Tree seedling dynamics in two abandoned tropical dry forests of differing successional status in Yucatán,Mexico: a field experiment with N and P fertilization

The effects of nitrogen (N) and phosphorus (P) fertilization on seedling survival, recruitment, species richness and diversity in two abandoned tropical dry forests (10-yr old, young forest, and c. 60-yr old, old forest) in Yucatán, Mexico, were studied over two years. The seedling dynamics in the control plots were found to be highly seasonal with highest recruitment and lowest death rates during the rainy season. A low percentage of seedlings were resprouts; this important mechanism for forest regeneration had a higher-than-expected survival when compared to seedlings regenerated from seeds. Nutrient addition had significant effects on seedling dynamics in both of the forest regeneration stages. In the young forest, N fertilization facilitated the increase of seedling density. In the old forest, the addition of P decreased seedling diversity, while it increased the recruitment of only a few species. In both forests, P fertilization increased the survival time of seedlings when interacting with light availability and bulk density on the topsoil layer. Results suggest that low nutrient availability combined with low light availability constrain forest succession in Yucatán, Mexico. This revised version was published online in August 2006 with corrections to the Cover Date.     

Natural regeneration of selected timber species in the Republic of Congo

Natural regeneration of timber species is critical to the sustainable management of tropical forests. To understand what determines regeneration success of timber species in the Congo Basin, we evaluated whether seedling recruitment rates differed between forest logged 30 years previously and unlogged forest and determined the environmental factors that influence seedling density, growth and survival. We monitored the fate of 2186 seedlings of seven timber species within 462, 25‐m² plots located along 21 transects. We characterized seedling plots by light availability, soil nutrient availability and pH, and abundance of mammalian herbivores and then used linear and generalized linear mixed models to evaluate the variables that influenced seedling density, growth and survival. Light availability and canopy openness were 18% and 81% higher in logged than unlogged forest, and concentration of soil nutrients varied between sites. Seedling density was 32% higher in unlogged than logged forest. Taking all species together, seedling survival was positively correlated with calcium and negatively with magnesium and available phosphorus. Rates of seedling growth increased with available light. Taken separately, seedlings of the selected timber species responded differently to abiotic and biotic factors, demonstrating species‐specific regeneration requirements.     

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.     

Experimental Assessment of Factors Limiting Seedling Recruitment of an Amazonian Understory Herb

Understory herbs are an important and species-rich component of tropical forests, but little is known about factors limiting recruitment of these herbs. In a Peruvian flood plain forest, we studied seedling recruitment of the widespread clonal herb Heliconia metallica . We analyzed natural recruitment and experimentally added seeds, excluded predators and removed the understory vegetation at occupied and unoccupied sites to test whether seed limitation, predation, disturbance, and the presence of mature conspecific plants influence seedling recruitment. The number of naturally recruited seedlings was higher at flooded sites and close to flowering H. metallica ramets. Predation caused considerable seed loss shortly after sowing. Seed survival strongly increased with the openness of a site, but was not affected by the presence of mature conspecifics. Seed addition increased the number of seedlings at both occupied and unoccupied sites. Two years after sowing, the number of established seedlings and their size increased with light availability at a site. In experimentally disturbed plots, light availability, seedling establishment, and survival were higher. The effect of disturbance on seedling survival was stronger at occupied sites, probably due to strong light competition from mature H. metallica plants. Herbivore damage on leaves of seedlings was lower at occupied sites, but seedlings grew faster at unoccupied sites. We conclude that seedling recruitment of H. metallica depends on natural disturbance and is limited by both seed availability and the presence of mature conspecific plants.     

Effects of Nutrient Limitation on Aboveground Carbon Dynamics during Tropical Dry Forest Regeneration in Yucatán,Mexico

Tree growth (as diameter increment), litterfall production, and litter biomass were studied in two secondary tropical dry forests of the Yucatán Peninsula under four treatments of nutrient addition. The studys objective was to assess how variations in the nutrient supply affect aboveground net primary production and carbon (C) accumulation on the floor of two forests in different stages of regeneration. The study included an area of young forest (10 years old) with phosphorus (P)-poor soils and an area of old forest (around 60 years old) where soil P was comparatively less limiting. Four replicate plots (12 × 12 m) at each forest were either left intact (controls) or fertilized with nitrogen (N), P, or N plus P during 3 consecutive years. After 3 years of fertilization, relaxation of the constraints on nutrient limitation resulted in increased trunk growth rates at both the young and old forests. This effect was more pronounced with the addition of P or N plus P (trunk growth doubled with respect to controls), whereas N addition increased tree growth by 60% in comparison to trees in plots without nutrient supplements. In both forests, there were no significant differences in litterfall production among treatments during the first 2 years after fertilization. In the 3rd year of nutrient addition, litterfall production was significantly higher in plots fertilized with N plus P compared to control plots at both forest sites; however, changes in litterfall were not accompanied by litter accumulation in the floor of the two forests. The results of this study support the hypothesis that there is nutrient limitation during tropical dry forest regeneration. They further show that it may be maintained in the long term during secondary succession.     

Seedling interactions in a tropical forest in Panama

Competition is believed to be a central force limiting local diversity and controlling the structure of plant communities. However, it has been proposed that the stressed understory environment limits total understory plant density to such low levels that competitive exclusion cannot be an important factor limiting the local diversity of understory plants. To evaluate the importance of inter-seedling competition, we performed a seedling competition experiment with five shade-tolerant species in a tropical moist forest in Panama. Three-month-old seedlings were transplanted into the forest singly or with their roots intertwined with a single conspecific or heterospecific seedling in all pairwise species combinations. If competition is important, performance (survival, stem height, and number of leaves after one and six years) would be expected to be lowest with a conspecific neighbor and greatest without a neighbor. The experiment was replicated in five 0.24-m² plots at each of 20 sites in tall secondary forest. To test whether seedling performance differed among treatments we fitted linear mixed models (LMM) and generalized linear mixed models (GLMM), treating species identity and microsite (site and plot) as random effects. The five shade-tolerant study species all experienced good establishment with relatively high survival and growth rates. The neighbor treatment consistently affected seedling performance, but the effect was always very small, both in absolute terms and relative to the much stronger species and microsite effects. Seedlings with a conspecific neighbor consistently performed worse than seedlings with a heterospecific neighbor, but having no neighbor generally did not cause superior performance relative to the other treatments. We conclude that direct competitive interactions are relatively unimportant among understory plants in humid tropical forests.     

Short dry spells in the wet season increase mortality of tropical pioneer seedlings

Variation in plant species performance in response to water availability offers a potential axis for temporal and spatial habitat partitioning and may therefore affect community composition in tropical forests. We hypothesized that short dry spells during the wet season are a significant source of mortality for the newly emerging seedlings of pioneer species that recruit in treefall gaps in tropical forests. An analysis of a 49-year rainfall record for three forests across a rainfall gradient in central Panama confirmed that dry spells of ≥10 days during the wet season occur on average once a year in a deciduous forest, and once every other year in a semi-deciduous moist and an evergreen wet forest. The effect of wet season dry spells on the recruitment of pioneers was investigated by comparing seedling survival in rain-protected dry plots and irrigated control plots in four large artificially created treefall gaps in a semi-deciduous tropical forest. In rain-protected plots surface soil layers dried rapidly, leading to a strong gradient in water potential within the upper 10 cm of soil. Seedling survival for six pioneer species was significantly lower in rain-protected than in irrigated control plots after only 4 days. The strength of the irrigation effect differed among species, and first became apparent 3–10 days after treatments started. Root allocation patterns were significantly, or marginally significantly, different between species and between two groups of larger and smaller seeded species. However, they were not correlated with seedling drought sensitivity, suggesting allocation is not a key trait for drought sensitivity in pioneer seedlings. Our data provide strong evidence that short dry spells in the wet season differentially affect seedling survivorship of pioneer species, and may therefore have important implications to seedling demography and community dynamics.     

Testing for Janzen–Connell Effects in a West African Montane Forest

The Janzen–Connell hypothesis proposes that density dependent seed and seedling mortality, combined with increasing seed and seedling survival away from the parent tree, together promote regular spacing of species and thus α diversity. This hypothesis has rarely been tested in tropical Africa, and rarely in montane forests anywhere. We tested this hypothesis using a combination of field experiments and observations in the most floristically diverse dry submontane forest in Nigeria. We investigated distance effects on seedling herbivory, seedling survival and seedling height growth. We found a significant decrease in herbivory with distance from conspecific adult trees for all three species of experimentally planted seedlings (Entandrophragma angolense, Deinbollia pinnata and Sterculia setigera), and also for naturally occurring seedlings of Pouteria altissima but not of Newtonia buchananii or Isolona pleurocarpa. The relative density of large seedlings/saplings of P. altissima, N. buchananii and I. pleurocarpa increased significantly at greater distance from conspecific adult trees; however, we found no significant distance effect on survival or height growth over 3 mo for all three experimentally planted species. Taken together, our results are some of the first to show that Janzen–Connell effects occur on the African continent and in particular montane tropical forest and suggest that such effects may be pantropical.     

Non‐Native Grass Removal and Shade Increase Soil Moisture and Seedling Performance during Hawaiian Dry Forest Restoration

Invasive non‐native species can create especially problematic restoration barriers in subtropical and tropical dry forests. Native dry forests in Hawaii presently cover less than 10% of their original area. Many sites that historically supported dry forest are now completely dominated by non‐native species, particularly grasses. Within a grass‐dominated site in leeward Hawaii, we explored the mechanisms by which non‐native Pennisetum setaceum, African fountain grass, limits seedlings of native species. We planted 1,800 seedlings of five native trees, three native shrubs, and two native vines into a factorial field experiment to examine the effects of grass removal (bulldozed vs. clipped plus herbicide vs. control), shade (60% shade vs. full sun), and water (supplemental vs. ambient) on seedling survival, growth, and physiology. Both grass removal and shade independently increased survival and growth, as well as soil moisture. Seedling survival and relative growth rate were also significantly dependent on soil moisture. These results suggest that altering soil moisture may be one of the primary mechanisms by which grasses limit native seedlings. Grass removal increased foliar nitrogen content of seedlings, which resulted in an increase in leaf‐level photosynthesis and intrinsic water use efficiency. Thus in the absence of grasses, native species showed increased productivity and resource acquisition. We conclude that the combination of grass removal and shading may be an effective approach to the restoration of degraded tropical dry forests in Hawaii and other ecologically similar ecosystems.     

Demography of the seedling bank of Manilkara zapota (L.) Royen,in a subtropical rain forest of Mexico

Selection of individuals in tropical trees, occurs mainly in the seedling phase, which in part explains the low densities of most species. The main objective of this work was to gain an insight into those factors that influence growth and survival of Manilkara zapota seedlings, one of the most abundant species in the lowland forests of Mexico and Central America. Eight 10-m² rectangular plots (5m×2m) were established, in which all < 35-cm-high M. zapota seedlings were marked, measured and enumerated at bimonthly intervals. In each census seedling height, number of leaves, length of largest leaf, type of damage and seedling death were registered. Seedling survival during 2 yr was high, reaching 82% including newly emerged seedlings. A maximum likelihood regression analysis showed that both number of leaves and length of largest leaf had a direct influence on seedling survival; however, neither type of damage nor seedling density and height had significant effect. Seedling height growth averaged 2.8 cm in the 2 yr of study. The combined effect of high seedling survival, reduced growth, and impact of physical damage mainly due to falling branches and leaves reveals the occurrence of a persistent seedling bank. Such a bank would contribute to recruitment of individuals in the juvenile and eventually in the adult stages. This seedling bank could explain the high density of individuals of M. zapota in the tropical forests of Mexico.     

The invasive grass,Melinis minutiflora,inhibits tree regeneration in a Neotropical savanna

Abstract Exotic grasses are becoming increasingly abundant in Neotropical savannas, with Melinis minutiflora Beauv. being particularly invasive. To better understand the consequences for the native flora, we performed a field study to test the effect of this species on the establishment, survival and growth of seedlings of seven tree species native to the savannas and forests of the Cerrado region of Brazil. Seeds of the tree species were sown in 40 study plots, of which 20 were sites dominated by M. minutiflora, and 20 were dominated by native grasses. The exotic grass had no discernable effect on initial seedling emergence, as defined by the number of seedlings present at the end of the first growing season. Subsequent seedling survival in plots dominated by M. minutiflora was less than half that of plots dominated by native species. Consequently, at the end of the third growing season, invaded plots had only 44% as many seedlings as plots with native grasses. Above‐ground grass biomass of invaded plots was more than twice that of uninvaded plots, while seedling survival was negatively correlated with grass biomass, suggesting that competition for light may explain the low seedling survival where M. minutiflora is dominant. Soils of invaded plots had higher mean Ca, Mg and Zn, but these variables did not account for the higher grass biomass or the lower seedling survival in invaded plots. The results indicate that this exotic grass is having substantial effects on the dynamics of the tree community, with likely consequences for ecosystem structure and function.     

长白山不同演替阶段温带森林林下草本植物对乔木幼苗的影响

为了解次生针阔混交林和阔叶红松林林下草本植物对幼苗生长和存活的影响,基于长白山次生针阔混交林样地(Ⅰ)和阔叶红松林样地(Ⅱ),以246个1 m×1 m幼苗样方中乔木幼苗为研究对象,通过去除草本植物的对照试验探究草本植物对乔木幼苗高度生长和存活率的影响。结果表明,(1)群落水平上,草本植物去除有助于林下乔木幼苗的高度生长。次生针阔混交林和阔叶红松林中幼苗高度生长量在除草后较对照组均有显著提高,且阔叶红松林中幼苗高度增长在对照组和处理组中均高于次生针阔混交林。(2)去除草本植物对不同年龄级水平乔木幼苗高度生长影响不同。次生针阔混交林中,去除草本显著促进四年生及以上幼苗高度生长,对一至三年生幼苗影响不显著;阔叶红松中去除草本显著促进一至三年生幼苗高度生长,对四年生及以上幼苗影响不显著。(3)除草处理后,水曲柳幼苗高度生长量在两处样地均显著增加,假色槭幼苗高度增长量只在次生针阔混交林中显著增加,而其他幼苗高度增长量只在阔叶红松林中显著增加。(4)次生针阔混交林中,幼苗存活率与草本多度和物种数呈正相关关系,与草本盖度无相关关系;阔叶红松林中幼苗存活率与草本物种数呈正相关关系,与草本多度和盖度无相关关系。结果表明,草本植物会抑制乔木幼苗高度生长;虽可能在一定程度上有助于改善微生境,但未显著提高幼苗存活率。     

Effects of position, understorey vegetation and coarse woody debris on tree regeneration in two environmentally contrasting forests of north-western Patagonia: a manipulative approach

Aim To investigate the differential effects of position within gaps, coarse woody debris and understorey cover on tree seedling survival in canopy gaps in two old‐growth Nothofagus pumilio (Poepp. & Endl.) Krasser forests and the response of this species to gaps in two forests located at opposite extremes of a steep rainfall gradient. Location Nahuel Huapi National Park, at 41° S in north‐western Patagonia, Argentina. Methods In both study sites, seedlings were transplanted to experimental plots in gaps in three different positions, with two types of substrate (coarse woody debris or forest floor), and with and without removal of understorey vegetation. Survival of seedlings was monitored during two growing seasons. Soil moisture and direct solar radiation were measured once in mid‐summer. Seedling aerial biomass was estimated at the end of the experiment. Results Mid‐summer soil water potential was lowest in the centre of gaps, in plots where the understorey had been removed, and highest at the northern edges of gaps. Direct incoming radiation was highest in gap centres and southern edges, and lowest at northern edges. Seedling mortality was highest in gap centres, in both sites. Coarse woody debris had a positive effect on seedling survival during summer in the mesic forest and during winter in the xeric forest. The removal of understorey cover had negative effects in gap centres during summer. Seedling final aerial biomass was positively affected by understorey removal and by soil substrate in both sites. In the dry forest gaps, seedling growth was highest in northern edges, whereas it was highest in gap centres in the mesic forest. Overall growth was positively related to survival in the xeric forest, and negatively related in the mesic forest. Main conclusions Survival and growth were facilitated by the shade of gap‐surrounding trees only in the xeric forest. Understorey vegetation of both forests facilitated seedling survival in exposed microsites but competed with seedling growth. Nurse logs were an important substrate for seedling establishment in both forests; however, causes of this pattern differed between forests. Water availability positively controls seedling survival and growth in the xeric forest while in the mesic forest, survival and growth are differentially controlled by water and light availability, respectively. These two contrasting old‐growth forests, separated by a relatively short distance along a steep rainfall gradient, had different yet unexpected microenvironmental controls on N. pumilio seedling survival and growth. These results underscore the importance of defining microscale limiting factors of tree recruitment in the context of large‐scale spatial variation in resources.     

Contrasting species responses to continued nitrogen and phosphorus addition in tropical montane forest tree seedlings

Global changes in nutrient deposition rates are likely to have profound effects on plant communities, particularly in the nutrient‐limited systems of the tropics. We studied the effects of increased nutrient availability on the seedlings of six tree species in montane forests of southern Ecuador in situ. After five years of continued N, P, or N+P addition, naturally grown seedlings of each of the two most common species at each elevation (1000, 2000, and 3000 m asl) were harvested for analyses of leaf morphology, nutrient content, herbivory, and tissue biomass allocation. Most species showed increased foliar N and P concentrations after addition of each respective element. Leaf tissue N:P ratios of >20 in the control plants of all species suggest that P is more growth‐limiting in these forests than N. Leaf morphological responses to nutrient addition were species and nutrient specific, with some species (Hedyosmum purparescens, Graffenrieda emarginata) exhibiting increased specific leaf area (SLA), and others (Graffenrieda harlingii) increased leaf area ratios (LAR). Pouteria torta (1000 m) had lower SLA and LAR after P addition. Increased herbivory was only evident in G. emarginata (after N and N+P addition). Only the species from 3000 m asl modified biomass allocation after nutrient addition. In general, N and N+P addition more strongly affected the species studied at the upper elevations, whereas P addition had a similar range of effects on the species at all elevations. We conclude that the responses of the studied tropical montane forest tree seedlings to chronic N and P addition are highly species‐specific and that successful adaptation to increased nutrient availability will depend on species‐specific morphological and physiological plasticity.     

Fertilization with nitrogen and phosphorus increases abundance of non-native species in Hawaiian montane forests

Weexamined the effects of fertilization on the diversity, abundance, and cover ofthe understory plant community of two montane wet forests in Hawaii. One siteoccupies a young substrate, where aboveground tree growth is limited bynitrogen(N), while the other site is on an older substrate, where aboveground treegrowth is limited by phosphorus (P). Both sites contained an on-going,long-termfactorial fertilization experiment in which plots were fertilized semi-annuallywith N, P, or N and P in combination. In each fertilization treatment, wemeasured density of species 0.5 m tall and percent cover ofspecies <0.5 m tall. Fertilization with N reducedspeciesrichness at the young, N-limited site, but none of the nutrient additionsaltered species richness at the older, P-limited site. Species diversity andevenness were not affected by fertilization at either site. At the site withlowN availability, plots fertilized with NP had higher densities of the non-nativeginger Hedychium gardnerianum, and at the site with lowP-availability, densities of the exotic shrub Rubusargutuswere higher in P- and NP-fertilized plots. Other effects included declines inmoss cover with fertilization at both sites, and reduced abundance of nativeseedlings in response to N and NP addition at the N-limited site. Continuedlong-term fertilization could lead to greater dominance of non-native speciesbyencouraging their growth at the expense of native species, which may sufferdecreased recruitment as fertilization and increased abundance of thenon-nativespecies may reduce suitable substrates for seedling establishment.     

Seedling growth declines in warmed tropical forest soils

The response of plants to a warming climate could have a large feedback on further climatic change. This feedback is especially important for tropical forests, where the global peak in plant productivity and biodiversity occurs. Here we test the response of tropical forest tree seedling growth, photosynthesis and herbivory to 3 years of in situ full-soil profile warming. We studied six species, three of which are known nitrogen-fixers and we hypothesized that the warming response of growth will be mediated by nutrient availability to plants. Across species, growth was significantly lower in warmed soil compared to soil at ambient temperature, and the same pattern was observed for light-saturated photosynthesis, pointing toward a growth decline associated with decreased C fixation. Within species, the relative growth decline was significant for two species, Inga laurina and Tachigali versicolor, both of which are N-fixers. Together our results suggest a growth decline may have resulted from a negative effect of warming on N-fixation, rather than via changes in nutrient mineralization from soil organic matter, which was unchanged for N and increased for P during the dry-to-wet season transition. Overall, our study demonstrates that belowground warming causes species-specific declines in the growth and photosynthesis of seedlings, with a suggestion—requiring further investigation—that this growth decline is larger in N-fixing species.     

不同氮磷添加浓度对豆科3种树木幼苗生长及生物量分配的影响

选取海南尖峰岭热带山地雨林中不同功能类群的豆科树木幼苗（代表低氮需求的长脐红豆Ormosia balansae Drake、中氮需求的荔枝叶红豆Ormosia semicastrata Hance f. litchifolia How和高氮需求的猴耳环Archidendron clypearia（Jack）I.C.Nielsen）为对象,设置5个浓度梯度的氮（N）添加和2个浓度梯度的磷（P）添加养分控制实验,研究苗木的生长表现。结果显示：（1）氮添加条件下,3个树种幼苗的苗高、总叶面积、根长、根表面积、生物量5个指标对中高浓度氮添加的敏感性大小均为长脐红豆 > 猴耳环 > 荔枝叶红豆;其中,叶总面积对氮肥浓度变化的响应最敏感,长脐红豆的根长、根表面积以及猴耳环根长的响应敏感性次之。（2）长脐红豆和猴耳环幼苗的根冠比受氮肥添加浓度的影响不显著;荔枝叶红豆幼苗的根冠比则随氮肥添加浓度的升高而增大,这种适应策略反映出荔枝叶红豆幼苗对添加中高浓度氮肥有较强的耐受能力。（3）磷添加条件下,长脐红豆和猴耳环幼苗的生长速率为低磷>高磷,表明这2个树种在幼苗阶段为低氮、低磷需求;荔枝叶红豆在低氮处理下的生长速率为高磷>低磷,表明该树种幼苗阶段为低氮、高磷需求。     

Rainfall seasonality and drought performance shape the distribution of tropical tree species in Ghana

Tree species distribution in lowland tropical forests is strongly associated with rainfall amount and distribution. Not only plant water availability, but also irradiance, soil fertility, and pest pressure covary along rainfall gradients. To assess the role of water availability in shaping species distribution, we carried out a reciprocal transplanting experiment in gaps in a dry and a wet forest site in Ghana, using 2,670 seedlings of 23 tree species belonging to three contrasting rainfall distributions groups (dry species, ubiquitous species, and wet species). We evaluated seasonal patterns in climatic conditions, seedling physiology and performance (survival and growth) over a 2‐year period and related seedling performance to species distribution along Ghana's rainfall gradient. The dry forest site had, compared to the wet forest, higher irradiance, and soil nutrient availability and experienced stronger atmospheric drought (2.0 vs. 0.6 kPa vapor pressure deficit) and reduced soil water potential (?5.0 vs. ?0.6 MPa soil water potential) during the dry season. In both forests, dry species showed significantly higher stomatal conductance and lower leaf water potential, than wet species, and in the dry forest, dry species also realized higher drought survival and growth rate than wet species. Dry species are therefore more drought tolerant, and unlike the wet forest species, they achieve a home advantage. Species drought performance in the dry forest relative to the wet forest significantly predicted species position on the rainfall gradient in Ghana, indicating that the ability to grow and survive better in dry forests and during dry seasons may allow species to occur in low rainfall areas. Drought is therefore an important environmental filter that influences forest composition and dynamics. Currently, many tropical forests experience increase in frequency and intensity of droughts, and our results suggest that this may lead to reduction in tree productivity and shifts in species distribution.     

Litter mixture effects on tropical tree seedling growth--a greenhouse experiment

Decomposing litter provides critical nutrients for plants, particularly in nutrient-poor ecosystems such as tropical forests. We hypothesised that decomposing litter improves the performance of a variety of tropical tree seedlings, and that this litter effect varies depending on the species of litter present in litter mixtures. We addressed these hypotheses with a large pot experiment manipulating a range of different litter mixtures of contrasting quality and using seedlings of four tree species from the Amazonian forest of French Guiana. In contrast to our initial hypothesis, decomposing litter had either neutral or negative impacts on seedling growth, despite strongly different growth rates, biomass allocation patterns and leaf and root traits among tree species. Tree species varied in their responses to litter additions, which were further modified by species identity of the added litter. Our data show litter species-specific effects on growth, biomass allocation and leaf and root traits of tropical tree seedlings. These results suggest that a net nutrient release from decomposing litter does not necessarily improve tree seedling growth, even under nutrient-limiting conditions. In conclusion, litter layer composition may affect seedling establishment and recruitment success beyond litter-derived plant nutrient availability, which may contribute to tree species composition and dynamics in the studied tropical forest.     

Seedling Mortality and Herbivory Damage in Subtropical and Temperate Populations: Testing the Hypothesis of Higher Herbivore Pressure Toward the Tropics

Herbivory rates are generally thought to be higher in tropical than in temperate forests. Nevertheless, tests of this biogeographic prediction by comparing a single plant species across a tropical-temperate range are scarce. Here, we compare herbivore damage between subtropical and temperate populations of the evergreen tree Aextoxicon punctatum (Olivillo), distributed between 30° and 43° S along the Pacific margin of Chile. To assess the impact of herbivory on Olivillo seedlings, we set up 29 experimental plots, 1.5 × 3 m: 16 in forests of Fray Jorge National Park (subtropical latitude), and 13 in Guabún, Chiloé Island (temperate latitude). Half of each plot was fenced around with chicken wire, to exclude small mammals, and the other half was left unfenced. In each half of the plots we planted 16 seedlings of Olivillo in December 2003, with a total of 928 plants. Seedling survival, leaf production and herbivory by invertebrates were monitored over the next 16 mo. Small mammal herbivores killed ca 30 percent of seedlings in both sites. Nevertheless, invertebrate herbivory was greater in the temperate forest, thus contradicting the expected trend of increasing herbivore impact toward the tropics. Seedling growth was greater in subtropical forest suggesting better conditions for tree growth or that higher invertebrate herbivory depressed seedling growth in the temperate forest. Invertebrate herbivory increased toward temperate latitudes while small mammal herbivory was similar in both sites. We suggest that comparison of single species can be useful to test generalizations about latitudinal patterns and allow disentangling factors controlling herbivory patterns across communities.