Insects and light interact to mediate vine colonization of fast growing Microberlinia bisulcata tree seedlings in gaps of an African rain forest

Vines thrive in lowland tropical forests, yet the biotic factors underlying their colonization of host tree seedlings and saplings remain surprisingly understudied. Insect herbivores presumably could influence this process, especially where disturbance has opened the canopy (i.e., gaps)—temporary areas of higher primary productivity favoring the recruitment of vines and trees and invertebrates in forests—but their impact on vine colonization has never been experimentally tested. Using data from an insect herbivore exclusion (mesh-netting cages) experiment conducted in an African rain forest (Korup, Cameroon), I logistically modeled the probability of vines colonizing seedlings of three co-dominant species (Microberlinia bisulcata vs. Tetraberlinia bifoliolata and T. korupensis) in paired shaded understory and sunny gap locations (41 blocks across 80 ha, starting n = 664 seedlings) in a 1–2-yr period (2008–2009). Vine colonization occurred almost exclusively in gaps, occurring on 16% of seedlings there. Excluding herbivores in gaps doubled colonization of the light-demanding and faster growing M. bisulcata but had negligible effects on the two shade-tolerant, slower growing and less palatable Tetraberlinia species, which together were twice as susceptible to vines under natural forest gap conditions (controls). When protected from herbivores in gaps, more light to individual seedlings strongly increased vine colonization of M. bisulcata whereas its well-lit control individuals supported significantly fewer vines. These results suggest vines preferably colonize taller seedlings, and because light-demanding tree species grow faster in height with more light, they are more prone to being colonized in gaps; however, insect herbivores can mediate this process by stunting fast growing individuals so that colonization rates becomes more similar between co-occurring slow and fast growing tree species. Further influencing this process might be associational resistance or susceptibility to herbivores linked to host species’ leaf traits conferring shade-tolerant ability as seedlings or saplings. A richer understanding of how vines differentially influence forest regeneration and species composition may come from investigating vine–tree–herbivore interactions across light gradients, ideally via long-term studies and intercontinental comparisons. Abstract in French is available with online material.     

Reproductive investment and seedling survival of the mast-fruiting rain forest tree,Microberlinia bisulcata A. chev.

In the southern part of Korup National Park, Cameroon, the mast fruitingtree Microberlinia bisulcata occurs as a codominant ingroves of ectomycorrhizal Caesalpiniaceae within a mosaic of otherwisespecies-rich lowland rain forest. To estimate the amount of carbon andnutrientsinvested in reproduction during a mast fruiting event, and the consequentialseed and seedling survival, three related field studies were made in 1995.Theseprovided a complete seed and seedling budget for the cohort. Seed productionwasestimated by counting woody pods on the forest floor. Trees produced on average26,000 (range 0–92,000) seeds/tree, with a dry mass of 16.6kg/tree. Seeds were contained in woody pods of mass 307kg/tree. Dry mass production of pods and seeds was 1034 kgha^–1, equivalent to over half (55%) of annual leaflitterfall for this species, and contained 13% of the nitrogen and 21% of thephosphorus in annual leaf litterfall. Seed and young-seedling mortality wasinvestigated with open quadratsand cages to exclude vertebrate predators, at two distances from the parenttree. The proportion of seeds on the forest floor which disappeared in thefirst6 wk after dispersal was 84%, of which 26.5% was due to likelyvertebrate removal, 36% to rotting, and 21.5% to other causes. Vertebratepredation was greater close to the stem than 5 m beyond the crown(41 vs 12% of seeds disappearing) where the seed shadow was less dense.Previousstudies have demonstrated an association between mast years at Korup and highdry-season radiation before flowering, and have shown lower leaf-litterfallphosphorus concentrations following mast fruiting. The emerging hypothesis isthat mast fruiting is primarily imposed by energy limitation for fruitproduction, but phosphorus supply and vertebrate predation are regulatingfactors. Recording the survival of naturally-regeneratingM. bisulcata seedlings (6-wk stage) showed that 21% ofseedlings survived to 31 mo. A simple three-stage recruitment model wasconstructed. Mortality rates were initially high and peaked again in each ofthenext two dry seasons, with smaller peaks in the two intervening wet seasons,these latter coinciding with annual troughs in radiation. The very poorrecruitment of M. bisulcata trees in Korup, demonstratedinprevious investigations, appears not to be due to a limitation in seed oryoung-seedling supply, but rather by factors operating at theestablished-seedling stage.     

Differences in light energy utilisation and dissipation between dipterocarp rain forest tree seedlings

 The light environment within tropical rain forests varies considerably both spatially and temporally, and photon flux density (PFD) is considered to be an important factor determining the growth and survival of rain forest tree seedlings. In this paper we examine the ability of four ecologically contrasting dipterocarps (Dryobalanops lanceolata, Shorea leprosula, Hopea nervosa and Vatica oblongifolia) to utilise and dissipate light energy when grown in different light environments in lowland dipterocarp rain forest in the Danum Valley Conservation Area, Sabah, East Malaysia. Specifically we report (i) photosynthetic light response curves and associated fluorescence characteristics, including quantum yield (ΦPSII) and non-photochemical quenching (qN) and (ii) the extent to which photoinhibition occurs when plants grown in either high or low light are exposed to short bursts of high PFD. When grown in low light (artificial or forest shade) all four species had low light saturated rates of photosynthesis which were achieved at low PFDs. In addition, values of ΦPSII and qN were similar over a range of measurement PFDs. D. lanceolata and S. leprosula were also grown at high PFD and showed marked differences in their responses. S. leprosula demonstrated an ability to increase its rate of photosynthesis and there was a small increase in capacity to dissipate excess light energy non-photochemically at high PFDs. Partitioning of this qN into its fast, photo-protective (qE) and slow, photoinhibitory (qI) components indicated that there was an increase in qE quenching. In contrast, although D. lanceolata survived in the high light environment, greater rates of photosynthesis were not observed and the plants showed a greater capacity to dissipate energy non-photochemically. Partitioning of qN revealed that the majority of this increase was attributable to the slower relaxing phases. Received: 10 February 1996 / Accepted: 14 June 1996     

西双版纳原始热带季节雨林生物量研究

用标准木回归分析法（ 乔木、木质藤本） 和样方收获法（ 灌木、草本） , 研究了西双版纳原始热带季节雨林生物量及其分配。雨林总生物量为６９２－５９０ ｔ／ｈｍ２ , 其分配为： 乔木层占９８－６６ ％ 、灌木层占０－７６ ％ 、木质藤本占０－５０ ％ 、草本层占０－０９ ％ , 生物量主要集中于乔木层。乔木层生物量的器官分配向树干和树根集中： 树干占６９－８０ ％ , 树根占２１－５６ ％ , 树枝占７－７７ ％ ,树叶占０－７７ ％ ; 生物量径级分配向中等径级（６０ ～７０ ｃｍ） 和最大径级（１５０ ～１６０ ｃｍ） 集中; 生物量垂直分配向上层集中; Ⅰ亚层（ 高度＞ ４０ ｍ） 占６０－５５ ％ 、Ⅱ亚层（２０ ～４０ ｍ） 占３６－７２ ％ 、Ⅲ亚层（３ ～２０ ｍ） 占２－７３ ％ ; 优势种番龙眼生物量占乔木层的２０－０７ ％ ; 乔木层叶面积指数为６－９１ 。     

The responses to shade of seedlings of very small-seeded tree and shrub species from tropical rain forest in Singapore

1. Newly germinated seedlings of six tree and shrub species with very small seeds (31–460 μg dry mass), one light-demanding and five shade-tolerant at the stage of establishment in the wild, were grown for 5 months in neutral shade houses with 0·5, 1, 3·5 and 7·5% daylight.
2. The ratio of yield in 7·5% to that in 1% was 8:1 for the light-demanding Melastoma malabathricum but only 2:1 for the confamilial shade-tolerator Pternandra echinata. The Pternandra, Urophyllum hirsutum, Ficus chartacea, Ficus grossularioides and Pellacalyx saccardianus showed a graded series of responses to irradiance, generally consistent with their apparent demands for light in the wild. In contrast, survival of very deep shade was not clearly related to light demand in nature.
3. The results support the conclusion drawn from observational studies that large seed size is not primarily adaptive in resisting shade but in resisting the associated risks of burial by litter, desiccation during dry spells, uprooting by birds and other kinds of damage by animals or falling debris.     

Nutrient cycling in a tropical seasonal rain forest of Xishuangbanna, Southwest China. Part 1: tree species: nutrient distribution and uptake

Tropical rain forests are characterized by large numbers of the species with diverse growth habits. The objective of the present study was to determine the distribution of nutrient content in the major trees of the tropical rain forests in Xishuangbanna. This will improve the understanding of the nutrient losses from such sites that result from harvesting and flow of nutrients within the ecosystem and lead to the development of effective and rational forest management strategies. Based on the results in this study, the distribution of nutrients among biomass components of trees varied: The ordering of major elements concentrations was K>N>Mg>Ca>P in branch, stem and root tissues but was N>K>Mg>Ca>P in leaves. The maximum amount of all nutrients per ha occurred in the stems followed by branches, roots and leaves. Of the total uptake of 6167.7 kg ha⁻¹ of all nutrients, the contribution of various nutrients was found to be N (2010.6 t ha⁻¹), P (196.3 t ha⁻¹), K (2123.8 kg ha⁻¹), Ca (832 kg ha⁻¹) and Mg (1005 kg ha⁻¹). However, comparing the nutrient uptake of other tropical and sub tropical forests, the results indicated that rates for the Xishuangbanna forests were 20–35% lower than previously reported values.     

The ecology of tree reproduction in an African medium altitude rain forest

The occurrence of flowering and fruiting in tropical trees will be affected by a variety of factors, linked to availability of resources and suitable climatic triggers, that may be affected by increasing global temperatures. Community‐wide flowering and fruiting of 2526 trees in 206 plots were monitored over 24 years in the Budongo Forest Reserve (BFR), Uganda. Factors that were assessed included the size of the tree, access to light, the impacts of liana load, effects of tree growth, and variation between guilds of trees. Most flowering occurs at the end of the long dry season from February to April. Trees that had access to more light flowered and fruited more frequently. Pioneer and non‐pioneer light‐demanding species tended to reproduce more frequently than shade‐bearing species. Trees that grew faster between 1993 and 2011 also fruited more frequently. When examining all factors, growth rate, tree size, and crown position were all important for fruiting, while liana load but not growth rate was important in reducing flowering. Trees in BFR show a large decline in fruiting over 24 years, particularly in non‐pioneer light demanders, shade‐bearers, and species that produce fleshy fruits eaten by primates. The decline in fruit production is of concern and is having impacts on primate diets and potential recruitment of mahogany trees. Whether climate change is responsible is unclear, but flowering of the guilds/dispersal types which show declines is correlated with months with the coolest maximum temperatures and we show temperature has been increasing in BFR since the early 1990s.     

Element loss on rain forest conversion in East Amazonia: comparison of balances of stores and fluxes

Nutrient loss from forest ecosystems by burning and leaching can be estimated by comparing nutrient stores before and after the disturbance, and by measuring the nutrient fluxes during disturbance. We applied both methods to a clear cutting experiment near Belem, Eastern Amazon, in a correct time series of 15 months duration. Nutrient flux measurements include wood export, loss to the atmosphere by burning, and leaching. The latter was based on water flux simulation with a soil water model, and on element analysis of soil water extracts (ceramic cup lysimeter). Two plots with 33 and 92tha^–1 of residual biomass (left after export of wood >7cm diameter) were compared with a forest control plot. Store and flux balances agreed well in case of Na, K, Mg, and S, and partially also for Ca and total P. Deviations seem to be caused by erratic but statistically not significant variations of the soil stores (mainly C and total N). Spacial variability of soil parameters demand very high sample replication, especially after disturbance. Flux measurements are better but more costly.     

Effects of dung and seed size on secondary dispersal,seed predation,and seedling establishment of rain forest trees

Seeds dispersed by tropical, arboreal mammals are usually deposited singly and without dung or in clumps of fecal material. After dispersal through defecation by mammals, most seeds are secondarily dispersed by dung beetles or consumed by rodents. These post-dispersal, plant-animal interactions are likely to interact themselves, as seeds buried by dung beetles are less likely to be found by rodents than unburied seeds. In a series of three experiments with seeds of 15 species in central Amazonia (Brazil), we determined (1) how presence and amount of dung associated with seeds influences long-term seed fate and seedling establishment, (2) how deeply dung beetles bury seeds and how burial depth affects seedling establishment, and (3) how seed size affects the interaction between seeds, dung beetles, and rodents. Our overall goal was to understand how post-dispersal plant-animal interactions determine the link between primary seed dispersal and seedling establishment. On average, 43% of seeds surrounded by dung were buried by dung beetles, compared to 0% of seeds not surrounded by dung (n=2,156). Seeds in dung, however, tended to be more prone than bare seeds to predation by rodents. Of seeds in dung, probability of burial was negatively related to seed size and positively related to amount of dung. Burial of seeds decreased the probability of seed predation by rodents three-fold, and increased the probability of seedling establishment two-fold. Mean burial depth was 4 cm (0.5–20 cm) and was not related to seed size, contrary to previous studies. Probability of seedling establishment was negatively correlated with burial depth and not related to seed size at 5 or 10 cm depths. These results illustrate a complex web of interactions among dung beetles, rodents, and dispersed seeds. These interactions affect the probability of seedling establishment and are themselves strongly tied to how seeds are deposited by primary dispersers. More generally, our results emphasize the importance of looking beyond a single type of plant-animal interaction (e.g., seed dispersal or seed predation) to incorporate potential effects of interacting interactions.     

Soil acidity and nutrient deficiency in central Amazonian heath forest soils

Experiments were carried out to test the effects of liming and nutrient additions on plant growth and soil processes such as C and N mineralisation in three contrasting forest types in central Amazonia: the stunted facies of heath forest (SHF), the tall facies of heath forest (THF) and the surrounding lowland evergreen rain forest (LERF). Calcium-carbonate additions increased soil respiration in the field plots in the SHF; in laboratory incubations, soil respiration was higher in the SHF when soils were fertilised with N, and in THF and LERF after S additions. The addition of N alone or in different combinations generally induced a net immobilisation of soil N. Net nitrification increased during the incubation in SHF and THF soils fertilised with N+P, and in LERF soils fertilised with either N, or P, or CaCO₃. In a field experiment using ingrowth bags, a higher fine root production was observed in all forest types when bags were fertilised with CaCl₂ or CaCO₃, suggesting that Ca may be a limiting nutrient in these soils. Calcium-carbonate addition in a glasshouse bioassay experiment with rice showed an overall positive effect on the survival and growth of the seedlings. In other treatments where soil pH was not raised, the rice showed acute toxicity symptoms, poor root and shoot growth and high mortality. Similar results were yielded in a field experiment, using naturally established seedlings in the field plots in SHF, THF and LERF. It is concluded that the acute H⁺ ion toxicity is a major growth-limiting factor for non-adapted plants in heath forest soils in central Amazonia.     

Arbuscular mycorrhizal fungi in the tree seedlings of two Australian rain forests: occurrence, colonization, and relationships with plant performance

The roots of rain forest plants are frequently colonized by arbuscular mycorrhizal fungi (AMF) that can promote plant growth in the nutrient poor soils characteristic of these forests. However, recent studies suggest that both the occurrence of AMF on rain forest plants and the dependence of rain forest plants on AMF can be highly variable. We examined the occurrence and levels of AMF colonization of some common seedling species in a tropical and a subtropical rain forest site in Queensland, Australia. We also used a long-term database to compare the growth and mortality rates of seedling species that rarely formed AMF with those that regularly formed AMF. In both forests, more than one-third of the seedling species rarely formed AMF associations, while 40% of species consistently formed AMF in the tropical site compared to 27% in the subtropical site. Consistent patterns of AMF occurrence were observed among plant families at the two sites. Variation among seedling species in AMF occurrence or colonization was not associated with differences in seed mass among species, variation in seedling size and putative age within a species, or lack of AMF inoculum in the soil. Comparisons of four seedling species growing both in the shaded understory and in small canopy gaps revealed an increase in AMF colonization in two of the four species in gaps, suggesting that light limitation partially explains the low occurrence of AMF. Seedling survival was significantly positively associated with seed biomass but not with AMF colonization. Furthermore, seedling species that regularly formed AMF and those that did not had similar rates of growth and survival, suggesting that mycorrhizal and nonmycorrhizal strategies were equivalent in these forests. Furthermore, the high numbers of seedlings that lacked AMF and the overall low rate of seedling growth (the average seedling required 6 years to double its height) suggest that most seedlings did not receive significant indirect benefits from AMF through connection to canopy trees via a common mycorrhizal network.     

Light environment,sapling architecture,and leaf display in six rain forest tree species

Architecture and leaf display were compared in saplings of six rain forest tree species differing in shade tolerance. Saplings were selected along the whole light range encountered in a forest environment. Species differed largely in realized height and crown expansion per unit support biomass, but this could not be related to differences in shade tolerance. The results demonstrate that there exist various solutions to an effective expansion of plant height and crown area. It is argued that choice of the study species and the ontogenetic trajectory regarded determine to a large extent the outcome of interspecific comparisons. No evidence was found that pioneers were characterized by a multilayered and shade tolerants by a monolayered leaf distribution. Yet, sun plants had a similar crown area, a deeper crown, and a higher leaf area index compared to shade plants and their leaves were more evenly distributed along the stem. This suggests that differences in leaf layering are found between plants growing in different light environments, rather than between species differing in shade tolerance.     

Effect of typhoon disturbance on fine litterfall and related nutrient input in a subtropical forest on Okinawa Island, Japan

Typhoons are frequent on Okinawa Island, southwestern Japan. The effects of typhoon disturbance on the patterns of fine litterfall and related nutrient inputs in a subtropical evergreen broad-leaved forest were studied over 5 years from May 1996 to April 2001. Annual fine litterfall averaged 7558 kg ha⁻¹ (range from 6188 to 9439 kg ha⁻¹) for six sampling plots over 5 years, which differed significantly among years (p<0.001) but not among plots (p=0.122). A seasonal maximum was most evident for leaf litter component. Woody litter fell more irregularly through the year, and peak fall varied with typhoon and windstorm. The mean ratio of annual litterfall mass of sexual organs to leaves was 0.06, much lower than that in other tropical and subtropical rain forests. Nutrient concentrations varied in litterfall components, but were not significantly different among plots. The lowest concentrations of N and P in leaf litter were observed in March, which is also the month with the greatest leaf fall. However, the highest concentrations were recorded in typhoon season. Nitrogen and P concentrations were 34% and 106% greater in the green leaves that fell during typhoons than in senescent leaves. Mean nutrient inputs by litterfall were: N 83, P 3.2, K 25, Ca 71, Mg 19, Al 12, Na 10, Fe 0.86 and Mn 3.9 kg ha⁻¹ yr⁻¹, and differed significantly among years for all elements (p<0.0005) and among plots only for K (p<0.05) and Mn (p<0.0001). Typhoon disturbance strongly affected annual fine litterfall and related nutrient inputs, which contributed an average of 30% of the annual litterfall mass, and from 30% to 39% (for different nutrient elements) of annual total nutrient inputs. The results from this study suggest that typhoon-driven maintenance of rapid cycling of P and N and their high availability in soil appears to be an important mechanism to maintain productivity in the subtropical forest on Okinawa Island.     

Formation and closure of canopy gaps in the rain forest at Nouragues, French Guiana

Formation and closure of canopy gaps was monitored for three years in 12 ha of primary rain forest at Nouragues, French Guiana. At the first inventory, in April 1991, 74 openings in the canopy > 4 m² (sensu Brokaw 1982a) were located; 60 of these gaps were formed before January 1990. Between January 1990 and December 1993, 5 to 15 gaps were annually formed, opening 0.64–1.33% of the forest canopy each year. Of all gaps, 41% were created by a falling, snapped tree, 34% by a falling, uprooted tree, 22% by a falling branch, and 3% by a falling dead stem. A refined nearest neighbour analysis showed that gaps formed after January 1990 were clustered: uprooting of trees seemed to be related to shallow soils, and relatively many other trees fell when a tree uprooted, independent of the dbh of the uprooted tree. In 37 gaps, canopy openness in the gap centre (determined by hemispherical photographs) was monitored over three years. In 54% of the gaps, canopy openness increased in two successive years. It is reasoned that edges of especially large gaps may frequently be re-disturbed by falling trees or branches. Results suggest that gaps have closed after around 15 years. More data are needed to verify this.     

西双版纳热带季节雨林的树种组成和群落结构动态

研究了西双版纳热带季节雨林1 hm2(hectare)动态监测样地1993年与2007年之间树种组成和群落结构的变化。对样地中胸径≥5 cm的乔木进行了每木调查。目前其树种组成的热带分布科、属所占比例分别为91%和94%,具有较高比例的热带植物区系性质。在1993年与2007年两次调查之间,树种数量由145种增至179种,仅有1到2个个体的稀有树种所占比例从54%降为51.1%。从森林的垂直结构来看,A、B、C三层的个体死亡率分别为12.8%、12.9%和19.0%,各层树木的增长率分别为-8.5%、-1.4%和44.8%。与此相对应,C层小径级的树木所占比例有较大提高。虽然小径级的树木在种类和数量上比例增大,但个体数量和种类组成相对稳定的A、B层优势树种变化不大,维持了群落结构的稳定性。14 a间,群落中新增加的具有先锋性质的树种不超过5个。1993年时,A、B两层尚有先锋树种存在,2007年已经从A、B两层中退出。因此,从14 a间树种组成和群落结构的变化来看,虽然具有树木的死亡和增补,但其物种成分和群落结构的总体格局没有明显的变化,处于动态平衡过程中。     

Seasonality of weather and tree phenology in a tropical evergreen mountain rain forest

Flowering and fruiting as phenological events of 12 tree species in an evergreen tropical mountain rain forest in southern Ecuador were examined over a period of 3–4 years. Leaf shedding of two species was observed for 12 months. Parallel to the phenological recordings, meteorological parameters were monitored in detail and related to the flowering and fruiting activity of the trees. In spite of the perhumid climate of that area, a high degree of intra- and inter-specific synchronisation of phenological traits was apparent. With the exception of one species that flowered more or less continuously, two groups of trees could be observed, one of which flowered during the less humid months (September to October) while the second group started to initiate flowers towards the end of that phase and flowered during the heavy rains (April to July). As reflected by correlation coefficients, the all-time series of meteorological parameters showed a distinct seasonality of 8–12 months, apparently following the quasi-periodic oscillation of precipitation and related cloudiness. As revealed by power spectrum analysis and Markov persistence, rainfall and minimum temperature appear to be the only parameters with a periodicity free of long-term variations. The phenological events of most of the plant species showed a similar periodicity of 8–12 months, which followed the annual oscillation of relatively less and more humid periods and thus was in phase or in counter-phase with the oscillations of the meteorological parameters. Periods of unusual cold or dryness, presumably resulting from underlying longer-term trends or oscillations (such as ENSO), affected the homogeneity of quasi-12-month flowering events, fruit maturation and also the production of germinable seeds. Some species show underlying quasi-2-year-oscillations, for example that synchronise with the development of air temperature; others reveal an underlying decrease or increase in flowering activity over the observation period, influenced for instance by solar irradiance. As Ecuador suffers the highest rate of deforestation in South America, there is an urgent need for indigenous plant material for reforestation. A detailed knowledge of the biology of reproduction in relation to governing external factors (mainly climate) is thus required.     

Biomass,production and nutrient distribution of a natural oak forest in central Korea

Biomass, production, and nutrient distribution of a pure Quercus variabilis Bl. stand (stand 1) and two mixed Q. variabilis–Q. mongolica Fisch. stands (stand 2 and 3) were investigated in central Korea. Stand 1 naturally occurred on a site with a southern aspect while stand 2 and stand 3 occurred on sites with a northern aspect. Total (overstory+understory vegetation) biomass (tha^-1) and annual production (tha^–1year^–1) were 137.8 and 11.1 for stand 1, 216.2 and 16.6 for stand 2, and 253.3 and 19.7 for stand 3. Nutrient contents (kgha^–1) in the vegetation were distributed as follows: K, 478–860; N, 471–839; Ca, 428–791; Mg, 72–125; Na, 77–141; and P, 37–71, and were greatest in stand 3 followed by stand 2, and stand 1. Stand density influenced the differences in biomass, annual production and nutrient contents in the vegetation. Forest floor dry mass and N content (kgha^–1) were 13400 and 169 for stand 1, 10400 and 133 for stand 2, and 11200 and 127 for stand 3. Total amounts of N, P and Na in the ecosystem were greatest in the upper 40cm of mineral soil followed by the vegetation and forest floor. However, the vegetation contained a greater amount of K than the mineral soil. It appeared that microenvironments, such as, aspect influenced the distribution of natural oak species within a relatively small area and resulted in differences in biomass, production and nutrient distribution among the stands.