Incipient ecological speciation between successional varieties of a dominant tree involves intrinsic postzygotic isolating barriers

Whereas disruptive selection imposed by heterogeneous environments can lead to the evolution of extrinsic isolating barriers between diverging populations, the evolution of intrinsic postzygotic barriers through divergent selection is less certain. Long‐lived species such as trees may be especially slow to evolve intrinsic isolating barriers. We examined postpollination reproductive isolating barriers below the species boundary, in an ephemeral hybrid zone between two successional varieties of the landscape‐dominant Hawaiian tree, Metrosideros polymorpha, on volcanically active Hawai'i Island. These archipelago‐wide sympatric varieties show the weakest neutral genetic divergence of any taxon pair on Hawai'i Island but significant morphological and ecological differentiation consistent with adaptation to new and old lava flows. Cross‐fertility between varieties was high and included heterosis of F₁ hybrids at the seed germination stage, consistent with a substantial genetic load apparent within varieties through low self‐fertility and a lack of self‐pollen discrimination. However, a partial, but significant, barrier was observed in the form of reduced female and male fertility of hybrids, especially backcross hybrids, consistent with the accumulation of genetic incompatibilities between varieties. These results suggest that partial intrinsic postzygotic barriers can arise through disruptive selection acting on large, hybridizing populations of a long‐lived species.     

Incipient radiation within the dominant Hawaiian tree Metrosideros polymorpha

Although trees comprise a primary component of terrestrial species richness, the drivers and temporal scale of divergence in trees remain poorly understood. We examined the landscape-dominant tree, Metrosideros polymorpha, for variation at nine microsatellite loci across 23 populations on young Hawai''i Island, sampling each of the island''s five varieties throughout its full geographic range. For four varieties, principal coordinate analysis revealed strong clustering of populations by variety across the 10 430 km² island, indicating partitioning of the species into multiple evolutionarily significant units. The single island-endemic form, riparian var. newellii, showed especially strong differentiation from other varieties despite occurring in sympatry with other varieties and likely evolved from a bog form on the oldest volcano, Kohala, within the past 500 000 years. Along with comparable riparian forms on other Pacific Islands, var. newellii appears to represent parallel incipient ecological speciation within Metrosideros. Greater genetic distance among the more common varieties on the oldest volcano and an inverse relationship between allelic diversity and substrate age appear consistent with colonization of Hawai''i Island by older, partially diverged varieties followed by increased hybridization among varieties on younger volcanoes. This study demonstrates that broad population-level sampling is required to uncover patterns of diversification within a ubiquitous and long-lived tree species. Hawaiian Metrosideros appears to be a case of incipient radiation in trees and thus should be useful for studies of divergence and the evolution of reproductive isolating barriers at the early stages of speciation.     

Short‐term Dynamics and the Effects of Biotic and Abiotic Factors on Plant Physiognomic Groups in a Hurricane‐impacted Lower Montane Tropical Forest

We assessed the short‐term effects of biotic (density, plant size) and abiotic factors (light), on the dynamics of physiognomically different plant groups (palms, tree ferns, lianas, and trees) in a hurricane‐impacted tropical wet montane forest, John Crow Mountains, Jamaica. All plants ≥2 cm (dbh) found within 45, 25 × 25 m permanent sample plots (2.8125 ha), established according to a randomized block design along an elevation gradient, were tagged and measured (dbh) in 2006 and re‐assessed in 2012 after Hurricane Dean (2007). Hemispheric light was measured in 2007 and 2008. Tree and liana size class distributions changed due to high mortality in the smallest size classes and their densities declined; however, palm and tree fern density remained unchanged. The dynamics of trees were only related to tree fern and liana dynamics (e.g., tree mortality was negatively related to liana recruitment etc.). Although pre‐ and posthurricane light was related to palm density and the density of the other plant groups, respectively, there were no significant changes in light. Tree survivorship increased with increasing dbh while posthurricane light and overall density influenced the growth and survivorship of tree species. Species importance value did not change, suggesting that direct regeneration may be the model of forest recovery following this small‐scale disturbance. Over the short term, tree species showed life history trade‐offs that aid species coexistence after this moderate/low disturbance event. Our study highlights that hurricanes with low impacts can have differential short‐ and possibly long‐term effects on different plant groups.     

Ecosystem development in short‐term postglacial chronosequences: N and P limitation in glacier forelands from Santa Inés Island,Magellan Strait

Glacier foreland moraines provide an ideal model to examine the patterns of ecosystem development and the evolution of nitrogen and phosphorous limitation over successional time. In this paper, we focus on a 400‐year soil chronosequence in the glacier forelands of Santa Inés Island in the Magellan Strait, southern Chile by examining forest development on phosphorus (P)‐poor substrates in a uniquely unpolluted region of the world. Results show a steady increase in tree basal area and a humped trend in tree species richness over four centuries of stand development. The increase in basal area suggests that the late successional tree species were more efficient nutrient users than earlier successional ones. Total contents of carbon (C) and nitrogen (N) in soils increased during the chronosequence, reaching an asymptote in late succession. The net increases in soil C : N, C : P and N : P ratios observed over successional time suggest that nutrient limitation is maximal in 400‐year‐old substrates. Foliar C : N and C : P ratios also increased over time to reach an asymptote in old‐growth stages, following soil stoichiometric relationships; however the foliar N‐to‐P ratio remained constant throughout the chronosequence. Biological N fixation was greater in early postglacial succession, associated with the presence of the symbiotic N‐fixer Gunnera magellanica. Declining trends of δ¹⁵N in surface soils through the 400‐year chronosequence are evidence of decreasing N losses in old‐growth forests. In synthesis, glacier foreland chronosequences at this high South American latitude provide evidence for increasing efficiency of N and P use in the ecosystem, with the replacement of shade‐intolerant pioneers by more efficient, shade‐tolerant tree species. This pattern of ecosystem development produces a constant foliar N : P ratio, regardless of variation in soil N‐to‐P ratio over four centuries.     

Larval habitat for the avian malaria vector Culex quinquefasciatus (Diptera: Culicidae) in altered mid‐elevation mesic‐dry forests in Hawai'i

Effective management of avian malaria (Plasmodium relictum) in Hawai'i's endemic honeycreepers (Drepanidinae) requires the identification and subsequent reduction or treatment of larval habitat for the mosquito vector, Culex quinquefasciatus (Diptera: Culicidae). We conducted ground surveys, treehole surveys, and helicopter aerial surveys from 2001–2003 to identify all potential larval mosquito habitat within two 100+ ha mesic‐dry forest study sites in Hawai'i Volcanoes National Park, Hawai'i; ‘Ainahou Ranch and Mauna Loa Strip Road. At ‘Ainahou Ranch, anthropogenic sites (43%) were more likely to contain mosquitoes than naturally occurring (8%) sites. Larvae of Cx. quinquefasciatus were predominately found in anthropogenic sites while Aedes albopictus larvae occurred less frequently in both anthropogenic sites and naturally‐occurring sites. Additionally, moderate‐size (≈20–22,000 liters) anthropogenic potential larval habitat had >50% probability of mosquito presence compared to larger‐ and smaller‐volume habitat (<50%). Less than 20% of trees surveyed at ‘Ainahou Ranch had treeholes and few mosquito larvae were detected. Aerial surveys at ‘Ainahou Ranch detected 56% (95% CI: 42–68%) of the potential larval habitat identified in ground surveys. At Mauna Loa Strip Road, Cx. quinquefasciatus larvae were only found in the rock holes of small intermittent stream drainages that made up 20% (5 of 25) of the total potential larval habitat. The volume of the potential larval habitat did not influence the probability of mosquito occurrence at Mauna Loa Strip Road. Our results suggest that Cx. quinquefasciatus abundance, and subsequently avian malaria, may be controlled by larval habitat reduction in the mesic‐dry landscapes of Hawai'i where anthropogenic sources predominate.     

A Strategy for Restoration of Montane Forest in Anthropogenic Fern Thickets in the Dominican Republic

Deforested tropical areas are often colonized by competitive ferns that inhibit forest succession. In thickets of such a fern (Dicranopteris pectinata), we investigated methods for initiating restoration of tropical montane forest in the Ébano Verde Scientific Reserve (Dominican Republic). In clearings in the thickets, growth and survivorship of 18 common early‐ and late‐successional woody species were tested, with and without fertilizer (poultry litter). Three years after sowing, life history did not affect survivorship, but early‐successional species grew faster than late‐successional species (height increase 153 ± 103 cm vs. 81 ± 67 cm [mean ± 1 SD]). Inga fagifolia, a late‐successional species, and Alchornea latifolia, an early‐successional species, had 160 ± 62 cm mean height increase, and low mortality rates (<4%). In contrast, four late‐successional species (Cyrilla racemiflora, Myrcia deflexa, Prestoea acuminata var. montana, and Mora abbottii), and one early‐successional species, Ocotea leucoxylon, had approximately 39% mortality and height increase of 43 ± 48 cm. Brunellia comocladifolia had high mortality (55%), but averaged 340 ± 201 cm height increase, and was the only species whose growth was improved by fertilization. Fertilization improved survivorship of only one species, Piper aduncum. After three years, soils in the clearings had low pH and available P and did not differ significantly from soils in thickets. However, based on the growth rates and survivorship of sown woody plants, these soils did not appear to be a barrier for restoration. Although a complementary study demonstrated substantial natural regeneration, active planting should be used to increase plant density and diversity, especially where natural regeneration is poor.     

Forest succession and prey availability influence the strength and scale of terrestrial‐aquatic linkages in a headwater salamander system

1. Trophic linkages between terrestrial and aquatic ecosystems are common and sensitive to disruption. However, there is little information on what causes variation in the strength and spatial scale of these linkages. 2. In the highly aquatic adults of the headwater salamander Gyrinophilus porphyriticus (family Plethodontidae), use of terrestrial prey decreases along a gradient from early‐ to late‐successional riparian forests. To understand the cause of this relationship, we tested the predictions that (i) terrestrial prey abundance is lower in late‐successional forests, and (ii) G. porphyriticus adults cannot move as far from the stream to forage in late‐successional forests, thus limiting access to terrestrial prey. 3. We established 100‐m long study reaches on six headwater streams in the Hubbard Brook Experimental Forest, New Hampshire. Three reaches were in early‐successional forests and three were in late‐successional forests. We conducted pitfall trapping for invertebrate prey in June and July of 2005, with three traps at 0, 2, 5 and 10 m from the stream at each reach. In June, July and August of 2004 and 2005, nighttime salamander surveys were conducted at each reach along ten, 10‐m long by 2.5‐m wide transects perpendicular to the stream. 4. Abundance of terrestrial prey was consistently lower in late‐successional forests, suggesting that consumption of terrestrial prey by G. porphyriticus is affected by prey abundance. Contrary to our prediction, G. porphyriticus adults moved farther from the stream in late‐successional forests, suggesting that habitat conditions in late‐successional forests do not limit movement away from the stream, and that lower abundances of terrestrial prey in these forests may cause salamanders to move farther from streams. 5. Our results provide novel insight on the extent of terrestrial habitat use by G. porphyriticus. More broadly, these results indicate that major habitat gradients, such as forest succession, can affect the strength and scale of terrestrial‐aquatic linkages. Application of this insight to the design of vegetation buffers along headwater streams would have widespread benefits to freshwater ecosystems.     

Coexistence of two congeneric tree species of Lauraceae in a secondary warm‐temperate forest on Miyajima Island,south‐western Japan

We analyzed the spatial distributions of two congeneric tree species, Neolistea aciculata and Neolistea sericea (Lauraceae), in a warm‐temperate forest on Miyajima Island, south‐western Japan. Both species were mainly found in valley sites on the island. Hence, these species shared the same topographic habitat niche. However, we found a clear difference between the spatial distributions of the two species in relation to the light environment. Neolistea aciculata was predominantly found in stands with low light, such as beneath the canopy of dense evergreen broadleaved forest. In contrast, N. sericea was predominantly associated with ample light, such as in secondary Pinus densiflora forest. In stands with moderate light conditions, both species were found. This habitat niche segregation in relation to light conditions presumably allows the coexistence of these two species in the predominantly successional forest on Miyajima Island.     

Butterfly Community Composition Across a Successional Gradient in a Human‐disturbed Afro‐tropical Rain Forest

Knowledge of the recovery of insect communities after forest disturbance in tropical Africa is very limited. Here, fruit‐feeding butterflies in a tropical rain forest at Kibale National Park, Uganda, were used as a model system to uncover how, and how fast, insect communities recover after forest disturbance. We trapped butterflies monthly along a successional gradient for one year. Traps were placed in intact primary forest compartments, heavily logged forest compartments with and without arboricide treatment approximately 43 years ago, and in conifer‐clearcut compartments, ranging from 9 to 19 years of age. The species richness, total abundance, diversity, dominance, and similarity of the community composition of butterflies in the eight compartments were compared with uni‐ and multivariate statistics. A total of 16,728 individuals representing 88 species were trapped during the study. Butterfly species richness, abundance, and diversity did not show an increasing trend along the successional gradient but species richness and abundance peaked at intermediate stages. There was monthly variation in species richness, abundance, diversity and composition. Butterfly community structure differed significantly among the eight successional stages and only a marginal directional change along the successional gradient emerged. The greatest number of indicator species and intact forest interior specialists were found in one of the primary forests. Our results show that forest disturbance has a long‐term impact on the recovery of butterfly species composition, emphasizing the value of intact primary forests for butterfly conservation.     

Testing the Performance of Fourteen Native Tropical Tree Species in Two Abandoned Pastures of the Lacandon Rainforest Region of Chiapas,Mexico

The rainforest of Mexico has been degraded and severely fragmented, and urgently require restoration. However, the practice of restoration has been limited by the lack of species‐specific data on survival and growth responses to local environmental variation. This study explores the differential performance of 14 wet tropical early‐, mid‐ or late‐successional tree species that were grown in two abandoned pastures with contrasting land‐use histories. After 18 months, seedling survival and growth of at least 7 of the 14 tree species studied were significantly higher in the site with a much longer history of land use (site 2). Saplings of the three early‐successional species showed exceptional growth rates. However, differences in performance were noted in relation to the differential soil properties between the experimental sites. Mid‐successional species generally showed slow growth rates but high seedling survival, whereas late‐successional species exhibited poor seedling survival at both the study sites. Stepwise linear regressions revealed that the species integrated response index combining survivorship and growth measurements, was influenced mostly by differences in soil pH between the two abandoned pastures. Our results suggest that local environmental variation among abandoned pastures of contrasting land‐use histories influences sapling survival and growth. Furthermore, the similarity of responses among species with the same successional status allowed us to make some preliminary site and species‐specific silvicultural recommendations. Future field experiments should extend the number of species and the range of environmental conditions to identify site “generalists” or more narrowly adapted species, that we would call “sensitive.”     

Steady and dynamic photosynthetic responses of seedlings from contrasting successional groups under low‐light growth conditions

To test the hypothesis that leaf‐level photosynthetic‐related traits might confer late successionals a competitive advantage over early successionals in low‐light growth conditions, steady photosynthetic assimilation‐ and dynamic photosynthetic induction‐related traits were examined in low‐light‐grown seedlings with contrasting successional status. Compared with the early successionals, late successionals as a group significantly exhibited lower leaf gas exchange rates. While late successionals required a longer time to respond to simulated sunflecks, they had lower rates of induction losses after sunflecks. Such photosynthetic induction traits allowed late successionals to more effectively utilize subsequent sunflecks. It was observed that plants with lower gas exchange rates responded more slowly to simulated sunfelcks, but they had lower rates of induction losses after sunflecks. In addition, the rate of response to sunflecks was positively correlated with the rate of induction loss after sunflecks across the successional status of species. A principal components analysis (PCA) demonstrated that early and late successionals were separated along the first axis of the PCA, and that early successionals were grouped on the right and were associated with higher gas exchange rates, fast responses to sunflecks, and rapid rates of induction loss after sunflecks; late successionals held an opposite pattern. Overall, our results suggest that smaller respiratory carbon losses and lower metabolic costs give late successionals a competitive advantage in low‐light growth conditions, that late successionals have an advantage over early successionals in utilizing sunflecks, and thus that the successional status of species are mainly associated with the leaf‐level photosynthetic‐related traits.     

Leaf damage and functional traits along a successional gradient in Brazilian tropical dry forests

Herbivory has significant impacts on individual plants and plant communities, both at ecological and evolutionary time scales. In this context, this study aims to evaluate herbivore damage and its relationship with leaf chemical and structural traits, nutritional status, and forest structural complexity along a successional gradient. We predicted that trees in early successional stages support conservative traits related to drought tolerance (high specific leaf mass and phenolics), whereas trees in light-limited, late successional stages tend to enhance light acquisition strategies (high nitrogen content). We sampled 261 trees from 26 species in 15 plots (50 × 20 m; five per successional stage). From each tree, twenty leaves were collected for leaf trait measures. Phenolic content increased whereas specific leaf mass and nitrogen content decreased from early to late stages. However, leaf damage did not differ among successional stages. Our results partially corroborate the hypothesis that early successional plants in tropical dry forests exhibit leaf traits involved in the conservative use of water. The unexpected decrease in nitrogen content along the chronosequence is likely related to the fact that thinner leaves with low specific leaf mass could have less nitrogen-containing mesophyll per unit area. Mechanisms affecting herbivory intensity varied across scales: at the species level, leaf damage was negatively correlated with tannin concentration and specific leaf mass; at the plot level, leaf damage was positively affected by forest structural complexity. Herbivory patterns in tropical forests are difficult to detect because abiotic factors and multiple top-down and bottom-up forces directly and indirectly affect herbivores.     

Shifts in the height‐related competitiveness of tree species following recent climate warming and implications for tree community composition: the case of common beech and sessile oak as predominant broadleaved species in Europe

Height growth is a trait that contributes to tree species fitness. How height growth responds to environmental changes may therefore provide indications on species ability to compete and maintain, and on changes in tree community composition. Common beech Fagus sylvatica and sessile oak Quercus petraea are the predominant late‐successional broadleaved species in Europe, and they differ in their shade‐tolerance. On common beech (a shade tolerant species), recent observations across Europe have shown a growth decline during recent climate warming. Because sessile oak is a warmth‐ and light‐demanding species, we therefore hypothesised that it may gain in competitiveness relative to common beech. We conducted analyses of historical height growth in several regions spanning the distributional range of the two species across a temperate‐continental gradient in France. Common beech and sessile oak were sampled in two and four regions, respectively, and were compared in two neighbouring regions. We documented the climatic and nutritional conditions of regional samples. Height growth of 408 trees of various ages was reconstituted from stem analyses. We estimated 20th‐century regional chronologies of height growth using a statistical modelling approach that filtered out the effects of ontogeny and site fertility. In regions where both species were sampled, modelled height trajectories were compared at different periods over the 20th century. Growth chronologies revealed 1) long‐term growth rate increases of a magnitude of 50–100% over 100 years in both species, more acute in the continental domain, 2) recurrent historical inversions in growth fluctuations between species, 3) a recent divergence, with growth decline in common beech versus a dramatic growth increase in sessile oak, more acute in colder regions. The analysis of height trajectories indicated a recent reduction in common beech competitiveness relative to sessile oak. In the face of future climate warming, we conclude that increased prevalence of beech–oak mixtures may arise.     

Canopy manipulation as a tool for restoring mature forest conifers under an early‐successional angiosperm canopy

Low‐light environments in early‐successional forests that have established after abandonment of farming often restrict the establishment of later successional species resulting in an arrested succession. This 6‐year study tested the potential of different canopy manipulations to facilitate the establishment of a light‐demanding canopy tree species, tōtara (Podocarpus totara), within a regenerating kānuka (Kunzea robusta) stand. Results highlighted the effectiveness of artificial gaps over other methods (ring‐barking and edge‐planting) in accelerating the growth of planted tōtara. Seedlings under gaps grew consistently taller and faster over time indicative of an improved understorey light environment. Ring‐barking did not have a significant effect on tōtara growth because only a portion of the treated trees died, and after 6 years dead trees remained standing with intact branches resulting in insignificant increases in light transmission. At the forest edge sites, tōtara growth was highly variable. Although some seedlings grew as tall as in the gaps, others did not. Survival was also lower in the edge sites than in other treatments, which was likely due to enhanced herbivory from ungulates which impacted some plants at these sites. Gap creation is likely to be an important tool for restoring late‐successional canopy species in regenerating stands both through providing ideal sites for the growth of light‐demanding species such as tōtara and through natural establishment of other future canopy trees into the gaps.     

Understorey vegetation development in North Finnish Picea forests after disturbance: re‐analysis of Sirén's data

Abstract. Sirén (1955) studied understorey species composition, tree stand properties and humus‐layer thickness in 64 unlogged forest stands on topographically and pedologically comparable sites. The stands were of even age (6 – 300 yr), stocked with the first or second tree generation after wildfire. The view of Sirén and several authors after him, that the vegetation of old‐growth boreal Picea forests is homogeneous on a broad scale, was examined by applying, in parallel, the partial variants of two ordination methods (DCA and PCA) to Sirén's vegetation data. Two main vegetation gradients were found: a major gradient running from recently burnt plots with prominence of pioneer species to plots with stand age > 100 yr, a well stocked tree layer and a thick humus layer, dominance of feather‐mosses and ample occurrence of shade‐tolerant as well as light‐preferring vascular plant species, and a second gradient along which first‐ and second‐generation plots segregate. The more prominent element of Betula trees in first‐ than in second‐generation stands < 100 yr contributed to the latter. A minor third gradient related to humus‐layer thickness was recovered by partial DCA only. The main vegetation gradient reappeared in separate ordinations of data from 47 mature forest stands (> 100 yr), but without being correlated with forest age. Variation among mature‐forest stands in the importance of pioneer species is considered mainly to be brought about by fine‐scale disturbance processes such as tree uprooting. Increasing importance of factors operating on within‐stand scales [development of a varied gap structure and stronger gradients in tree influence (radiation at ground level), soil moisture, soil depth and nutrient availability] with time is also reflected in the second and third mature‐forest ordination axes. Possible implications of the results for conservation of biological diversity and monitoring of changes in boreal forests are discussed.     

Colonization dynamics and facilitative impacts of a nitrogen‐fixing shrub in primary succession

Abstract. Whether nitrogen‐fixing plants facilitate or inhibit species change in primary succession is best resolved by examining their impacts throughout the plant's entire life cycle from arrival to senescence. We experimentally examined two aspects of the successional impacts of a nitrogen‐fixing shrub, Coriaria arborea, on Mt. Tarawera, a volcano in New Zealand: factors limiting Coriaria colonization and impacts of Coriaria‐induced soil changes on a later successional tree, Griselinia littoralis. Coriaria germination was promoted by artificial wind protection and by the presence of heath shrubs. Transplanted Coriaria seedlings survived only if nodulated with Frankia, and the addition of Coriaria‐enriched soils slowed Coriaria seedling growth and did not improve seedling survival. This explained why Coriaria seedlings were found mostly in protected habitats away from adult Coriaria, and suggested that Coriaria thickets are not self‐replacing. Coriaria increased soil fertility by developing a 4 cm thick organic soil horizon that was richer in nitrogen (tenfold) and phosphorus (threefold) than pre‐Coriaria stages. These soil changes resulted in three‐ to sixfold increases in growth of Griselinia when it was grown in Coriaria‐enriched soils in a glasshouse. Coriaria's net effect on primary succession is facilitative, but the establishment of Coriaria is itself facilitated by the amelioration of the physical habitat by earlier colonists, suggesting facilitation is important throughout the life cycle of Coriaria. Sequential facilitative events determine the order of species replacements in this study but inhibition, linked to the developmental stages of Coriaria, may determine the rate of species change.     

Shoot life span in relation to successional status in deciduous broad-leaved tree species in a temperate forest

In trees, leaf life span is closely related to successional status. Although leaves are attached to shoots, shoot life span has been insufficiently studied in the context of ecological systems. Interspecific variation in shoot survivorship was investigated over 27 months in 15 temperate hardwood tree species. Relationships between shoot architecture and shoot survival were also investigated. Shoot life span was shortest in early successional species, and longest in late successional species, in each of the families Betulaceae and Fagaceae. In Salicaceae, all of which were early successional species, shoot life span was longer in mountainous than in riparian species. Early successional or riparian species distributed longer shoots densely, even in proximal positions on mother shoots, resulting in mutual shading and consequent early and massive shoot shedding. By contrast, late successional or mountainous species concentrated shoots in distal positions, allowing shoots to receive equally favorable light, resulting in a longer life span. These results reveal close relationships between shoot life span and environmental resource availability or successional status and suggest a causal relationship between shoot shedding and shoot architecture.     

Increased forest carbon storage with increased atmospheric CO2 despite nitrogen limitation: a game‐theoretic allocation model for trees in competition for nitrogen and light

Changes in resource availability often cause competitively driven changes in tree allocation to foliage, wood, and fine roots, either via plastic changes within individuals or through turnover of individuals with differing strategies. Here, we investigate how optimally competitive tree allocation should change in response to elevated atmospheric CO₂ along a gradient of nitrogen and light availability, together with how those changes should affect carbon storage in living biomass. We present a physiologically‐based forest model that includes the primary functions of wood and nitrogen. From a tree's perspective, wood is an offensive and defensive weapon used against neighbors in competition for light. From a biogeochemical perspective, wood is the primary living reservoir of stored carbon. Nitrogen constitutes a tree's photosynthetic machinery and the support systems for that machinery, and its limited availability thus reduces a tree's ability to fix carbon. This model has been previously successful in predicting allocation to foliage, wood, and fine roots along natural productivity gradients. Using game theory, we solve the model for competitively optimal foliage, wood, and fine root allocation strategies for trees in competition for nitrogen and light as a function of CO₂ and nitrogen mineralization rate. Instead of down‐regulating under nitrogen limitation, carbon storage under elevated CO₂ relative to carbon storage at ambient CO₂ is approximately independent of the nitrogen mineralization rate. This surprising prediction is a consequence of both increased competition for nitrogen driving increased fine root biomass and increased competition for light driving increased allocation to wood under elevated CO₂.     

Regional‐scale directional changes in abundance of tree species along a temperature gradient in Japan

Climate changes are assumed to shift the ranges of tree species and forest biomes. Such range shifts result from changes in abundances of tree species or functional types. Owing to global warming, the abundance of a tree species or functional type is expected to increase near the colder edge of its range and decrease near the warmer edge. This study examined directional changes in abundance and demographic parameters of forest trees along a temperature gradient, as well as a successional gradient, in Japan. Changes in the relative abundance of each of four functional types (evergreen broad‐leaved, deciduous broad‐leaved, evergreen temperate conifer, and evergreen boreal conifer) and the demography of each species (recruitment rate, mortality, and population growth rate) were analyzed in 39 permanent forest plots across the Japanese archipelago. Directional changes in the relative abundance of functional types were detected along the temperature gradient. Relative abundance of evergreen broad‐leaved trees increased near their colder range boundaries, especially in secondary forests, coinciding with the decrease in deciduous broad‐leaved trees. Similarly, relative abundance of deciduous broad‐leaved trees increased near their colder range boundaries, coinciding with the decrease in boreal conifers. These functional‐type‐level changes were mainly due to higher recruitment rates and partly to the lower mortality of individual species at colder sites. This is the first report to show that tree species abundances in temperate forests are changing directionally along a temperature gradient, which might be due to current or past climate changes as well as recovery from past disturbances.