The role of tree size in the leafing phenology of a seasonally dry tropical forest in Belize, Central America

Leafing phenology of two dry-forest sites on soils of different depth (S = shallow, D = deep) at Shipstern Reserve, Belize, were compared at the start of the rainy season (April-June 2000). Trees greater than or equal to 2.5 cm dbh were recorded weekly for 8 wk in three 0.04-ha plots per site. Ten species were analysed individually for their phenological patterns, of which the three most common were Bursera simaruba, Metopium brownei and Jatropha gaumeri. Trees were divided into those in the canopy (> 10 cm dbh) and the subcanopy (less than or equal to 10 cm dbh). Site S had larger trees on average than site D. The proportion of trees flushing leaves at any one time was generally higher in site S than in site D, for both canopy and subcanopy trees. Leaf flush started 2 wk earlier in site S than site D for subcanopy trees, but only 0.5 wk earlier for the canopy trees. Leaf flush duration was 1.5 wk longer in site S than site D. Large trees in the subcanopy flushed leaves earlier than small ones at both sites but in the canopy just at site D. Large trees flushed leaves earlier than small ones in three species and small trees flushed leaves more rapidly in two species. Bursera and Jatropha followed the general trends but Metopium, with larger trees in site D than site S, showed the converse with onset of flushing I wk earlier in site D than site S. Differences in response of the canopy and subcanopy trees on each site can be accounted for by the predominance of spring-flushing or stem-succulent species in site S and a tendency for evergreen species to occur in site D. Early flushing of relatively larger trees in site D most likely requires access to deeper soil water reserves but small and large trees utilize stored tree water in site S.

Tree size and fecundity influence ballistic seed dispersal of two dominant mast-fruiting species in a tropical rain forest

Seed production, seed dispersal, and seedling recruitment are integral to forest dynamics, especially in masting species. Often these are studied separately, yet scarcely ever for species with ballistic dispersal even though this mode of dispersal is common in legume trees of tropical African rain forests. Here, we studied two dominant main-canopy tree species, Microberlinia bisulcata and Tetraberlinia bifoliolata (Caesalpinioideae), in 25 ha of primary rain forest at Korup, Cameroon, during two successive masting events (2007/2010). In the vicinity of c. 100 and 130 trees of each species, 476/580 traps caught dispersed seeds and beneath their crowns c. 57,000 pod valves per species were inspected to estimate tree-level fecundity. Seed production of trees increased non-linearly and asymptotically with increasing stem diameters. It was unequal within the two species’ populations, and differed strongly between years to foster both spatial and temporal patchiness in seed rain. The M. bisulcata trees could begin seeding at 42–44 cm diameter: at a much larger size than could T. bifoliolata (25 cm). Nevertheless, per capita life-time reproductive capacity was c. five times greater in M. bisulcata than T. bifoliolata owing to former’s larger adult stature, lower mortality rate (despite a shorter life-time) and smaller seed mass. The two species displayed strong differences in their dispersal capabilities. Inverse modelling (IM) revealed that dispersal of M. bisulcata was best described by a lognormal kernel. Most seeds landed at 10–15 m from stems, with 1% of them going beyond 80 m (<100 m). The direct estimates of fecundity significantly improved the models fitted. The lognormal also described well the seedling recruitment distribution of this species in 121 ground plots. By contrast, the lower intensity of masting and more limited dispersal of the heavier-seeded T. bifoliolata prevented reliable IM. For this species, seed density as function of distance to traps suggested a maximum dispersal distance of 40–50 m, and a correspondingly more aggregated seedling recruitment pattern ensued than for M. bisulcata. From this integrated field study, we conclude that the reproductive traits of M. bisulcata give it a considerable advantage over T. bifoliolata by better dispersing more seeds per capita to reach more suitable establishment sites, and combined with other key traits they explain its local dominance in the forest. Understanding the linkages between size at onset of maturity, individual fecundity, and dispersal capability can better inform the life-history strategies, and hence management, of co-occurring tree species in tropical forests.

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.

A novel comparative research platform designed to determine the functional significance of tree species diversity in European forests

One of the current advances in functional biodiversity research is the move away from short-lived test systems towards the exploration of diversity-ecosystem functioning relationships in structurally more complex ecosystems. In forests, assumptions about the functional significance of tree species diversity have only recently produced a new generation of research on ecosystem processes and services. Novel experimental designs have now replaced traditional forestry trials, but these comparatively young experimental plots suffer from specific difficulties that are mainly related to the tree size and longevity. Tree species diversity experiments therefore need to be complemented with comparative observational studies in existing forests. Here we present the design and implementation of a new network of forest plots along tree species diversity gradients in six major European forest types: the FunDivEUROPE Exploratory Platform. Based on a review of the deficiencies of existing observational approaches and of unresolved research questions and hypotheses, we discuss the fundamental criteria that shaped the design of our platform. Key features include the extent of the species diversity gradient with mixtures up to five species, strict avoidance of a dilution gradient, special attention to community evenness and minimal covariation with other environmental factors. The new European research platform permits the most comprehensive assessment of tree species diversity effects on forest ecosystem functioning to date since it offers a common set of research plots to groups of researchers from very different disciplines and uses the same methodological approach in contrasting forest types along an extensive environmental gradient. (C) 2013 Elsevier GmbH. All rights reserved.

Neighbourhood abundance and small-tree survival in a lowland Bornean rainforest

Conspecific effects of neighbours on small-tree survival may have a role in tree population dynamics and community composition of tropical forests. This notion was tested with data from two 4-ha plots in lowland forest at Danum, Sabah (Borneo), for a 21-year interval (censuses at 1986, 1996, 2001, 2007). Species with ≥45 focal trees 10 to <100 cm stem girth per plot in 1986 were selected. Logistic regressions fitted mean focal tree size and mean inverse-distance-weighted basal area abundance of neighbours (within 20 m), for the periods over which each focus tree was alive. Coefficients of variation of neighbourhood basal area abundance, both spatially and temporally, quantified the changing environment of each focus tree. Fits were critically and individually evaluated, with corrections for spatial autocorrelation. Conspecific effects at Danum was generally very weak or non-existent: species’ mortality rates varied also across plots. The main reasons appear to be that (1) species were not dense enough to interact despite frequent although weak spatial aggregation, and their neighbourhoods were highly differing in species composition; and (2) these neighbourhoods were highly variable temporally, meaning that focus trees experienced stochastically fluctuating neighbourhood environments. Only one species, Dimorphocalyx muricatus, showed strong conspecific effects (varying between plots) which can be explained by its distinct ecology. This understorey species is highly aggregated on ridges and is drought-tolerant. That this functionally and habitat-specialized species, has implied intraspecific density-dependent feedback in its dynamics is a remarkable indication of the overall processes maintaining stability of the Danum forest.

On the detection of dynamic responses in a drought-perturbed tropical rainforest in Borneo

The dynamics of aseasonal lowland dipterocarp forest in Borneo is influenced by perturbation from droughts. These events might be increasing in frequency and intensity in the future. This paper describes drought-affected dynamics between 1986 and 2001 in Sabah, Malaysia, and considers how it is possible, reliably and accurately, to measure both coarse- and fine-scale responses of the forest. Some fundamental concerns about methodology and data analysis emerge. In two plots forming 8 ha, mortality, recruitment, and stem growth rates of trees ≥10 cm gbh (girth at breast height) were measured in a ‘pre-drought’ period (1986–1996), and in a period (1996–2001) including the 1997–1998 ENSO-drought. For 2.56 ha of subplots, mortality and growth rates of small trees (10–<50 cm gbh) were found also for two sub-periods (1996–1999, 1999–2001). A total of c. 19 K trees were recorded. Mortality rate increased by 25% while both recruitment and relative growth rates increased by 12% for all trees at the coarse scale. For small trees, at the fine scale, mortality increased by 6% and 9% from pre-drought to drought and on to ‘post-drought’ sub-periods. Relative growth rates correspondingly decreased by 38% and increased by 98%. Tree size and topography interacted in a complex manner with between-plot differences. The forest appears to have been sustained by off-setting elevated tree mortality by highly resilient stem growth. This last is seen as the key integrating tree variable which links the external driver (drought causing water stress) and population dynamics recorded as mortality and recruitment. Suitably sound measurements of stem girth, leading to valid growth rates, are needed to understand and model tree dynamic responses to perturbations. The proportion of sound data, however, is in part determined by the drought itself.

Experimental evidence for trait utility of gill raker number in adaptive radiation of a north temperate fish

North temperate fish in post-glacial lakes are textbook examples for rapid parallel adaptive radiation into multiple trophic specialists within individual lakes. Speciation repeatedly proceeded along the benthic – limnetic habitat axis, and benthic – limnetic sister species diverge in the number of gill rakers. Yet, the utility of different numbers of gill rakers for consuming benthic vs. limnetic food has only very rarely been experimentally demonstrated. We bred and raised families of a benthic – limnetic species pair of whitefish under common garden conditions to test whether these species (i) show heritable differentiation in feeding efficiency on zooplankton, and (ii) whether varia- tion in feeding efficiency is predicted by variation in gill raker numbers. We used zooplankton of three different size classes to investigate prey size dependency of divergence in feeding efficiency and to investigate the effect strength of variation in the number of gill rakers. Our results show strong interspecific differences in feeding efficiency. These differences are largest when fish were tested with the smallest zooplankton. Importantly, feeding efficiency is significantly positively correlated with the number of gill rakers when using small zooplankton, also when species identity is statistically controlled for. Our results support the hypothesis that a larger number of gill rakers are of adaptive significance for feeding on zooplankton and pro- vide one of the first experimental demonstrations of trait utility of gill raker number when fish feed on zooplankton. These results are consistent with the suggested importance of divergent selection driven feeding adaptation during adaptive radiation of fish in post-glacial lakes.

Contrasting vulnerability and resilience to drought-induced decline of densely planted vs. natural rear-edge Pinus nigra forests

The southernmost European natural and planted pine forests are among the most vulnerable areas to warming-induced drought decline. Both drought stress and management factors (e.g., stand origin or reduced thinning) may induce decline by reducing the water available to trees but their relative importances have not been properly assessed. The role of stand origin - densely planted vs. naturally regenerated stands - as a decline driver can be assessed by comparing the growth and vigor responses to drought of similar natural vs. planted stands. Here, we compare these responses in natural and planted Black pine (Pinus nigra) stands located in southern Spain. We analyze how environmental factors - climatic (temperature and precipitation anomalies) and site conditions - and biotic factors - stand structure (age, tree size, density) and defoliation by the pine processionary moth - drive radial growth and crown condition at stand and tree levels. We also assess the climatic trends in the study area over the last 60 years. We use dendrochronology, linear mixed-effects models of basal area increment and structural equation models to determine how natural and planted stands respond to drought and current competition intensity. We observed that a temperature rise and a decrease in precipitation during the growing period led to increasing drought stress during the late 20th century. Trees from planted stands experienced stronger growth reductions and displayed more severe crown defoliation after severe droughts than those from natural stands. High stand density negatively drove growth and enhanced crown dieback, particularly in planted stands. Also pine processionary moth defoliation was more severe in the growth of natural than in planted stands but affected tree crown condition similarly in both stand types. In response to drought, sharp growth reduction and widespread defoliation of planted Mediterranean pine stands indicate that they are more vulnerable and less resilient to drought stress than natural stands. To mitigate forest decline of planted stands in xeric areas such as the Mediterranean Basin, less dense and more diverse stands should be created through selective thinning or by selecting species or provenances that are more drought tolerant. (C) 2013 Elsevier B.V. All rights reserved.

Does proximity to conspecific adults influence the establishment of ectomycorrhizal trees in rain forest?

Three ectomycorrhizal legume trees, Microberlinia bisulcata, Tetraberlinia bifoliolata and T. moreliana, form discrete groves in the southern part of Korup National Park, in southwest Cameroon and contribute c. 45–70% of stand basal area locally in a matrix of otherwise species-rich arbuscular mycorrhizal forest. A transplant experiment was performed to assess the importance of ectomycorrhizal infection associated with proximity to parents in seedling establishment of the grove-forming species. Nonectomycorrhizal seedlings of the three species were transplanted into plots of two forest types, one of high (HEM, within-grove) and one of very low (LEM, outside the grove) abundance of all three species as adult trees. For two species (T. moreliana and M. bisulcata) there was no difference in survival over 16 months, but for the third (T. bifoliolata) survival was best in HEM forest, and correlated with the basal area of adult trees of ectomycorrhizal species. Only one species (T. moreliana) increased in biomass over the experimental period; the others declined. There was no effect of forest type on overall growth of any species, but the survivors of two (T. moreliana and M. bisulcata) had heavier stems in the HEM forest. Differences in survival and growth of transplants between the three species were in accord with the ecology of the species as inferred from the frequency distributions of adult tree size in the forest. Seedlings became infected with ectomycorrhizas in both forest types; where there was a difference in extent of infection (T. moreliana) this was not related to survival or growth; and where there was a difference in survival (T. bifoliolata) this was not related to extent of infection. These results confirm that mycorrhizal inoculum associated with conspecific adults is neither a prerequisite nor a guarantee of seedling establishment, but indicates that in some circumstances there might be benefits of being close to parents. Further research is required to unravel the complexities of ectomycorrhizal community structure in this spatially and temporally heterogeneous forest, and to clarify the extent to which the various hosts share ectomycorrhizal partners.

Different-sized grazers have distinctive effects on plant functional composition of an African savannah

Grazing ungulates play a key role in many ecosystems worldwide and can form diverse assemblages, such as in African savannahs. In many of these ecosystems, present-day ungulate communities are impoverished subsets of once diverse assemblages. While we know that excluding all ungulates from grasslands can exert major effects on both the structure and composition of the vegetation, how different individual ungulate species may have contrasting effects on grassland communities remains poorly understood. Here, we performed a long-term ‘Russian doll’ grazing exclosure experiment in an African savannah to test for the effects of different size classes of grazers on grassland structure and composition. At five sites, grazer species of decreasing size class (ranging from white rhino to scrub hare) were excluded using four fence types, to experimentally create different realized grazer assemblages. The vegetation structure and the grass functional community composition were characterized in 6 different years over a 10-year period. Additionally, animal footprints were counted to quantify the abundance of different ungulate species in each treatment. We found that while vegetation height was mostly driven by total grazing pressure of all species together, ungulate community composition best explained the functional community composition of grasses. In the short term, smaller ungulate species (‘mesoherbivores’) had strongest effects on vegetation composition, by shifting communities towards dominance by species with low specific leaf area and low nutritional value. In the long term, large grazers had stronger but similar effects on the functional composition of the system. Surprisingly, the largest ‘mega-herbivore’, the white rhinoceros, did not have strong effects on the vegetation structure or composition. Synthesis. Our results support the idea that different size classes of grazers have varying effects on the functional composition of grassland plant communities. Therefore, the worldwide decline in the diversity of ungulate species is expected to have (had) major impacts on community composition and functioning of grassland ecosystems, even if total grazing pressure has remained constant, for example, due to replacement by livestock.

Light and seed size affect establishment of grove-forming ectomycorrhizal rain forest tree species

• Regeneration of the dominant ectomycorrhizal tree Microberlinia bisulcata in groves in Korup, Central Africa, is very poor. The hypothesis was tested that this species is more shade intolerant than other co-occurring species. • In two 1-yr trials, each with M. bisulcata and four other species at a nursery close to Korup, growth was measured under five PAR levels, with ± added P and ± watering in the dry season. In parallel experiments the effects of PAR with two R : FR ratios were investigated. • Increasing PAR had a consistent effect on the rates of increase in plant mass and on changes in the other variables. Doubling soil P, watering and halving the R : FR ratio had almost no effect. However, across species, mass at low PAR and relative growth rate related positively and negatively, respectively, to seed mass. • One contributing factor for the poor recruitment of M. bisulcata is therefore its low survival and slow growth at low PAR, due to its small seed size. The two codominant ectomycorrhizal grove species of Tetraberlinia, with larger seeds, were less affected by low PAR.

Using scenario tree modelling for targeted herd sampling to substantiate freedom from disease

BACKGROUND: In order to optimise the cost-effectiveness of active surveillance to substantiate freedom from disease, a new approach using targeted sampling of farms was developed and applied on the example of infectious bovine rhinotracheitis (IBR) and enzootic bovine leucosis (EBL) in Switzerland. Relevant risk factors (RF) for the introduction of IBR and EBL into Swiss cattle farms were identified and their relative risks defined based on literature review and expert opinions. A quantitative model based on the scenario tree method was subsequently used to calculate the required sample size of a targeted sampling approach (TS) for a given sensitivity. We compared the sample size with that of a stratified random sample (sRS) with regard to efficiency. RESULTS: The required sample sizes to substantiate disease freedom were 1,241 farms for IBR and 1,750 farms for EBL to detect 0.2% herd prevalence with 99% sensitivity. Using conventional sRS, the required sample sizes were 2,259 farms for IBR and 2,243 for EBL. Considering the additional administrative expenses required for the planning of TS, the risk-based approach was still more cost-effective than a sRS (40% reduction on the full survey costs for IBR and 8% for EBL) due to the considerable reduction in sample size. CONCLUSIONS: As the model depends on RF selected through literature review and was parameterised with values estimated by experts, it is subject to some degree of uncertainty. Nevertheless, this approach provides the veterinary authorities with a promising tool for future cost-effective sampling designs.

Climatic drivers of hourly to yearly tree radius variations along a 6°C natural warming gradient

Climate affects the timing, rate and dynamics of tree growth, over time scales ranging from seconds to centuries. Monitoring how a tree's stem radius varies over these time scales can provide insight into intra-annual stem dynamics and improve our understanding of climate impacts on tree physiology and growth processes. Here, we quantify the response of radial conifer stem size to environmental fluctuations via a novel assessment of tree circadian cycles. We analyze four years of sub-hourly data collected from 56 larch and spruce trees growing along a natural temperature gradient of ∼6 °C in the central Swiss Alps. During the growing season, tree stem diameters were greatest at mid-morning and smallest in the late evening, reflecting the daily cycle of water uptake and loss. Along the gradient, amplitudes calculated from the stem radius cycle were ∼50% smaller at the upper site (∼2200 m a.s.l.) relative to the lower site (∼800 m a.s.l.). We show changes in precipitation, temperature and cloud cover have a substantial effect on typical growing season diurnal cycles; amplitudes were nine times smaller on rainy days (>10 mm), and daily amplitudes are approximately 40% larger when the mean daily temperature is 15–20 °C than when it is 5–10 °C. We find that over the growing season in the sub-alpine forests, spruce show greater daily stem water movement than larch. However, under projected future warming, larch could experience up to 50% greater stem water use, which may severely affect future growth on already dry sites. Our data further indicate that because of the confounding influences of radial growth and short-term water dynamics on stem size, conventional methodology probably overstates the effect of water-linked meteorological variables (i.e. precipitation and relative humidity) on intra-annual tree growth. We suggest future studies use intra-seasonal measurements of cell development and consider whether climatic factors produce reversible changes in stem diameter. These study design elements may help researchers more accurately quantify and attribute changes in forest productivity in response to future warming.

Density-dependent dynamics of a dominant rain forest tree change with juvenile stage and time of masting

Although negative density dependence (NDD) can facilitate tree species coexistence in forests, the underlying mechanisms can differ, and rarely are the dynamics of seedlings and saplings studied together. Herein we present and discuss a novel mechanism based on our investigation of NDD predictions for the large, grove-forming ectomycorrhizal mast fruiting tree, Microberlinia bisulcata (Caesalpiniaceae), in an 82.5-ha plot at Korup, Cameroon. We tested whether juvenile density, size, growth and survival decreases with increasing conspecific adult basal area for 3245 ‘new’ seedlings and 540 ‘old’ seedlings (< 75-cm tall) during an approximately 4-year study period (2008–2012) and for 234 ‘saplings’ (≥ 75-cm tall) during an approximately 6-year study period (2008–2014). We found that the respective densities of new seedlings, old seedlings and saplings were positively, not and negatively related to increasing BA. Maximum leaf numbers and heights of old seedlings were negatively correlated with increasing basal areas, as were sapling heights and stem diameters. Whereas survivorship of new seedlings decreased by more than one-half with increasing basal area over its range in 2010–2012, that of old seedlings decreased by almost two-thirds, but only in 2008–2010, and was generally unrelated to conspecific seedling density. In 2010–2012 relative growth rates in new seedlings’ heights decreased with increasing basal area, as well as with increasing seedling density, together with increasing leaf numbers, whereas old seedlings’ growth was unrelated to either conspecific density or basal area. Saplings of below-average height had reduced survivorship with increasing basal area (probability decreasing from approx. 0.4 to 0.05 over the basal area range tested), but only sapling growth in terms of leaf numbers decreased with increasing basal area. These static and dynamic results indicate that NDD is operating within this system, possibly stabilizing the M. bisulcata population. However, these NDD patterns are unlikely to be caused by symmetric competition or by consumers. Instead, an alternative mechanism for conspecific adult–juvenile negative feedback is proposed, one which involves the interaction between tree phenology and ectomycorrhizal linkages.