data could only be explained by the coldest temperature scenario with a sensitivity of 0.3‰/°C, corresponding to an anomaly of − 15 °C. Once the taxa were present, their temporal pattern was shaped by competition. The later arrival of Pinus could not be explained even by the coldest temperatures, and its timing had to be prescribed by first observations in the pollen record. After the arrival into the simulation area, the expansion of Pinus was further influenced by competitors and minor climate oscillations. The rapid change in the simulated species composition went along with a drastic change in forest structure, leaf area, albedo, and surface roughness. Pollen increased only shortly after biomass. Based on our simulations, two alternative potential scenarios for the pollen pattern can be given: either very cold climate suppressed most species in the Oldest Dryas, or they were delayed by soil formation or migration. One taxon, Pinus, was delayed by migration and then additionally hindered by competition. Community dynamics affected the pattern in two ways: potentially by facilitation, i.e. by nitrogen-fixing pioneer species at the onset, whereas the later pattern was clearly shaped by competition. The simulated structural changes illustrate how vegetation on a larger scale could feed back to the climate system. For a better understanding, a more integrated simulation approach covering also the immigration from refugia would be necessary, for this combines climate-driven population dynamics, migration, individual pollen production and transport, soil dynamics, and physiology of individual pollen production.

Simulating Carbon Stocks and Fluxes of an African Tropical Montane Forest with an Individual-Based Forest Model

Tropical forests are carbon-dense and highly productive ecosystems. Consequently, they play an important role in the global carbon cycle. In the present study we used an individual-based forest model (FORMIND) to analyze the carbon balances of a tropical forest. The main processes of this model are tree growth, mortality, regeneration, and competition. Model parameters were calibrated using forest inventory data from a tropical forest at Mt. Kilimanjaro. The simulation results showed that the model successfully reproduces important characteristics of tropical forests (aboveground biomass, stem size distribution and leaf area index). The estimated aboveground biomass (385 t/ha) is comparable to biomass values in the Amazon and other tropical forests in Africa. The simulated forest reveals a gross primary production of 24 tcha-1yr-1. Modeling above- and belowground carbon stocks, we analyzed the carbon balance of the investigated tropical forest. The simulated carbon balance of this old-growth forest is zero on average. This study provides an example of how forest models can be used in combination with forest inventory data to investigate forest structure and local carbon balances.

Primary lowland dipterocarp forest at Danum Valley, Sabah, Malaysia. Species composition and patterns in the understorey

The spatial distributions of species of tree ≥10 cm gbh were examined in two 4 ha plots and related to the local variation in topography and soil chemistry. The plots were similar in their species composition, particularly in terms of the densities of small trees, and they showed very similar edaphic characteristics. Size class distributions varied little within and between plots. Ordination of 0.25 ha subplots highlighted parallel gradients in the vegetation of both plots when the densities of trees ≥10 cm gbh were considered. Focusing on understorey trees in the 10-<50 cm gbh class at the 0.04 ha subplot scale showed a similar vegetation gradient in both plots closely associated with change from lower slope to ridge. No relationship with soil chemistry was found. On the ridges a special group of understorey species formed clumps and these species contributed importantly to the ordinations. Borneo has a regional history of occasionally severe droughts. It is suggested here that the observed patterns in the understorey are due to differential responses to low soil water supply, the ridges probably tending to dryness more than the lower slopes. Within the large and diverse family Euphorbiaceae, which dominates the understorey at Danum, there may be ecophysiological groupings of species. The long-term effects of disturbance interacting with local edaphic factors on forest structure and composition are discussed.

Placing unprecedented recent fir growth in a European-wide and Holocene-long context

Forest decline played a pivotal role in motivating Europe's political focus on sustainability around 35 years ago. Silver fir (Abies alba) exhibited a particularly severe dieback in the mid-1970s, but disentangling biotic from abiotic drivers remained challenging because both spatial and temporal data were lacking. Here, we analyze 14 136 samples from living trees and historical timbers, together with 356 pollen records, to evaluate recent fir growth from a continent-wide and Holocene-long perspective. Land use and climate change influenced forest growth over the past millennium, whereas anthropogenic emissions of acidic sulfates and nitrates became important after about 1850. Pollution control since the 1980s, together with a warmer but not drier climate, has facilitated an unprecedented surge in productivity across Central European fir stands. Restricted fir distribution prior to the Mesolithic and again in the Modern Era, separated by a peak in abundance during the Bronze Age, is indicative of the long-term interplay of changing temperatures, shifts in the hydrological cycle, and human impacts that have shaped forest structure and productivity.

Differential Responses of Herbivores and Herbivory to Management in Temperate European Beech

Forest management not only affects biodiversity but also might alter ecosystem processes mediated by the organisms, i.e. herbivory the removal of plant biomass by plant-eating insects and other arthropod groups. Aiming at revealing general relationships between forest management and herbivory we investigated aboveground arthropod herbivory in 105 plots dominated by European beech in three different regions in Germany in the sun-exposed canopy of mature beech trees and on beech saplings in the understorey. We separately assessed damage by different guilds of herbivores, i.e. chewing, sucking and scraping herbivores, gall-forming insects and mites, and leaf-mining insects. We asked whether herbivory differs among different forest management regimes (unmanaged, uneven-aged managed, even-aged managed) and among age-classes within even-aged forests. We further tested for consistency of relationships between regions, strata and herbivore guilds. On average, almost 80 of beech leaves showed herbivory damage, and about 6 of leaf area was consumed. Chewing damage was most common, whereas leaf sucking and scraping damage were very rare. Damage was generally greater in the canopy than in the understorey, in particular for chewing and scraping damage, and the occurrence of mines. There was little difference in herbivory among differently managed forests and the effects of management on damage differed among regions, strata and damage types. Covariates such as wood volume, tree density and plant diversity weakly influenced herbivory, and effects differed between herbivory types. We conclude that despite of the relatively low number of species attacking beech; arthropod herbivory on beech is generally high. We further conclude that responses of herbivory to forest management are multifaceted and environmental factors such as forest structure variables affecting in particular microclimatic conditions are more likely to explain the variability in herbivory among beech forest plots.

The ecoclimatology of Danum, Sabah, in the context of the world's rainforest regions, with particular reference to dry periods and their impact

Climatic records for Danum for 1985 to 1998, elsewhere in Sabah since 1879, and long monthly rainfall series from other rainforest locations are used to place the climate, and particularly the dry period climatology, of Danum into a world rainforest context. The magnitude frequency and seasonality of dry periods are shown to vary greatly within the world's rainforest zone. The climate of Danum, which is aseasonal but subject, as in 1997 to 1998, to occasional drought, is intermediate between less drought–prone north–western Borneo and the more drought–prone east coast. Changes through time in drought magnitude frequency in Sabah and rainforest locations elsewhere in South–East Asia and in the Neotropics are compared. The 1997 to 1998 ENSO–related drought event in Sabah is placed into a historical context. The effects of drought on tree growth and mortality in the tropics are assessed and a model relating intensity and frequency of drought disturbance to forest structure and composition is discussed.

Structure and inferred dynamics of a large grove of Microberlinia bisulcata trees in central African rain forest: the possible role of periods of multiple disturbance events

A 272-ha grove of dominant Microberlinia bisulcata (Caesalpinioideae) adult trees greater than or equal to 50 cm stem diameter was mapped in its entirety in the southern part of Korup National Park, Cameroon. The approach used an earlier-established 82.5-ha permanent plot with a new surrounding 50-m grid of transect lines. Tree diameters were available from the plot but trees on the grid were recorded as being greater than or equal to 50 cm. The grove consisted of 1028 trees in 2000. Other species occurred within the grove. including the associated subdominants Tetraberlinia bifoliolata and T. korupensis. Microberlinia bisulcata becomes adult at a stein diameter of c. 50 cm and at an estimated age of 50 y. Three oval-shaped subgroves with dimensions c. 8 50 in x 13 50 in (90 ha) were defined. For two of them (within the plot) tree diameters were available. Subgroves differed in their scales and intensities of spatial tree patterns, and in their size frequency distributions, these suggesting differing past dynamics. The modal scale of clumping was 40-50 m. Seed dispersal by pod ejection (to c. 50 in) was evident from the semi-circles of trees at the grove's edge and from the many internal circles (100-200 m diameter). The grove has the capacity. therefore, to increase at c. 100 m per century. To form its present extent and structure. it is inferred that it expanded and infilled from a possibly smaller area of lower adult-tree density. This possibly happened in three waves of recruitment, each one determined by a period of several intense disturbances. Climate records for Africa show that 1740-50 and 1820-30 were periods of drought, and that 1870-1895 was also regionally very dry. Canopy openings allow the light-demanding and fast-growing ectomycorrhizal M. bisulcata to establish, but successive releases are thought to be required to achieve effective recruitment. Nevertheless, in the last 50 y there were no major events and recruitment in the grove was very poor. This present study leads to a new hypothesis of the role of periods of multiple extreme events being the driving factor for the population dynamics of many large African tree species such as M. bisulcata.

Buttress form of the central African rain forest tree Microberlinia bisulcata, and its possible role in nutrient acquisition

Buttressing is a trait special to tropical trees but explanations for its occurrence remain inconclusive. The two main hypotheses are that they provide structural support and/or promote nutrient acquisition. Studies of the first are common but the second has received much less attention. Architectural measurements were made on adult and juvenile trees of the ectomycorrhizal species Microberlinia bisulcata, in Korup (Cameroon). Buttressing on this species is highly distinctive with strong lateral extension of surface roots of the juveniles leading to a mature buttress system of a shallow spreading form on adults. This contrasts with more vertical buttresses, closer to the stem, found on many other tropical tree species. No clear relationship between main buttress and large branch distribution was found. Whilst this does not argue against the essential structural role of buttresses for these very large tropical trees, the form on M. bisulcata does suggest a likely second role, that of aiding nutrient acquisition. At the Korup site, with its deep sandy soils of very low phosphorus status, and where most nutrient cycling takes place in a thin surface layer of fine roots and mycorrhizas, it appears that buttress form could develop from soil-surface root exploration for nutrients by juvenile trees. It may accordingly allow M. bisulcata to attain the higher greater competitive ability, faster growth rate, and maximum tree size that it does compared with other co-occurring tree species. For sites across the tropics in general, the degree of shallowness and spatial extension of buttresses of the dominant species is hypothesized to increase with decreasing nutrient availability.

Transient dominance in a central African rain forest

The large-crowned emergent tree Microberlinia bisulcata dominates rain forest groves at Korup National Park, Cameroon, along with two codominants, Tetraberlinia bifoliolata and T. korupensis. M. bisulcata has a pronounced modal size frequency distribution around 110 cm stem diameter: its recruitment potential is very poor. It is a long-lived light-demanding species, one of many found in African forests. Tetraberlinia species lack modality, are more shade tolerant, and recruit better. All three species are ectomycorrhizal. M. bisulcata dominates grove basal area, even though it has similar numbers of trees (≥50 cm stem diameter) as each of the other two species. This situation presented a conundrum that prompted a long-term study of grove dynamics. Enumerations of two plots (82.5 and 56.25 ha) between 1990 and 2010 showed mortality and recruitment of M. bisulcata to be very low (both rates 0.2% per year) compared with Tetraberlinia (2.4% and 0.8% per year), and M. bisulcata grows twice as fast as the Tetraberlinia. Ordinations indicated that these three species determined community structure by their strong negative associations while other species showed almost none. Ranked species abundance curves fitted the Zipf-Mandelbrot model well and allowed “overdominance” of M. bisulcata to be estimated. Spatial analysis indicated strong repulsion by clusters of large (50 to <100 cm) and very large (≥100 cm) M. bisulcata of their own medium-sized (10 to <50 cm) trees and all sizes of Tetraberlinia. This was interpreted as competition by M. bisulcata increasing its dominance, but also inhibition of its own replacement potential. Stem coring showed a modal age of 200 years for M. bisulcata, but with large size variation (50–150 cm). Fifty-year model projections suggested little change in medium, decreases in large, and increases in very large trees of M. bisulcata, accompanied by overall decreases in medium and large trees of Tetraberlinia species. Realistically increasing very-large-tree mortality led to grove collapse without short-term replacement. M. bisulcata most likely depends on climatic events to rebuild its stands: the ratio of disturbance interval to median species' longevity is important. A new theory of transient dominance explains how M. bisulcata may be cycling in abundance over time and displaying nonequilibrium dynamics.

Secondary succession and dipterocarp recruitment in Bornean rain forest after logging

To understand succession in dipterocarp rain forest after logging, the structure, species composition and dynamics of primary (PF) and secondary (SF) forest at Danum were compared. In 10 replicate 0.16-ha plots per forest type trees >= 10 cm gbh (3.2 cm dbh) were measured in 1995 and 2001. The SF had been logged in 1988, which allowed successional change to be recorded at 8 and 13 years. In 2001, saplings (1.0-3.1 cm dbh) were measured in nested quadrats. The forest types were similar in mean radiation at 2 m height, and in density, basal area and species number of all trees. Among small (10 <= 31.4) and large ( >= 31.4 cm gbh) trees, in both 1995 and 2001, there were 10- and 3-fold more dipterocarps in SF than PF respectively; and averaging over the two dates, there were correspondingly ca. 10- and 18-fold more pioneers. Mortality was ca. 60% higher in SF than PF, largely due to a seven-fold difference for pioneers: for dipterocarps there was little difference. Recruitment was similar in PF and SE Stem growth rates were 37% higher in SF than PF for all trees, although dipterocarps showed the opposite trend. Among saplings, dipterocarps dominated SF with a 10-fold higher density than in PF. For dipterocarps, the light (LH) and medium-heavy (MHH) canopy hardwoods, and the shade-tolerant, smaller-stature other (OTH) species (e.g. Hopea and Vatica) were in the ratios ca. 40:15:45 in SF and 85: < 1:15 in PF. LHs had higher mortality than OTHs in SE In PF ca. 80% of the saplings were LH: in SF ca. 70% were OTH. The predominance of OTHs in SF is explained by the logging of primary rain forest which was in a likely late stage of recovery from natural disturbance, plus the continuing shaded conditions in the understorey promoted by dense pioneer vegetation. At 13 years after logging succession appeared to be inhibited: LHs were being suppressed but MHHs and OTHs persisted. Succession in lowland dipterocarp, rain forests may therefore depend on the successional state of the primary forest when it is logged. A review of logged versus unlogged studies in Borneo highlights the need for more detailed ecological comparisons.

Temperate Mountain Forest Biodiversity under Climate Change: Compensating Negative Effects by Increasing Structural Complexity

Species adapted to cold-climatic mountain environments are expected to face a high risk of range contractions, if not local extinctions under climate change. Yet, the populations of many endothermic species may not be primarily affected by physiological constraints, but indirectly by climate-induced changes of habitat characteristics. In mountain forests, where vertebrate species largely depend on vegetation composition and structure, deteriorating habitat suitability may thus be mitigated or even compensated by habitat management aiming at compositional and structural enhancement. We tested this possibility using four cold-adapted bird species with complementary habitat requirements as model organisms. Based on species data and environmental information collected in 300 1-km2 grid cells distributed across four mountain ranges in central Europe, we investigated (1) how species’ occurrence is explained by climate, landscape, and vegetation, (2) to what extent climate change and climate-induced vegetation changes will affect habitat suitability, and (3) whether these changes could be compensated by adaptive habitat management. Species presence was modelled as a function of climate, landscape and vegetation variables under current climate; moreover, vegetation-climate relationships were assessed. The models were extrapolated to the climatic conditions of 2050, assuming the moderate IPCC-scenario A1B, and changes in species’ occurrence probability were quantified. Finally, we assessed the maximum increase in occurrence probability that could be achieved by modifying one or multiple vegetation variables under altered climate conditions. Climate variables contributed significantly to explaining species occurrence, and expected climatic changes, as well as climate-induced vegetation trends, decreased the occurrence probability of all four species, particularly at the low-altitudinal margins of their distribution. These effects could be partly compensated by modifying single vegetation factors, but full compensation would only be achieved if several factors were changed in concert. The results illustrate the possibilities and limitations of adaptive species conservation management under climate change.