Population structures and individual performances of <Emphasis Type="Italic">Trillium grandiflorum</Emphasis> in hedgerow and forest habitats

In agricultural landscapes, linear habitats, such as hedgerows at field margins increase structural connectivity among forest patches, potentially providing dispersal corridors for forest herbs. The spatial structure of linear habitats, however, also results in edge effects and perturbations that can influence the individual and population performance of forest plants. This study compares the stage structure and components of growth and reproduction of 14 Trillium grandiflorum populations in hedgerows and forests. Hedgerow Trillium tended to grow faster and, when mature, produced more flowers and more ovules per flowers than forest Trillium, a pattern possibly associated to differences in nutrients and light availability between the two habitats. Seed production and germination rate, however, did not differ between hedgerows and forests. At the population level, seedlings and juveniles were proportionally less abundant in hedgerows than in forests. Although well-established plants can thrive in hedgerows, reduced recruitment may eventually limit the capacity to establish new populations and therefore hamper migration along hedgerow-corridors. Considering the strategies by which plants persist in linear habitats becomes particularly relevant at a time when species are expected to be much in need of dispersal corridors because of climatic stress.     

Effects of chronic herbivory and historic land use on population structure of a forest perennial,Trillium catesbaei

Question: How have long-term herbivory and past land use impacted the population structure of Trillium catesbaei, a long-lived rhizomatous herb? Location: Western Great Smoky Mountains National Park, Tennessee, USA. Methods: We examined T. catesbaei populations at three sites: (1) Cades Cove (CC), an area of intensive historic land use that has been maintained as open fields and woodlots with a history of chronic deer herbivory, (2) Whiteoak Sink (WOS), a reference area with similar land-use history, geology, and soils that has succeeded to closed-canopy forest with relatively low levels of deer herbivory, and (3) Leadbetter Ridge (LBR), an area of primary forest that has never received significant anthropogenic disturbance. Trillium catesbaei is the most common Trillium species at the three study sites, but smaller in stature, shorter lived, and more of a habitat generalist. Results: Chronic herbivory in CC has created a highly-truncated age structure with no plants older than 9 years, while plant ages at the other sites were more evenly distributed. Compared to WOS, plants in CC were younger at a given height and more likely to flower when younger. Across all life stages, populations at CC contained 68 × fewer plants than WOS. The age structures of WOS and LBR were similar. Compared to published age estimates for other Trillium species, our results suggest that T. catesbaei is relatively short lived within the genus. Conclusions: Chronic herbivory had pronounced effects on the population structure of a perennial herb. Other long-lived herbaceous species may exhibit similar truncated age structures and flowering by younger and smaller plants. Habitat generalist species within a genus, such as T. catesbaei, that are able to reproduce more quickly may persist longer under chronic herbivory. However, chronic herbivory has likely caused the loss of herbaceous species from CC and may eventually cause the local extirpation of T. catesbaei populations.     

THE CYTO-ECOLOGY OF FOUR SPECIES OF TRILLIUM FROM WESTERN NORTH CAROLINA

The major center of variability in the genus Trillium occurs in the southern Appalachian mountains of the eastern United States. The karyotypic variation existing within T. cuneatum Raf., T. grandiflorum (Michx.) Salisb., T. erectum L., and T. gleasoni Fern., the four most abundant Trillium species in western North Carolina, was analyzed and compared with the variation in gross morphology and the ecological associations of the respective species. The number of types of each kind of chromosome (A-E), as exhibited following “nucleic acid starvation” by cold treatment, varied widely from species to species. Chromosome types varied within each species and within populations in the numbers, sizes, and positions of euchromatic and heterochromatic segments. No 10-chromosome karyotype and only one 5-chromosome complement was found to be duplicated in any two plants which belonged to different species. Trillium cuneatum with eight chromosome types was morphologically stable and occurred in a limited geographic area. Trillium gleasoni, characteristically adapted to elevations near 2,000 feet, contained 32 chromosome types and exhibited a high morphological variability. Trillium erectum (25 types) and T. grandiflorum (23 types) were also intermediate in morphological variability. These two species had the widest geographical range. Karyotypes were analyzed from pure stands of each of the four species and for each species from mixed stands of two or more of the species growing in close association. These Trillium species are maintaining a high level of karyotypic and gross morphological variability within comparatively limited geographical areas. None of the observed karyotypes in any species is likely to have arisen by recent interspecific hybridization. The composite karyotype of each species could be used as an aid to classification in the genus Trillium.     

A field study of seed germination in the endangered Trillium reliquum Freeman (Trilliaceae)

Previous studies examining the seeds of most Trillium species have reported double dormancy, a type of seed dormancy where two cold periods and one warm period are needed for complete germination. In the present paper, we describe a field study examining the federally endangered Trillium reliquum Freeman (Trilliaceae) in which moderate to high numbers of seeds germinated after one winter following seed production. Sixteen baskets with seeds were placed in four T. reliquum populations (four baskets in each population) in Georgia, USA, in June 2005. In spring 2006, all seed baskets contained seedlings. Germination percentages ranged from 33.3 to 83.3% across sites with a mean of 56.9 ± 3.9%. Trillium reliquum had higher germination percentages compared with other field‐based germination studies with other Trillium species. Our findings will inform future demographic studies of T. reliquum and suggest that double dormancy in seeds may not be as widespread as previously reported within the genus Trillium.     

Forest age correlates with fine-scale spatial structure of Matsutake mycorrhizas

Examining the fine-scale spatial structure of fungal populations can tell us much about how individual species reproduce and disperse throughout natural landscapes. Here we study the fine-scale genetic structure of Tricholoma matsutake, a prized edible and medicinal mushroom, by systematic sampling of mycorrhizas within fairy rings in 50-y-old and old-growth forests in two villages. Using single nucleotide polymorphism DNA markers we show that mycorrhizas in both forest age classes in both villages showed high levels of genotypic diversity, consistent with a reproductive life history predominated by outcrossing via basidiospore dispersal. Both the percentage of polymorphic loci within fairy rings, as well as genotype diversity were higher in old-growth compared to 50-y-old forests. Fifty-year-old forests showed significant spatial autocorrelation between pairs of mycorrhizas up to 42 m, and a pattern consistent isolation-by-distance structure. Spatial patterns in old-growth forests were random. Furthermore, AMOVA analysis indicates that 11 % of molecular variance in 50-y-old forests is partitioned between villages, whereas no significant variance is partitioned between villages in old-growth forests. We conclude that populations of T. matsutake in 50-y-old forests are the result of a founder effect maintained by local inoculation sources. This pattern attenuates as forests age and accumulate inocula from more distance sources. We speculate on how genetic mosaicism within T. matsutake fairy rings may structure populations within a chronosequence. Finally, we discuss how population spatial dynamics and dispersal strategy in T. matsutake contrast with other ectomycorrhizal species.     

Presidential Address—Some Reflections On The Nature Of Species

Background: Variation in the distribution and abundance of woody plants as consequence of disturbances such as fire may be explained by lineage age.

Aims: We tested whether lowland tropical tree lineages that colonise secondary forests are more late-diverging than clades from old-growth forests, and whether tree phylogenetic beta diversity from old-growth to secondary forests is higher in burned than non-burned secondary forests.

Methods: We sampled tree communities in old-growth forests and in secondary forests with distinct disturbance histories (burned and unburned). We calculated mean family age in each plot, and tested for differences among forest types using ANOVA. A phylogenetic fuzzy-weighting procedure was employed to generate a matrix describing the abundance of tree clades per plot, which was then analysed using a principal coordinate analysis.

Results: Most clades found in old-growth forests were underrepresented in secondary forests, which have been heavily colonised by a single species from a young lineage that is not found in old-growth forests. Phylogenetic beta diversity was higher between unburned secondary forests and old-growth forests than between burned secondary forests and old-growth forests.

Conclusions: The capacity of Neotropical trees to colonise secondary forests and persist after fire disturbance may be related to the age of distinct lineages.     

Spatial patterns and interspecific associations of dominant tree species in two old-growth karst forests,SW China

Spatial patterns and interspecific associations of plant species in forests are important for revealing how species interact with each other and with the environment, and hence have important implications for optimal forest management and restoration in degraded forest ecosystems. In this paper, the O-ring statistics were used to characterize the spatial patterns and interspecific associations of eight dominant tree species in two 1-ha old-growth karst forest plots in Maolan National Natural Reserve, southwestern China. We found that most of the eight dominant tree species in two forests were continuously regenerating populations. Six species (Platycarya longipes, Acer wangchii, Clausena dunniana, Castanopsis carlesii var. spinulosa, Distylium myricoides, and Rhododendron latoucheae) exhibited significant aggregations at the majority of scales while others (Celtis biondii and Cyclobalanopsis myrsinaefolia) showed a random distribution pattern at most scales. Negative association was a dominant pattern for most species pairs in the two plots, while positive associations were found at most scales for only two species pairs (Platycarya–Clausena and Castanopsis–Rhododendron). Results also indicated that the two main factors of habitat complexity and heterogeneity—the elevation and rock-bareness rate—play important roles in determining spatial distribution patterns and interspecific associations of tree species in karst forests of Maolan. Thus, the observed spatial patterns among the eight tree species are influenced by habitat heterogeneity in the context of karst topographical variations. The partitioning of habitat niches contributes to the promoting species coexistence in species-rich karst forests. The differences of species features in spatial patterns and associations should be paid more attention when planning forest management and developing restoration strategies.     

Genetic Diversity and Spatial Genetic Structure of an Epiphytic Bromeliad in Costa Rican Montane Secondary Forest Patches

Information on genetic variation and its distribution in tropical plant populations relies mainly on studies of ground‐rooted species, while genetic information of epiphytic plants is still limited. Particularly, the effect of forest successional condition on genetic diversity and structure of epiphytes is scanty in the literature. We evaluated the genetic variation and spatial genetic structure of the epiphytic bromeliad Guzmania monostachia (Bromeliaceae, Tillandsioideae) in montane secondary forest patches in Costa Rica. The sampling design included plants on the same trees (i.e., populations), populations within forest patches and patches within secondary forest at two different successional stages (early vs. mid‐succession). Six microsatellites revealed low levels of population genetic variation (A = 2.06, A_E = 1.61, H_E = 0.348), a marked deficiency of heterozygotes (H_O = 0.031) and high inbreeding (f = 0.908). Genetic differentiation was negligible among populations within the same forest patch, but moderate (G_ST = 0.123 ± 0.043) among forest patches. Genetic relatedness between individuals was significantly higher for plants located within the same forest patch and separated by <60 m and decreased as distance between plants increased, becoming significantly negative at distances >400 m. An analysis of molecular variance (AMOVA) showed significant genetic variation between forest patches, but non‐significant variation between successional stages. The selfing breeding system and limited seed dispersal capabilities in G. monostachia could explain the observed levels and partitioning of genetic diversity at this geographic scale. However, these results also suggest that forest fragmentation is likely to influence the degree of local genetic structuring of epiphytic plants by limiting gene flow.     

Investigating above-ground biomass in old-growth and secondary montane forests of the Cameroon Highlands

Tropical montane forests can store and sequester substantial amounts of carbon in above-ground biomass (AGB), but variations in this storage related to location or degradation have not been quantified in the Cameroon Highlands. We established 25 permanent plots (20 m × 40 m) and sampled all trees ≥10 cm diameter following standard RAINFOR protocols. We estimated AGB and investigated variations related to taxonomic and structural forest attributes, including the height–diameter allometry in five forest types (four old-growth dominated by different species and one secondary forest). Secondary forests had significantly lower AGB than old-growth forests (49.4 ± 2.5 vs. >476.3 ± 168.7 Mg/ha, respectively), mostly related to lower basal area and tree height. Significant differences in species composition but not in forest structure or AGB were found between the four types of old-growth forests studied, located at different altitudes and mountains. We discuss the importance of these montane forests for carbon storage and, considering their high diversity and current threats, their potential for carbon finance mechanisms related to both avoided deforestation and forest restoration.     

Structural heterogeneity and tree spatial patterns in an old-growth deciduous lowland forest in Cantabria, northern Spain

Old-growth deciduous forests in western Europe, for the most part, consist of small tracts that often may be atypical due to human disturbance, poor soil productivity or inaccessibility. In addition, very little information on tree age distributions, structural heterogeneity and tree spatial patterns appears to be available for west-European forests. Characterization of the structural features of tree populations in these old-growth stands can provide the basis to design conservation plans and also inform on how present forests might look in the absence of human interference. Four old-growth stands in a deciduous forest in the Cantabrian lowlands, northern Spain, were surveyed to determine forest structure and spatial patterns. Live and dead trees were identified, measured and mapped, and live trees were cored for age estimation. Structural heterogeneity was analyzed by means of the spatial autocorrelation of tree diameter, height and age, and the uni- and bivariate spatial patterns of trees were analyzed. The dominant species, Fagus sylvatica and Quercus robur, showed reverse-J shaped size distributions but discontinuous age distributions, with maximum ages of 255–270 yr. Tree ages suggested that the forest was largely modified by past changes in forest-use, especially by temporal variation in grazing intensity. Spatial autocorrelation revealed that former parkland stands were heterogeneous with respect to tree height only, while high forest stands were composed of patches of even-aged and even-sized trees. Young trees were clumped at varying distances and establishment occurred preferentially in canopy gaps, except for Ilex aquifolium that mainly occurred beneath mature Quercus trees. Surviving trees became less intensely clumped in the dominant species, and more strongly clumped in understorey ones, which may have been due to the effects of intraspecific competition and of canopy trees on tree survival, respectively. The spatial associations between species varied within the forest, probably as a consequence of specific establishment preferences and competitive interactions.     

Spatial patterns and interspecific associations among trees at different stand development stages in the natural secondary forests on the Loess Plateau,China

Quercus wutaishansea populations on the Loess Plateau are currently becoming more dominant in natural secondary forests, whereas Pinus tabulaeformis is declining. In the present paper, the diameter class (instead of age) was used to classify the different growth stages as juvenile, subadult, or adult, and the univariate function g(r) was used to analyze the dynamic changes in spatial patterns and interspecific associations in three 1‐ha tree permanent plots on the Loess Plateau, NW China. Our results suggested that the niche breadth changed with the development stage. The diameter distribution curve was consistent with the inverted “J” type, indicating that natural regeneration was common in all three plots. There was a close relationship between the spatial pattern and scale, which showed significant aggregation at small distances, and became more random as distance increased, but in the Pinus + Quercus mixed forests, the whole species were aggregated at distances up to 50 m. The degree of spatial clumping decreased from juvenile to subadult and from subadult to adult. The spatial pattern also differed at different growth stages, likely due to strong intraspecific competition. Associations among different growth stages were positively correlated at small scales. Our study is important to the understanding of the development of the Q. wutaishansea forests; thus, the spatial dynamic change features should be received greater attention when planning forest management and developing restoration strategies on the Loess Plateau.     

Effects of Sika Deer (Cervus nippon) and Dwarf Bamboo (Sasa senanensis) on Trillium Populations in Akan National Park,Eastern Hokkaido,Japan

The competition between Trillium spp. and other species could modify the utility of Trillium spp. as indicators of the impact of deer on forest vegetation. To evaluate whether Trillium spp. (T. camschatcense and T. tschonoskii) are appropriate indicators of the impact of sika deer (Cervus nippon) in the presence of dwarf bamboo (Sasa senanensis), which competes with Trillium populations, we examined the size class of large‐growth‐stage Trillium plants, the frequency of the presence of deer pellets and the coverage of S. senanensis in deer‐excluded sites (where deer have been eliminated since 1995) and unfenced control sites in Akan National Park, eastern Hokkaido, Japan. The result of a cumulative link mixed model (CLMM) suggests that size class at deer‐excluded sites is influenced by competitive understory species, even without deer grazing. The CLMM result for unfenced control sites suggests that C. nippon negatively influences the size class of Trillium populations; however, the negative effect weakened with increasing S. senanensis coverage. Thus, S. senanensis can function as an unpalatable neighbor that makes it difficult for C. nippon to detect Trillium plants. We conclude that the size class structure of Trillium spp. can be a useful index of C. nippon usage under sparse or medium S. senanensis understories; however, it is difficult to use these indices under dense S. senanensis understories because of the weakened negative effect of C. nippon grazing.     

广东南岭天然常绿阔叶林林下光环境对林下幼树功能性状的影响

光环境与幼树功能性状的关系对天然林的更新与演替具有重要的生态学意义。以广东南岭区域天然常绿阔叶林下不同林龄(幼龄林,中龄林,老龄林)的森林群落为研究对象,通过监测冠层结构、林下光照数据和林下幼树功能性状等指标,研究林龄梯度下其冠层结构与林下光环境之间的关系,以及林下幼树功能性状对光环境的响应。结果表明:(1)中龄林叶面积指数显著高于幼龄林和老龄林(P0.05),随着林龄的增长,林冠开度和透光率逐渐下降,林龄梯度下透光率、R/FR(红光/远红光比值)、Bw/Rw(宽带蓝光/宽带红光比值)差异极显著(P0.001);(2)天然常绿阔叶林中透光率与光质之间极显著相关(P0.001),R/FR随着透光率的增加而增加,Bw/Rw随着透光率的增加而减少。(3)林下幼树功能性状在光环境之间差异显著(P0.05),老龄林林下幼树叶片氮含量显著高于幼龄林,而叶片重叠率显著低于幼龄林;(4)在本试验地中,R/FR和Bw/Rw的变化对林下幼树的高径比和光合作用并无显著影响,光强对同种植物不同光环境下最大净光合速率的影响较大。总体而言,林龄梯度冠层结构和光环境的差异能在一定程度上解释幼树功能性状的差异,这将有助于我们理解光环境对林下幼树更新的影响机制,同时为天然植被恢复和森林经营提供指导。     

High litterfall in old-growth and secondary upper montane forest of Costa Rica

Tropical upper montane forests usually comprise trees of small stature with a relatively low aboveground productivity. In contrast to this rule, in the Cordillera de Talamanca (Costa Rica), tall trees (>35 m in height and more than 60 cm in diameter) are characteristic for the upper montane old-growth oak forests which are growing at an altitude of almost 3,000 m close to the alpine timberline. For these exceptional forests, productivity data are not yet available. In this study, we analyzed litterfall and its components (tree leaves, litter of epiphytic vascular and non-vascular plants, mistletoes, twigs and other canopy debris) in three forest stands belonging to different successional stages and related seasonal changes in litterfall to micrometeorological variables. The studied stands were early-successional forest (10–15-year-old), mid-successional forest (40-year-old), and old-growth forest. The stands are dominated by Quercus copeyensis and are located at 2,900-m altitude. Total litterfall was highest in the mid-successional forest (1,720 g m⁻² y⁻¹), and reached 1,288 g m⁻² y⁻¹ in the old-growth forest and 934 g m⁻² y⁻¹ in the early-successional forest. Litter mass was dominated by leaves in all stages (56–84% of total litterfall). In the old-growth forest, however, twigs and small canopy debris particles (33%), epiphytes (6%), and mistletoes (5%) also contributed substantially to litter mass. Leaf litterfall showed a clear seasonal pattern with a negative correlation to monthly precipitation and highest values in the dry season (January–April). However, the strongest correlation existed with minimum air temperature (negative), probably because temperatures already dropped at the end of the rainy season, when precipitation had not yet declined and leaf shedding already increased. In contrast, litterfall of epiphyte mass, and twigs and other debris was mostly dependent on occasional strong winds. We conclude that the upper montane oak forests of the Cordillera de Talamanca are exceptional with respect to the large tree size and the relatively high productivity as indicated by litterfall. Litter mass was especially high in the mid-successional and old-growth forests, where the observed annual totals are among the highest recorded for tropical forests so far.     

Seasonally flooded,and terra firme in northern Congo: Insights on their structure,diversity and biomass

In the Congo basin, considerable uncertainty remains about the amount and spatial variation of carbon stocks. We studied two types of seasonally flooded forests (dominated by Guibourtia demeusei and Lophira alata) and nearby terra firme forests in northern Congo. We sampled 1.25 ha per forest type and a total of 1,400 trees ≥5 cm diameter. AGB ranged from 207–343 Mg/ha, with no significant differences between forest types. Few significant differences were observed in vegetation structure or tree diversity between forest types. Species richness and stem density of small trees were lower, and dominance was higher in Guibourtia plots, which are subject to greater flooding than Lophira plots. Guibourtia was absent from smaller diameter class in Guibourtia forests; and Uapaca spp. were more abundant in terra firme than in seasonally flooded plots. We show that both types of seasonally flooded forests store important quantities of AGB and should also be considered in forest conservation programmes. We recommend more research on seasonally flooded forests, on larger geographical extent, which assesses flood depth and duration, and measures tree height in the field, as we took a conservative approach to AGB estimates, and AGB could be even greater than we report here.     

Structure and spatial patterns of trees in old-growth northern hardwood and mixed forests of northern Maine

Stand structure including spatial patterns was studied in northern hardwood and mixed forest types in the 2000-ha old-growth Big Reed Forest Reserve in northern Maine using complete stem mapping, dendrochronology, and spatial analyses on 0.5 plots. The inclusion of saplings, dead wood, age distributions, spatial pattern, and interactions provided some idea of underlying processes and temporal change. Structural characteristics were most comparable to spruce-northern hardwood forests of northern New England and New York, and most characteristics matched expected patterns for old-growth forests of the region.Results indicated smaller maximum-tree sizes, lower basal areas (26–34 ) and downed-wood volumes (29–64), higher densities (475–649), but similar species longevities compared to other mesic old-growth forests further south and in the Lake States. The stands were dominated by very shade-tolerant tree species, including Fagus grandifolia Ehrh., Acer saccharum Marsh.,Picea rubens Sarg. and Abies balsamea(L.) Mill, with each species found in many crown positions and age and size classes. The sapling layer was dominated by Fagus grandifolia followed by Picea rubens. Most species had reverse-J shaped diameter distributions, but age distributions were indicative of synchronous, episodic recruitment. In most plots, Acer saccharum diameter distributions were skewed towards the mid-larger size classes. Lack of young and small Acer saccharum stems suggested change in forest composition towards Fagus grandifolia dominance. Most species formed small-scale clusters (15 ) perhaps in response to small gap disturbances. Snags were the dominant dead wood type and were randomly to regularly distributed in most plots. Logfall directions were unrelated to hurricane paths. Recent small-scale disturbance events and topographic position appear to be important in explaining current structure and dynamics of the hardwood and mixed forests of Big Reed Forest Reserve in northern Maine. The continued effects of beech bark disease had a greater effect on hardwood plots, whereas a recent spruce budworm outbreak had a greater effect on plots with higher conifer density.The dominance of very shade tolerant tree species in small-scale clusters, and randomly distributed snags rather than clustered uproots were indicative of the prevalence of small scale gap disturbance regimes in the hardwood and mixed forests of Big Reed Forest Reserve in northern Maine. Varying topographic position may allow for slight changes in disturbance regime leading to consequent variation in structure and dynamics. H1, a more open plot on upper exposed slopes, had distinctly different characteristics such as lower live and dead tree and sapling densities than the other plots, but more uprooted trees and Acer saccharum saplings. Such small scale gap disturbance regimes operating on an episodic basis, and effects of slight variations in this regime on stand composition and structure have significant implications for silvicultural interventions and management of these forest types.     

Spatial genetic structure of aquatic bryophytes in a connected lake system

Using genetic markers, we investigated the genetic structure of three clonal aquatic moss species, Calliergon megalophyllum Mikut., Fontinalis antipyretica Hedw. and F. hypnoides Hartm. on two scales: among populations in a connected lake system (large‐scale spatial genetic structure) and among individuals within populations (fine‐scale spatial genetic structure). Mean genetic diversities per population were 0.138, 0.247 and 0.271, respectively, and total diversities equalled 0.223, 0.385 and 0.421, respectively. Relative differentiation levels (F_ST values of 0.173, 0.280 and 0.142, respectively) were significant but showed that there is a moderate amount of gene flow taking place within the lake system connected with narrow streams. Bayesian STRUCTURE analysis provided some indication that the direction of water flow influences population genetic structuring in the studied aquatic mosses. We propose that dispersal leading to gene flow in C. megalophyllum, F. antipyretica and F. hypnoides takes place both along water via connecting streams and by animal vectors, such as waterfowl. Nevertheless, the slight genetic structuring pattern along the direction of water flow suggests that dispersal of shoots or their fragments along water is a means of dispersal in these mosses. The absence of sexual reproduction and spores may have caused the observed spatial genetic structure within populations, including aggregations of similar genotypes (clones or closely related genotypes) at short distances in populations otherwise showing an isolation by distance effect. Regardless of the results pointing to the dominance of vegetative propagation, it is impossible to completely rule out the potential role of rare long‐distance spore dispersal from areas where the species are fertile.     

Seeing the forest for its multiple ecosystem services: Indicators for cultural services in heterogeneous forests

The ecosystem service (ES) framework is gaining traction in ecosystem management as a means to recognize the multiple benefits that ecosystems provide. In forested ecosystems, many structural attributes (trees, understory plants and woody debris) create heterogeneous ecosystems that provide numerous ecosystem services, including many that are culturally important. However, application of the ES framework to forest management is challenged by difficulties measuring and comparing multiple ES across diverse and heterogeneous forest conditions. Indicators can help bring the ES approach to forest management by providing a means for accurate ES inventory and mapping. We measured 10 forest ES in contrasting forest types to investigate the effects of past forest harvesting in coastal temperate rainforest of Vancouver Island, BC, Canada. Our objectives were to build a systematic set of ES indicators for coastal temperate forests based on forest structural features, including trees, coarse woody debris, and understory plants. To achieve this, we 1) analyzed field data to compare the effects of forest age (old-growth vs. second-growth) and ecological site conditions (riparian vs. upland forest) on the bundle of ES provided by different forest types; and 2) worked with a local indigenous wood carver to identify attributes of cedar trees (Thuja plicata) essential for traditional uses, including canoe carving. Forest age and forest type had significant and major effects on bundles of ES. Old-growth forests provided three times higher carbon storage, nine times higher wood volume, and eighteen times higher canopy habitat services than recovering forests. Within old-growth forests, the proportion of trees suitable for traditional indigenous wood carving was significantly higher in riparian stands. Yet of 456 trees measured, only 17 were cedar with potential traditional uses. Of those, trees for canoe carving were the least frequent (n = 3), which we identified as large (>110 cm DBH) trees of exceptional quality. In general, old-growth riparian forests were a hotspot of ES, providing for example nearly three times as much carbon storage as old-growth forests on upland sites and 12 times the amount of carbon storage as found in second-growth forests on upland sites. These results indicate that typical inventories of forest ES, which usually generalize across heterogeneity, may oversimplify dramatic variations in ES bundles in forested landscapes. Our novel set of stand-level ES indicators can improve the accuracy of ES assessments, incorporate important cultural ES, and help address the role of landscape heterogeneity in influencing ES.     

Separating forest continuity from tree age effects on plant diversity in the ground and epiphyte vegetation of a Central European mountain spruce forest

Forest continuity has been identified as an important factor influencing the structure and diversity of forest vegetation. Primary forests with centuries of continuity are usually more diverse than young secondary forests as forest are colonized only slowly and because the former are richer in old tree individuals. In the present study, performed in unmanaged high-elevation spruce forests of the Harz Mountains, Germany, we had the unique opportunity to separate the effects of forest continuity and tree age on plant diversity. We compared an old-growth spruce forest with century-long habitat continuity with an adjacent secondary spruce forest, which had naturally established on a former bog after 1796 when peat exploitation halted. Comparative analysis of the ground and epiphyte vegetation showed that the plant diversity of the old-growth forest was not higher than that of the secondary forest with a similar tree age of >200 years. Our results suggest that a period of >200 years was sufficient for the secondary forest to be colonized by the whole regional species pool of herbaceous and cryptogam forest plants and epiphytes. Therefore, it is likely that habitat structure, including the presence of old and decaying trees, was more important for determining plant diversity than the independent effect of forest continuity. Our results are probably not transferrable to spruce forests younger than 200 years and highly fragmented woodlands with long distances between new stands and old-growth forests that serve as diaspore sources. In addition, our results might be not transferable to remote areas without notable air pollution, as the epiphyte vegetation of the study area was influenced by SO₂ pollution in the second half of the 20th century.     

Habitat partitioning among neotropical pioneers: a consequence of differential susceptibility to browsing herbivores?

Four species of fast-growing pioneer tree species in the genus Cecropia exist in the forests in central Panama. Cecropia insignis is dominant in old-growth forests but absent from nearby secondary forests; C. obtusifolia, and C. peltata are abundant in secondary forests but rare in old-growth forest, and C. longipes is uncommon in both. To determine whether Cecropia habitat associations are a consequence of local dispersal or differences in recruitment success, we grew seedlings of these species in common gardens in large treefall gaps in secondary and old-growth forest. In contrast to the observed adult distribution, only C. insignis grew significantly over 16 months in secondary forests; remaining species were heavily browsed by herbivores. C. insignis also grew and survived best in old-growth forest. Differences in susceptibility to herbivory did not result from an ant defence mutualism; none of the plants were colonised by ants during the experiment. To test whether C. insignis, the species least susceptible to herbivory, trades off investment in growth in favour of defence, we also grew the four Cecropia species in a screened growing house under light conditions comparable to large forest gaps. Contrary to expectation, species growth rates were similar; only C. peltata grew significantly faster than C. insignis. These results suggest that (1) conditions in ~40-year-old secondary forests no longer support the recruitment of Cecropia species, which are canopy dominants there; and (2) among congeners, differences in plant traits with little apparent cost to growth can have large impacts on recruitment by affecting palatability to herbivores.