Effects of Light Gaps and Litter Removal on the Seedling Performance of Six African Timber Species1

Valuable timber tree species frequently show poor regeneration after selective logging in tropical forests. Small size of logging gaps, lack of soil disturbance, and limited seed availability have each been blamed for observed regeneration failures. We investigated seed germination and seedling performance using a split‐plot factorial design involving light availability and litter removal for six Central African timber tree species, hypothesizing that canopy gaps and litter removal would improve seedling establishment, and that less shade‐tolerant species would show stronger responses to both factors. Contrary to our expectations, significantly more germinants established on intact litter than on exposed mineral soil 3 mo after seeding. After 18 mo, seedling survival, height and diameter growth, leaf area, and rooting depth were all much higher in gap plots than in the understory for all species, with the exception of Gilbertiodendron dewevrei, a highly shade‐tolerant species whose survival was higher in the understory. Leaf production was negatively influenced by litter removal in the least shade‐tolerant species, Nauclea diderrichii, with weak or positive effects in other species. G. dewevrei, while displaying a low‐light threshold for growth, exhibited a surprisingly high growth response to increasing light comparable to more shade‐intolerant species, a response that may help explain its local competitive dominance in the region. Due to the rapid closure of small gaps, we suggest that shade‐intolerant species such as N. diderrichii, Khaya anthotheca, and Entandrophragma utile might benefit from more intensive silvicultural practices that create larger canopy gaps.     

Carbon Translocation as Affected by Shade in Saplings of Shade Tolerant and Intolerant Species

Carbon translocation was affected by shade in different tropical tree species differing in successional status and degree of shade tolerance. Plants of the early-successional shade-intolerant species Cecropia pachystachya and Schizolobium parahyba and of the late-successional shade-tolerant species Myroxylon peruiferum and Hymenaea courbaril were grown under full sun (FS) and natural shade treatments (NS) and assessed for [¹⁴C]-sucrose translocation. Most of the ¹⁴C was retained in the fed leaf after a 24 h translocation period. Under FS, the growing apical part of the plant was the most intense sink for most species. Shade affected growth and sink intensity differently in early and late successional species. Growth was more markedly affected in the early species. Whereas these continued to invest carbon into the growing apical part of the plant under shade conditions, the late successional species invested relatively more into other sinks. This revised version was published online in July 2006 with corrections to the Cover Date.     

Antioxidant and Pigment Composition during Autumnal Leaf Senescence in Woody Deciduous Species Differing in their Ecological Traits

Abstract: Photoprotection mechanisms have been studied during autumnal senescence in sun and shade leaves of woody plants with different ecological characteristics and senescence patterns. Three of them belonging to the same family, Betulaceae: the shade‐intolerant and early successional species (Betula alba L.), the shade‐tolerant and late successional species (Corylus avellana L.), and an N‐fixing tree with low N resorption efficiency (Alnus glutinosa L.). The other two species: a shade‐intolerant (Populus tremula L.) and a shade‐tolerant (Cornus sanguinea L.), were chosen because of their ability to accumulate anthocyanins during autumnal leaf senescence. The study of plants with different ecological strategies allowed us to establish general trends in photoprotection mechanisms during autumnal senescence, when nutrient remobilisation occurs, but also during whole leaf ontogeny. We have not found a clear relationship between shade tolerance and the level of photoprotection; the main difference between both groups of species being the presence of α‐carotene in shade leaves of shade‐tolerant species. Preceding autumn, nitrogen resorption started in mid‐summer and occurred in parallel with a slight and continuous ascorbate, chlorophyll and carotenoid degradation. However, the ascorbate pool remained highly reduced and lipid oxidation did not increase at this time. Contrasting with ascorbate, α‐tocopherol accumulated progressively in all species. Only during the last stages of senescence was chlorophyll preferentially degraded with respect to carotenoids, leading to the yellowing of leaves, except in A. glutinosa in which a large retention of chlorophyll and N took place. Senescing leaves were characterised, except in C. sanguinea, by a relative increase in the proportion of de‐epoxidised xanthophylls: zeaxanthin, antheraxanthin and lutein. The light‐induced accumulation of anthocyanins in C. sanguinea could play an additional protective role, compensating for the low retention of de‐epoxidised xanthophylls. These different strategies among deciduous species are consistent with a role for photoprotective compounds in enhancing nitrogen remobilization and storage for the next growing season.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

Seed size and the regeneration niches of one rare (Melaleuca deanei) and three common (Melaleuca styphelioides,Melaleuca thymifolia and Melaleuca nodosa) Melaleuca (Myrtaceae) species of the Sydney region

Melaleuca species occupy varied habitats across Australia with 16 members of the genus occurring in the Sydney district including the rare species Melaleuca deanei F. Muell. Little is known of their germination and recruitment requirements. This paper reports on experiments assessing the effects of temperature, water potential, fire cues, light and shade levels on germination in four species of Melaleuca native to the Sydney region. For the shade experiment, seedling survival at 12 months is also reported. The experiments tested the hypothesis that M. deanei, which exhibits few seedlings in the field, has limited seedling recruitment because of its particular requirements for germination and establishment. Further, that it differs in these requirements from three common congenerics: M. nodosa (Sol ex Gaertn.) Sm., M. thymifolia Sm. and M. styphelioides Sm. Results indicate that M. deanei has a substantially similar temperature and water potential range for germination to the common congeners from 15 to 35°C, and 0 to ?0.65 ψ. Melaleuca nodosa displayed the broadest regeneration niche on all factors assessed. Germination of M. styphelioides was significantly reduced in the dark and M. styphelioides and M. deanei were most sensitive to shade in the seedling establishment/post‐germination phase. Melaleuca deanei had significantly larger seed that was slower to germinate in all experiments. Germination of the four species was unaffected by the application of heat and smoke. The substantially similar germination parameters exhibited by the four species, despite differences in habitat, may reflect their close phylogenetic affinities. The effects of fire on stimulating seed release from the canopy, the high‐light environment post‐fire together with adequate follow‐up rainfall may all be critical in seedling establishment for M. deanei.     

Allometry and Shade Tolerance in Pole‐Sized Trees of Two Contrasting Dipterocarp Species in Sabah,Malaysia1

We compared the allometry of two contrasting late‐successional dipterocarp species to test whether a monolayer (shade‐tolerant)–multilayer (shade‐intolerant) model applies to pole‐sized trees. Crown traits of the more shade‐tolerant species (Vatica micrantha) did not conform to either of the familiar monolayer or multilayer models for pole‐sized trees, but instead were consistent with a “persistent multilayer” model. Species differences in crown traits may be influenced more by future rather than present light environments.     

Regeneration niche of three epiphytic species of Gesneriaceae from Chilean rainforests: implications for the evolution of growth habits in Coronanthereae

Ecological and evolutionary studies of the epiphytic growth habit in angiosperms are limited. In this article, we assess the relationship between growth habit and regeneration niche in Coronanthereae (Gesneriaceae) and discuss its implications for the evolution of epiphytism in this lineage. In the temperate rainforest of southern Chile, we quantified the vertical distribution and experimentally examined the regeneration niche of three endemic species of Coronanthereae. One species was a holoepiphyte, which was more frequent in the upper canopy, and two species were secondary hemiepiphytes, which decreased in abundance with tree height. Seed germination of the holoepiphyte was higher on tree bark substrates and under open canopy than on forest soil and in the shade. In contrast, seed germination of both secondary hemiepiphytes did not differ between substrates (bark vs. soil) or light conditions (light vs. shade). Seedling survival percentage of secondary hemiepiphytes was higher on forest soil and under a closed canopy, thus behaving as shade‐tolerant species. In turn, the holoepiphyte behaved as a shade‐intolerant species. The reconstruction of the ancestral growth habits and regeneration niches on the inferred phylogenetic tree of Coronanthereae revealed that the specialized regeneration niche of Sarmienta repens, characterized by requirements of shade intolerance and germination on tree bark, was coupled with the evolution of the holoepiphytic growth habit. We conclude that differentiation in the regeneration niche is a key process in the evolution of epiphytic growth habits in Coronanthereae. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 79–92.     

Factors Affecting Seed Germination and Seedling Establishment of Fen-Meadow Species

Availability of seeds and provision of “safe sites” for seedling recruitment are essential for successful restoration of seminatural grassland communities. Inability to provide species‐specific conditions for seedling recruitment appears to be a major factor limiting establishment of fen‐meadow species on restoration sites. This contention was tested in the field for both germination and establishment conditions for a selection of fen‐meadow species. A Cirsio‐Molinietum fen meadow and an agriculturally semi‐improved species‐poor grass dominated rush pasture were used. Seeds of Carex ovalis, Cirsium dissectum, Molinia caerulea, Succisa pratensis, and Holcus lanatus were sown onto treatments comprising either irrigation or no irrigation, presence or absence of existing vegetation canopy, and presence or absence of soil disturbance. Germination of all except H. lanatus was higher in the fen meadow than in the rush pasture. The fen‐meadow site was less susceptible to drought, provided more light to the seed environment, and showed a stronger day–night variation in relative humidity compared with the rush pasture. All the fen‐meadow species responded strongly to the experimental treatments, whereas H. lanatus showed only a small response. Soil disturbance was the major factor that increased germination. Removal of the vegetation canopy improved germination only in S. pratensis. Conditions affecting survival of seedlings were different from those affecting seed germination. Seedling survival was greater on the fen‐meadow site than on the rush pasture. Canopy presence was the major factor that reduced seedling survival. Few seedlings survived in the presence of the rush pasture canopy. Irrigation and soil disturbance were of minor importance for seedling survival on both sites. Safe sites for seed germination and seedling establishment of fen‐meadow species existed on the fen meadow even without soil disturbance and gap creation. Safe sites for seedling recruitment were not present in the rush pasture. The need for species‐specific definition of safe site characteristics at the two stages of seedling recruitment (i.e., for seed germination and for seedling survival) was demonstrated. The implications of these findings for restoration of seminatural grasslands are discussed.     

Changes in woody vegetation abundance and diversity after natural disturbances causing different levels of mortality

Questions: How does woody vegetation abundance and diversity differ after natural disturbances causing different levels of mortality? Location: Abies balsamea–Betula papyrifera boreal mixed‐wood stands of southeast Quebec, Canada. Methods: Woody vegetation abundance and diversity were quantified and compared among three disturbance‐caused mortality classes, canopy gap, moderate‐severity disturbances, and catastrophic fire, using redundancy analysis, a constrained linear ordination technique, and diversity indices. Results: Substantial changes in canopy tree species abundance and diversity only occurred after catastrophic fire. Shade‐tolerant, late‐successional conifer species remained dominant after canopy gap and moderate‐severity disturbances, whereas shade‐intolerant, early‐successional colonizers dominated canopy tree regeneration after catastrophic fire. Density and diversity of mid‐tolerant and shade‐intolerant understory tree and shrub species increased as the impact of disturbance increased. Highest species richness estimates were observed after catastrophic fire, with several species establishing exclusively under these conditions. Relative abundance of canopy tree regeneration was most similar after canopy gap and moderate‐severity disturbances. For the sub‐canopy tree and shrub community, relative species abundances were most similar after moderate‐severity disturbances and catastrophic fire. Vegetation responses to moderate‐severity disturbances thus had commonalities with both extremes of the disturbance‐caused mortality gradient, but for different regeneration layers. Conclusions: Current spatio‐temporal parameters of natural disturbances causing varying degrees of mortality promote the development of a complex, multi‐cohort forest condition throughout the landscape. The projected increase in time intervals between catastrophic fires may lead to reduced diversity within the system.     

A test of ecological succession hypotheses using 55‐year time‐series data for 361 boreal forest stands

Aim There has been much work on succession over many decades, but succession fundamentals are still debated because of the reliance on chronosequences and dendrochronological reconstruction, both of which are problematic approaches. Here we use time‐series data to test four hypotheses that lie at the heart of successional theory: (1) the neighbourhood effect hypothesis – tree species abundance is time dependent; (2) the density‐dependence hypothesis – a rare species is more favoured over time; (3) the resource ratio hypothesis – species that can grow at the lowest resource level tend to dominate resource limited sites through succession; and (4) the intermediate disturbance hypothesis – intermediate disturbances increase the abundance of rare species. Location Central boreal forest in Canada (47°50′–50°10′ N; 80°10′–85°50′ W). Methods We used repeated measurements from sequential aerial photography and ground surveys for 361 fire‐origin stands that were measured over a c. 55‐year period. Results Shade‐intolerant Pinus banksiana decreased, tolerant Thuja occidentalis increased, intolerant Populus spp. and Betula papyrifera displayed a U‐shaped trend, and intermediate‐tolerant Picea spp. and tolerant Abies balsamea did not change with time since fire, showing evidence of negative, positive, or neutral neighbourhood effects. Species either persisted for longer or increased more in non‐conspecific stands, and had higher increases in abundance when associated with species of contrasting shade tolerance, supporting the density‐dependence hypothesis and indicating shade‐tolerance complementarity as a mechanism for coexistence. Resource‐poor soils favoured those species capable of tolerating limited resources, whereas rich soils permitted invasion and promoted mixtures, supporting the resource ratio hypothesis. Intermediate disturbances increased the invasibility of rare species. Main conclusions Contrary to previous studies where time since a stand‐replacing disturbance is the sole predictor for succession, our study shows that time alone is either an insufficient predictor or is irrelevant to species dynamics in the boreal forest. Rather, density dependence, site resource and intermediate disturbances are key mechanisms in species dynamics and coexistence over time.     

Ant-mediated expansion of an obligate seeder species during the first years after fire

Most obligate seeder species build up a soil seed bank that is associated with massive seed germination in the year immediately after a fire. These species are also shade‐intolerant and disappear when vegetation cover closes, creating unsuitable conditions for seedling recruitment. The only way for these plants to expand their populations is when habitats suitable for seedling recruitment arise (i.e. in years immediately after a fire). However, short primary seed dispersal of obligate seeders does not allow these plants to colonise the suitable habitats, and these habitats can only be colonised by secondary seed dispersion. We hypothesised that Fumana ericoides, an obligate‐seeding small shrub, not only establishes abundantly in the first year after fire, but also expands its local range in the following years due to secondary dispersal by ants while suitable habitats are still available. We tested this hypothesis using experimental studies and a simulation model of potential population expansion in a recently burned area. Results showed that F. ericoides not only established prolifically in the year immediately after fire, but was also able to recruit new individuals and expand its population in the years following the fire, despite a low germination rate and short primary seed dispersal. Ant‐mediated seed dispersal and availability of suitable habitats were key factors in this phenomenon: ants redistributed seeds in suitable habitats while they were available, which accelerated the expansion of F. ericoides because new plants established far away from the core population.     

Tree seedling responses to in situ CO2-enrichment differ among species and depend on understorey light availability

Seedlings of six major European temperate forest tree species (Fagus sylvatica, Acer pseudoplatanus, Quercus robur, Taxus baccata, Abies alba, Pinus sylvestris) were exposed to 360, 500, and 660 μL CO₂ L^?1 in the understorey of a 120‐y‐old forest over two growing seasons. Seedlings rooted in the natural forest soil within 36 open‐top chambers (12 OTCs per CO₂ treatment), each with a different known quantum flux density (QFD) ranging from 0.36 to 2.16 mol m^?2 d^?1 (= 0.8% to 4.8% of full sun). In contrast to a frequent assumption the natural CO₂ concentration in the understorey is close to the ambient concentration in the free atmosphere during daytime. The CO₂‐effect on seedling growth differed greatly among species and was strongly codetermined by microsite‐specific QFD. Biomass production in the deep‐shade tolerant species Fagus and Taxus increased by 73% and 37% under elevated CO₂ in low QFD microsites but was not significantly different among CO₂‐treatments in high QFD microsites. The less shade‐tolerant species Acer, Quercus, and Abies showed no significant response to elevated CO₂ in low QFD microsites, but increased their biomass by 39%, 25%, and 55% in high QFD microsites. In the shade‐intolerant Pinus, seedling survival was too low for a safe conclusion. Our data showed that the largest relative responses to increasing CO₂ occurred at a comparatively small increase from 360 to 500 μL L^?1 with only small and non‐significant changes with a further increase to 660 μL L^?1. Subtle shifts in the availability of light can totally reverse interspecific differences in the CO₂ response. Given these different responses, we conclude that increasing atmospheric CO₂ is likely to induce changes in species composition of temperate forests due to altered chances of recruitment. However, these shifts will depend on light patterns in the understorey, and thus on canopy structure, disturbance patterns and forest management.     

Seed germination in temperate rain forest species of southern Chile: chilling and gap-dependency germination

Canopy gap and chilling requirements for seed germination were assessed in 61 and 44 species, respectively, in the temperate rain forest of southern Chile. Germination assays within canopy gap and understorey were carried out under natural conditions. Germination tests in cold stratified and non-stratified seeds were performed under laboratory conditions. Seeds were collected of common trees, shrubs, vines and perennial herbs of forests in southern Chile. Final percent germination was significantly enhanced under canopy gap conditions in 19 species, and significantly reduced in 11 species. Germination proved indifferent under gap vs. understorey conditions in half the species tested. Cold stratification affected germination in 11 species: significantly increasing final germination of four species, and significantly decreasing final percent germination of seven species. A Principal Components Analysis (PCA) was applied with the purpose of identifying groups of species with similar germination strategies. Four attributes were included in the PCA: final germination percentage in canopy gap, germination rate in the laboratory, and gap-and chilling-dependency indices (EGAP and STRAT, respectively). The first axis separated species mainly on EGAP variation while the second axis separated them mainly according to STRAT variation; the two axes together explaining 73% of the among-species variation. A small group of trees and vines germinating best in the understorey and neutral to chilling could clearly be distinguished from the remaining species analyzed. Multifactorial ANOVAs were used for evaluating the combined effect of successional status, seed mass, dispersal period, life form, phylogenetic categories, and dispersal syndrome on EGAP and STRAT variation. The EGAP value of secondary successional species proved significantly greater than that of primary successional species and species with endozoochorous seed dispersal were significantly less dependent on chilling (according to STRAT values) compared to species with mainly wind-dispersed seeds. The possible implications of these results for seedling establishment are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Recovery from clearing,cyclone and fire in rain forests of Tonga,South Pacific: Vegetation dynamics 1995–2005

Abstract In late 2001 a category 3 cyclone impacted forest plots that were established in Tonga in 1995, and additionally, one plot was accidentally burned by an escaped land‐clearing fire. Subsequent surveys provide observations of 10 years of forest dynamics in this poorly studied region, and the first reported observations of large interannual variation in juvenile (seedling and sapling) abundance in the western tropical Pacific. The severely disturbed (burned) plot was initially colonized by a non‐native early pioneer, Carica papaya L., but 3.5 years later a native pioneer, Macaranga harveyana (Muell. Arg.) Muell. Arg., was the most abundant tree species. The seedling layer included some long‐lived pioneers and shade‐tolerant species. Two mature forest plots affected only by the cyclone changed very little over a decade. Late‐successional shade‐tolerant species that dominated the overstory were also abundant as seedlings and saplings. This is in contrast with a 30‐ to 40‐year‐old, formerly cultivated, secondary forest plot that still shows no recruitment of late‐successional dominants, in spite of the proximity of remnant forest patches. This study suggests differing pathways of succession following shifting cultivation versus cyclone and fire disturbances in Tonga. Land use legacies appear to have a long‐lasting effect on community composition.     

Are variations of direct and indirect plant interactions along a climatic gradient dependent on species’ strategies? An experiment on tree seedlings

Investigating how interactions among plants depend on environmental conditions is key to understand and predict plant communities’ response to climate change. However, while many studies have shown how direct interactions change along climatic gradients, indirect interactions have received far less attention. In this study, we aim at contributing to a more complete understanding of how biotic interactions are modulated by climatic conditions. We investigated both direct and indirect effects of adult tree canopy and ground vegetation on seedling growth and survival in five tree species in the French Alps. To explore the effect of environmental conditions, the experiment was carried out at 10 sites along a climatic gradient closely related to temperature. While seedling growth was little affected by direct and indirect interactions, seedling survival showed significant patterns across multiple species. Ground vegetation had a strong direct competitive effect on seedling survival under warmer conditions. This effect decreased or shifted to facilitation at lower temperatures. While the confidence intervals were wider for the effect of adult canopy, it displayed the same pattern. The monitoring of micro‐environmental conditions revealed that competition by ground vegetation in warmer sites could be related to reduced water availability; and weak facilitation by adult canopy in colder sites to protection against frost. For a cold‐intolerant and shade‐tolerant species (Fagus sylvatica), adult canopy indirectly facilitated seedling survival by suppressing ground vegetation at high temperature sites. The other more cold tolerant species did not show this indirect effect (Pinus uncinata, Larix decidua and Abies alba). Our results support the widely observed pattern of stronger direct competition in more productive climates. However, for shade tolerant species, the effect of direct competition may be buffered by tree canopies reducing the competition of ground vegetation, resulting in an opposite trend for indirect interactions across the climatic gradient.     

Characteristics of the Psidium cattleianum invasion of secondary rainforests

Strawberry guava (Psidium cattleianum) is a shade‐tolerant shrub or small tree invader in tropical and subtropical regions and is considered among the world's top 100 worst invasive species. Studies from affected regions report deleterious effects of strawberry guava invasion on native vegetation. Here we examine the life history demographics and environmental determinants of strawberry guava invasions to inform effective weed management in affected rainforest regions. We surveyed the vegetation of 8 mature rainforest and 33 successional sites at various stages of regeneration in the Australian Wet Tropics and found that strawberry guava invasion was largely restricted to successional forests. Strawberry guava exhibited high stem and seedling densities, represented approximately 8% of all individual stems recorded and 20% of all seedlings recorded. The species also had the highest basal area among all the non‐native woody species measured. We compared environmental and community level effects between strawberry guava‐invaded and non‐invaded sites, and modelled how the species basal area and recruitment patterns respond to these effects. Invaded sites differed from non‐invaded sites in several environmental features such as aspect, distance from intact forest blocks, as well as supported higher grass and herb stem densities. Our analysis showed that invasion is currently ongoing in secondary forests, and also that strawberry guava is able to establish and persist under closed canopies. If left unchecked, strawberry guava invasion will have deleterious consequences for native regenerating forest in the Australian Wet Tropics.     

Multiple successional pathways of boreal forest stands in central Canada

Predicting forest composition change through time is a key challenge in forest management. While multiple successional pathways are theorized for boreal forests, empirical evidence is lacking, largely because succession has been inferred from chronosequence and dendrochronological methods. We tested the hypotheses that stands of compositionally similar overstory may follow multiple successional pathways depending on time since last stand‐replacing fire (TSF), edaphic conditions, and presence of intermediate disturbances. We used repeated measurements from combining sequential aerial photography and ground surveys for 361 boreal stands in central Canada. Stands were measured in 8–15 yr intervals over a ~ 60 yr period, covering a wide range of initial stand conditions. Multinomial logistic regression was used to analyze stand type transitions. With increasing TSF, stands dominated by shade‐intolerant Pinus banksiana, Populus sp., and Betula papyrifera demonstrated multiple pathways to stands dominated by shade‐tolerant Picea sp., Abies balsamea, and Thuja occidentalis. Their pathways seemed largely explained by neighborhood effects. Succession of stands dominated by shade‐tolerant species, with an exception of stands dominated by Picea sp., was not related to TSF, but rather dependent on edaphic conditions and presence of intermediate disturbances. Varying edaphic conditions caused divergent pathways with resource limited sites being dominated by nutrient‐poor tolerant species, and richer sites permitting invasion of early successional species and promoting species mixtures during succession. Intermediate disturbances promoted deciduous persistence and species diversity in A. balsamea and mixed‐conifer stands, but no evidence was detected to support “disturbance accelerated succession”. Our results demonstrate that in the prolonged absence of stand‐replacing disturbance boreal forest stands undergo multiple succession pathways. These pathways are regulated by neighborhood effects, resource availability, and presence of intermediate disturbance, but the relative importance of these regulators depends on initial stand type. The observed divergence of successional pathways supports the resource‐ratio hypothesis of plant succession.     

Seed to seedling transitions in successional habitats across a tropical landscape

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed‐dispersal limitation and seedling‐establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; Φ_i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (E_it) were higher for animal‐dispersed than wind‐dispersed species, and for non‐pioneer than pioneer species. Non‐pioneers and animal‐dispersed species had higher expected relative recruit abundance (ε_it) in primary forest (median of 4 seeds recruit^?1) whereas in secondary forest wind‐dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind‐dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (Φ_i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of Φ_i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); Φ_i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.     

Seed germination and seedling growth in the arrow bamboo <Emphasis Type="Italic">Fargesia qinlingensis</Emphasis>

Improving natural regeneration of bamboos after they die following mass flowering is critical for conservation of giant pandas. However, little is known about factors that affect seed germination and seedling growth of bamboos. We studied seed germination and seedling growth in Fargesia qinlingensis, which mass flowered in a giant panda habitat in the Qinling Mountains of China in early 2000, in laboratory and greenhouse conditions. Seed germination rate was tested under light and dark conditions 5 and 12 months after seed collection. Germination rate displayed no significant difference under light or dark conditions 5 months after seed collection, but was significantly greater in the dark than under light 12 months after seed collection, suggesting light inhibition of seed germination. A 2×2 factorial design was conducted to test the effects of nitrogen (N fertilization and non-N fertilization) and light [full sun and shade (i.e., 14% full sun)] on seedling growth and biomass allocation. N fertilization significantly increased seedling growth, resulting in greater seedling height, more branches, more leaves, greater stem biomass, and greater leaf biomass. Seedlings under 14% full sun conditions had a significantly lower percentage of biomass allocated to the stem. The root/shoot ratio was significantly greater in non-N/shade than non-N/full sun, while there was no significant difference in this ratio between N/shade and N/full sun, suggesting that nitrogen fertilization compensated for the effect of shade on biomass allocation. Our results suggest that N fertilization could be employed in restoration of F. qinlingensis stands after die-off following mass flowering.     

Leaf structure of Syzygium spp. (Myrtaceae) in relation to site affinity within a tropical rain forest

This study examined four species of Syzygium (S. firmum, S. makul, S. operculatum, S. rubicundum) Myrtaceae, a tree genus that dominates the canopy of rain forests of south‐west Sri Lanka. Syzygium spp. occupy differing habitats with relation to succession and forest topography. We examined differences in leaf morphology and physiology in response to amount of shade, an important environmental variable affecting Syzygium distribution within the forest. To study change in leaf structure and physiology, environmental shelters were constructed simulating forest shade that differed in quality, quantity and duration. Seedlings were exposed to: (i) 0% shade (full sun, FS), red : far red (R : FR) ratio 1.27; (ii) 65% shade (large opening, LO) with direct sunlight similar to the centre of a large canopy opening, R : FR ratio 1.27; (iii) 82% shade (small opening, SO) with direct sunlight similar to the centre of a small canopy opening, R : FR ratio 1.27; (iv) 58% uniform light shade (LS) with a quality similar to the outside edge of a large canopy opening, R : FR ratio 1.05; (v) 85% uniform medium shade (MS) with a quality similar to the inside forest edge of a large canopy opening, R : FR ratio 0.97; (vi) 99% uniform deep shade (DS) similar to that of the forest understorey, R : FR ratio 0.23. The shelters were constructed in a large open area at the field station of the Sinharaja World Heritage site, Sri Lanka. Seedlings of each species were grown for two years in their respective shade treatments before physiological, morphological and anatomical measurements were made on leaves. Variation in leaf structure and physiology between the species was associated with differences in shade‐tolerance and water‐use. All species increased in photosynthesis rates and dimensions in leaf structure (leaf blade and cuticle thickness, stomatal density, thickness of upper and lower epidermis, and thickness of palisade mesophyll) with decrease in shade. In contrast, stomatal conductivity was highest in the DS (99% shade) treatment. Leaves of Syzygium firmum were thickest and largest in area. S. firmum also had highest photosynthesis in the SO (82% shade) treatment. S. firmum was the most shade‐tolerant of all species: it grows well in low shade and its leaf structure suggests it to be the most conservative in water‐use of the Syzygium spp. In the forest S. firmum can persist in the forest shade as established seedlings, but grows best within canopy openings of late‐seral rain forest. Leaves of S. operculatum were thinnest but had highest stomatal densities of the four species. S. operculatum is considered shade‐intolerant, with a leaf structure suggesting it to be prone to desiccation, and by implication susceptible to drought. S. operculatum is found along streams within early seral rain forest habitat, often originating on stream banks after land clearance for cultivation. In the FS (0% shade) treatment, S. rubicundun had highest photosynthesis rates and greatest number of leaves but smallest leaf area of the Syzygium species. S. rubicundum is more shade‐intolerant but more efficient in water‐use than S. operculatum. S. rubicundum is a mid‐seral canopy tree of the midslope stands that are thought to have originated after catastrophic windthrows or swidden cultivation. The leaf physiology and structure of S. makul suggests it to be both moderately shade‐tolerant and conservative in water‐use. It is the most widely distributed Syzygium species across the topography of late‐seral rain forest. We suggest forest disturbance and hydrology are important environmental factors that influence distribution of Syzygium species across the topography. Results from this study contribute to a body of knowledge suggesting that canopy tree species of rain forests in south‐west Sri Lanka have discrete affinities to topography and differences in successional status, and that adaptations in leaf structure and physiology are indicative of such phenomena. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141 , 365–377.