Do fungal pathogens drive density-dependent mortality in established seedlings of two dominant African rain forest trees?

Where one or a few tree species reach local high abundance, different ecological factors may variously facilitate or hinder their regeneration. Plant pathogens are thought to be one of those possible agents which drive intraspecific density-dependent mortality of tree seedlings in tropical forests. Experimental evidence for this is scarce, however. In an African rain forest at Korup, we manipulated the density of recently established seedlings (~5–8 wk old; low vs. high-density) of two dominant species of contrasting recruitment potential, and altered their exposure to pathogens using a broad-spectrum fungicide. Seedling mortality of the abundantly recruiting subcanopy tree Oubanguia alata was strongly density-dependent after 7 mo, yet fungicide-treated seedlings had slightly higher mortality than controls. By contrast, seedling mortality of the poorly recruiting large canopy-emergent tree Microberlinia bisulcata was unaffected by density or fungicide. Ectomycorrhizal colonization of M. bisulcata was not affected by density or fungicide either. For O. alata, adverse effects of fungicide on its vesicular arbuscular mycorrhizas may have offset any possible benefit of pathogen removal. We tentatively conclude that fungal pathogens are not a likely major cause of density dependence in O. alata, or of early post-establishment mortality in M. bisulcata. They do not explain the latter's currently very low recruitment rate at Korup.

Seed fate and seedling dynamics after masting in two African rain forest trees

How the effects of biotic factors are moderated by abiotic factors, and their consequences for species interactions, is generally understudied in ecology. A key abiotic feature of forests is regular canopy disturbances that create temporary patches, or “gaps,” of above-average light availability. Co-occurring in lowland primary forest of Korup National Park (Cameroon), Microberlinia bisulcata and Tetraberlinia bifoliolata are locally dominant, ectomycorrhizal trees whose seeds share predator guilds in masting years. Here, we experimentally tested the impact of small mammal predators upon seedling abundance, growth, and survivorship. In 2007, we added a fixed density of seeds of each species to exclosures at 48 gap–understory locations across 82.5 ha within a large Microberlinia grove, and at 15 locations outside it. For both species, small mammals removed more seeds in gaps than in understory, whereas this was reversed for seeds killed by invertebrates. Nonetheless, Microberlinia lost twice as many seeds to small mammals, and more to invertebrates in exclosures, than Tetraberlinia, which was more prone to a pathogenic white fungus. After six weeks, both species had greater seedling establishment in gaps than understory, and in exclosures outside compared to exclosures inside the grove. In the subsequent two-year period, seedling growth and survivorship peaked in exclosures in gaps, but Microberlinia had more seedlings' stems clipped by animals than Tetraberlinia, and more than twice the percentage of leaf area damaged. Whereas Microberlinia seedling performance in gaps was inferior to Tetraberlinia inside the grove, outside it Microberlinia had reduced leaf damage, grew taller, and had many more leaves than Tetraberlinia. No evidence was found for “apparent mutualism” in the understory as seedling establishment of both species increased away from (>25 m) large stems of either species, pointing to “apparent competition” instead. In gaps, Microberlinia seedling establishment was lower near Tetraberlinia than conspecific adults because of context-dependent small mammal satiation. Stage-matrix analysis suggested that protecting Microberlinia from small mammals could increase its population growth rate by 0.06. In the light of prior research we conclude that small mammals and canopy gaps play an important role in promoting species coexistence in this forest, and that their strong interaction contributes to Microberlinia's currently very poor regeneration.

Mast fruiting of large ectomycorrhizal African rain forest trees: importance of dry season intensity, and the resource limitation hypothesis

Mast fruiting is a distinctive reproductive trait in trees. This rain forest study, at a nutrient-poor site with a seasonal climate in tropical Africa, provides new insights into the causes of this mode of phenological patterning. •  At Korup, Cameroon, 150 trees of the large, ectomycorrhizal caesalp, Microberlinia bisulcata, were recorded almost monthly for leafing, flowering and fruiting during 1995–2000. The series was extended to 1988–2004 with less detailed data. Individual transitions in phenology were analysed. •  Masting occurred when the dry season before fruiting was drier, and the one before that was wetter, than average. Intervals between events were usually 2 or 3 yr. Masting was associated with early leaf exchange, followed by mass flowering, and was highly synchronous in the population. Trees at higher elevation showed more fruiting. Output declined between 1995 and 2000. •  Mast fruiting in M. bisulcata appears to be driven by climate variation and is regulated by internal tree processes. The resource-limitation hypothesis was supported. An ‘alternative bearing’ system seems to underlie masting. That ectomycorrhizal habit facilitates masting in trees is strongly implied.

Herbivores differentially limit the seedling growth and sapling recruitment of two dominant rain forest trees

Resource heterogeneity may influence how plants are attacked and respond to consumers in multiple ways. Perhaps a better understanding of how this interaction might limit sapling recruitment in tree populations may be achieved by examining species’ functional responses to herbivores on a continuum of resource availability. Here, we experimentally reduced herbivore pressure on newly established seedlings of two dominant masting trees in 40 canopy gaps, across c. 80 ha of tropical rain forest in central Africa (Korup, Cameroon). Mesh cages were built to protect individual seedlings, and their leaf production and changes in height were followed for 22 months. With more light, herbivores increasingly prevented the less shade-tolerant Microberlinia bisulcata from growing as tall as it could and producing more leaves, indicating an undercompensation. The more shade-tolerant Tetraberlinia bifoliolata was much less affected by herbivores, showing instead near to full compensation for leaf numbers, and a negligible to weak impact of herbivores on its height growth. A stage-matrix model that compared control and caged populations lent evidence for a stronger impact of herbivores on the long-term population dynamics of M. bisulcata than T. bifoliolata. Our results suggest that insect herbivores can contribute to the local coexistence of two abundant tree species at Korup by disproportionately suppressing sapling recruitment of the faster-growing dominant via undercompensation across the light gradient created by canopy disturbances. The functional patterns we have documented here are consistent with current theory, and, because gap formations are integral to forest regeneration, they may be more widely applicable in other tropical forest communities. If so, the interaction between life-history and herbivore impact across light gradients may play a substantial role in tree species coexistence.

Distribution of cutin and suberin biomarkers under forest trees with different root systems

Background and aims Differences in chemical composition of root compounds and root systems among tree species may affect organic matter (OM) distribution, source and composition in forest soils. The objective of this study was to elucidate the contribution of species specific cutin and suberin biomarkers as proxies for shoot- and root-derived organic carbon (OC) to soil OM at different depths with increasing distance to the stems of four different tree species. Methods The contribution of cutin- and suberin-derived lipids to OM in a Cutanic Alisol was analyzed with increasing soil depth and distance to the stems of Fagus sylvatica L., Picea abies (L.) Karst., Quercus robur L. and Pseudotsuga menziesii (Mirb.) Franco. Cutin and suberin monomers of plants and soils were analyzed by alkaline hydrolysis and subsequent gas chromatography–mass spectrometry. Results The amount and distribution of suberin-derived lipids in soil clearly reflected the specific root system of the different tree species. The amount of cutin-derived lipids decreased strongly with soil depth, indicating that the input of leaf/needle material is restricted to the topsoil. In contrast to the suberin-derived lipids, the spatial pattern of cutin monomer contribution to soil OM did not depend on tree species. Conclusions Our results document the importance of tree species as a main factor controlling the composition and distribution of OM in forest soils. They reveal the impact of tree species on root-derived OM distribution and the necessity to distinguish among different zones when studying soil OM storage in forests.

Tree architecture in a Bornean lowland rain forest: intraspecific and interspecific patterns

Intraspecific and interspecific architectural patterns were studied for eight tree species of a Bornean rain forest. Trees 5--19 m tall in two 4-ha permanent sample plots in primary forest were selected, and three light descriptors and seven architectural traits for each tree were measured. Two general predictions were made: (1) Slow growing individuals (or short ones) encounter lower light, and have flatter crowns, fewer leaf layers, and thinner stems, than do fast growing individuals (or tall ones). (2) Species with higher shade-tolerance receive less light and have flatter crowns, fewer leaf layers, and thinner stems, than do species with lower shade-tolerance. Shade-tolerance is assumed to decrease with maximum growth rate, mortality rate, and adult stature of a species.

Island Invasion by a Threatened Tree Species: Evidence for Natural Enemy Release of Mahogany (Swietenia macrophylla) on Dominica, Lesser Antilles

Despite its appeal to explain plant invasions, the enemy release hypothesis (ERH) remains largely unexplored for tropical forest trees. Even scarcer are ERH studies conducted on the same host species at both the community and biogeographical scale, irrespective of the system or plant life form. In Cabrits National Park, Dominica, we observed patterns consistent with enemy release of two introduced, congeneric mahogany species, Swietenia macrophylla and S. mahagoni, planted almost 50 years ago. Swietenia populations at Cabrits have reproduced, with S. macrophylla juveniles established in and out of plantation areas at densities much higher than observed in its native range. Swietenia macrophylla juveniles also experienced significantly lower leaf-level herbivory (~3.0%) than nine co-occurring species native to Dominica (8.4–21.8%), and far lower than conspecific herbivory observed in its native range (11%–43%, on average). These complimentary findings at multiple scales support ERH, and confirm that Swietenia has naturalized at Cabrits. However, Swietenia abundance was positively correlated with native plant diversity at the seedling stage, and only marginally negatively correlated with native plant abundance for stems ≥1-cm dbh. Taken together, these descriptive patterns point to relaxed enemy pressure from specialized enemies, specifically the defoliator Steniscadia poliophaea and the shoot-borer Hypsipyla grandella, as a leading explanation for the enhanced recruitment of Swietenia trees documented at Cabrits.

Mapping tree density at a global scale

The global extent and distribution of forest trees is central to our understanding of the terrestrial biosphere. We provide the first spatially continuous map of forest tree density at a global scale. This map reveals that the global number of trees is approximately 3.04 trillion, an order of magnitude higher than the previous estimate. Of these trees, approximately 1.39 trillion exist in tropical and subtropical forests, with 0.74 trillion in boreal regions and 0.61 trillion in temperate regions. Biome-level trends in tree density demonstrate the importance of climate and topography in controlling local tree densities at finer scales, as well as the overwhelming effect of humans across most of the world. Based on our projected tree densities, we estimate that over 15 billion trees are cut down each year, and the global number of trees has fallen by approximately 46% since the start of human civilization.

On seeing the trees and the forest: Single-signal and multisignal analysis of periictal intracranial EEG

Epileptic seizures are associated with a dysregulation of electrical brain activity on many different spatial scales. To better understand the dynamics of epileptic seizures, that is, how the seizures initiate, propagate, and terminate, it is important to consider changes of electrical brain activity on different spatial scales. Herein we set out to analyze periictal electrical brain activity on comparatively small and large spatial scales by assessing changes in single intracranial electroencephalography (EEG) signals and of averaged interdependences of pairs of EEG signals.

Central European hardwood trees in a high-CO 2 future: synthesis of an 8-year forest canopy CO 2 enrichment project

Rapidly increasing atmospheric CO2 is not only changing the climate system but may also affect the biosphere directly through stimulation of plant growth and ecosystem carbon and nutrient cycling. Although forest ecosystems play a critical role in the global carbon cycle, experimental information on forest responses to rising CO2 is scarce, due to the sheer size of trees. Here, we present a synthesis of the only study world-wide where a diverse set of mature broadleaved trees growing in a natural forest has been exposed to future atmospheric CO2 levels (c. 550ppm) by free-air CO2 enrichment (FACE). We show that litter production, leaf traits and radial growth across the studied hardwood species remained unaffected by elevated CO2 over 8years. CO2 enrichment reduced tree water consumption resulting in detectable soil moisture savings. Soil air CO2 and dissolved inorganic carbon both increased suggesting enhanced below-ground activity. Carbon release to the rhizosphere and/or higher soil moisture primed nitrification and nitrate leaching under elevated CO2; however, the export of dissolved organic carbon remained unaltered.Synthesis. Our findings provide no evidence for carbon-limitation in five central European hardwood trees at current ambient CO2 concentrations. The results of this long-term study challenge the idea of a universal CO2 fertilization effect on forests, as commonly assumed in climate-carbon cycle models.

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.

Rainfall input, throughfall and stemflow of nutrients in a central African rain forest dominated by ectomycorrhizal trees

Incident rainfall is a major source of nutrient input to a forest ecosystem and the consequent throughfall and stemflow contribute to nutrient cycling. These rain-based fluxes were measured over 12 mo in two forest types in Korup National Park, Cameroon, one with low (LEM) and one with high (HEM) ectomycorrhizal abundances of trees. Throughfall was 96.6 and 92.4% of the incident annual rainfall (5370 mm) in LEM and HEM forests respectively; stemflow was correspondingly 1.5 and 2.2%. Architectural analysis showed that ln(funneling ratio) declined linearly with increasing ln(basal area) of trees. Mean annual inputs of N, P, K, Mg and Ca in incident rainfall were 1.50, 1.07, 7.77, 5.25 and 9.27 kg ha(-1), and total rain-based inputs to the forest floor were 5.0, 3.2, 123.4, 14.4 and 37.7 kg ha-1 respectively. The value for K is high for tropical forests and that for N is low. Nitrogen showed a significantly lower loading of throughfall and stemflow in HEM than in LEM forest, this being associated in the HEM forest with a greater abundance of epiphytic bryophytes which may absorb more N. Incident rainfall provided c. 35% of the gross input of P to the forest floor (i. e., rain-based plus small litter inputs), a surprisingly high contribution given the sandy P-poor soils. At the start of the wet season leaching of K from the canopy was particularly high. Calcium in the rain was also highest at this time, most likely due to washing off of dry-deposited Harmattan dusts. It is proposed that throughfall has an important `priming' function in the rapid decomposition of litter and mineralization of P at the start of the wet season. The contribution of P inputted from the atmosphere appears to be significant when compared to the rates of P mineralization from leaf litter.

Drought avoidance and the effect of local topography on trees in the understorey of Bornean lowland rain forest

The water relations of two tree species in the Euphorbiaceae were compared to test in part a hypothesis that the forest understorey plays an integral role in drought response. At Danum, Sabah, the relatively common species Dimorphocalyx muricatus is associated with ridges whilst another species, Mallotus wrayi, occurs widely both on ridges and lower slopes. Sets of subplots within two 4 -ha permanent plots in this lowland dipterocarp rain forest, were positioned on ridges and lower slopes. Soil water potentials were recorded in 1995-1997, and leaf water potentials were measured on six occasions. Soil water potentials on the ridges (-0.047 MPa) were significantly lower than on the lower slopes (-0.012 MPa), but during the driest period in May 1997 they fell to similarly low levels on both sites (-0.53 MPa). A weighted 40-day accumulated rainfall index was developed to model the soil water potentials. At dry times, D. muricatus (ridge) had significantly higher pre-dawn (-0.21 v. -0.57 MPa) and mid-day (-0.59 v. -1.77 MPa) leaf water potentials than M. wrayi (mean of ridge and lower slope). Leaf osmotic potentials of M. wrayi on the ridges were lower (-1.63 MPa) than on lower slopes (-1.09 MPa), with those for D. muricatus being intermediate (-1.29 MPa): both species adjusted osmotically between wet and dry times. D. muricatus trees were more deeply rooted than M. wrayi trees (97 v. 70 cm). M. wrayi trees had greater lateral root cross-sectional areas than D. muricatus trees although a greater proportion of this sectional area for D. muricatus was further down the soil profile. D. muricatus appeared to maintain relatively high water potentials during dry periods because of its access to deeper water supplies and thus it largely avoided drought effects, but M. wrayi seemed to be more affected yet tolerant of drought and was more plastic in its response. The interaction between water availability and topography determines these species' distributions and provides insights into how rain forests can withstand occasional strong droughts.