Structural and floristic characteristics of some monodominant and adjacent mixed rainforests in New Caledonia

Nothofagus spp. dominate the upper canopy of some rainforests on ultramafic soils in New Caledonia. These monodominant forests typically occur within, or contiguous with, larger areas of mixed‐canopy rainforest. In this study the structure, diversity and composition of six Nothofagus‐dominated plots were investigated, and comparisons were made with three adjacent mixed rainforest plots. Stand density and basal area (all stems ≥ 1.3 m high) in the Nothofagus plots were in the range 16,056–27,550 stems/ha and 43.1–69.9 m²/ha, respectively. There was no significant difference (P ≥ 0.05) in total stand density or basal area between the paired Nothofagus and mixed rainforests, but there were consistently fewer trees and less basal area of trees ≥ 40 cm d.b.h. in the Nothofagus forests. Species richness, species diversity (Shannon‐Wiener, based on basal area) and equitability (based on basal area) of trees ≥ 20 cm d.b.h. on 0.1 ha Nothofagus plots were in the range 4–17, 0.96–3.76 and 0.45–0.87, respectively. No significant differences (P ≥ 0.05) were recorded in these three parameters between the paired Nothofagus and mixed rainforests, although species diversity was consistently lower in the paired Nothofagus forests. Comparison of dominance by density and basal area indicated that although the uppermost canopy of the Nothofagus forests was dominated by Nothofagus (70–95%), the basal area and density contribution was ≤ 55% except at Col de Yaté (≈ 85%). Analysis of similarity indicated no significant difference in stand composition of trees ≥ 20 cm d.b.h. (following removal of Nothofagus from the data set) between Nothofagus and mixed rainforests using basal area, density or presence‐absence data. It is concluded that the Nothofagus‐dominated forests differ from the adjacent mixed rainforests mainly by (1) dominance of the uppermost canopy, without necessarily dominance of the stand by basal area or density, and (2) the smaller basal area contributed by large trees (all species).     

Effects of disturbance and size structure on the regeneration process in a sub-boreal coniferous forest,northern Japan

The stand structure and disturbance history in a sub-boreal coniferous forest dominated byPicea jezoensis, Picea glehnii andAbies sachalinensis were investigated in four study plots set up in Taisetsuzan National Park, Japan. The effect of stand characteristics on the growth and mortality rates of understory trees was examined. Although all the stands showed inverse J-shape d.b.h. (diameter at breast height) distributions, the age structure and disturbance history differed amongst the stands. The stands with wide d.b.h. distribution (i.e. large CV and skewness) were more uneven-aged than those with narrow d.b.h. distribution (i.e. small CV and skewness). The disturbance-return interval based on the model of Hett and Loucks was 31 to 65 years. The gap ratio in the canopy was also different among the stands. These suggest that the variations in stand structure represent different occurrences of natural disturbances. Furthermore, the structural features such as size structure, canopy gap ratio and density of canopy trees also affected the growth dynamics of understory trees (≥2 m in height and <10 cm in diameter at breast height). The growth and mortality rates of understory trees changed with the canopy gap ratio and canopy tree density. The understory trees of stands with wide canopy d.b.h. distribution had higher growth and canopy recruitment rates than those of stands with narrow canopy d.b.h. distribution, contributing to the maintenance of continuous stand stratification. The understory trees of stands with narrow canopy d.b.h. distribution showed lower growth and higher mortality rates than those of stands with narrow canopy d.b.h. distribution, leading to the formation of a single-canopy structure. It is suggested that natural disturbance governs the regeneration process in the future by affecting the growth and mortality patterns of understory trees through the stand structure (size and age structure, canopy tree density, canopy gap ratio).     

Linking radial growth patterns and moderate‐severity disturbance dynamics in boreal old‐growth forests driven by recurrent insect outbreaks: A tale of opportunities,successes, and failures

In boreal landscapes, emphasis is currently placed on close‐to‐nature management strategies, which aim to maintain the biodiversity and ecosystem services related to old‐growth forests. The success of these strategies, however, depends on an accurate understanding of the dynamics within these forests. While moderate‐severity disturbances have recently been recognized as important drivers of boreal forests, little is known about their effects on stand structure and growth. This study therefore aimed to reconstruct the disturbance and postdisturbance dynamics in boreal old‐growth forests that are driven by recurrent moderate‐severity disturbances. We studied eight primary old‐growth forests in Québec, Canada, that have recorded recurrent and moderately severe spruce budworm (Choristoneura fumiferana [Clem.]) outbreaks over the 20th century. We applied an innovative dendrochronological approach based on the combined study of growth patterns and releases to reconstruct stand disturbance and postdisturbance dynamics. We identified nine growth patterns; they represented trees differing in age, size, and canopy layer. These patterns highlighted the ability of suppressed trees to rapidly fill gaps created by moderate‐severity disturbances through a single and significant increase in radial growth and height. Trees that are unable to attain the canopy following the disturbance tend to remain in the lower canopy layers, even if subsequent disturbances create new gaps. This combination of a low stand height typical of boreal forests, periodic disturbances, and rapid canopy closure often resulted in stands constituted mainly of dominant and codominant trees, similar to even‐aged forests. Overall, this study underscored the resistance of boreal old‐growth forests owing to their capacity to withstand repeated moderate‐severity disturbances. Moreover, the combined study of growth patterns and growth release demonstrated the efficacy of such an approach for improving the understanding of the fine‐scale dynamics of natural forests. The results of this research will thus help develop silvicultural practices that approximate the moderate‐severity disturbance dynamics observed in primary and old‐growth boreal forests.     

Forest dynamics following eastern hemlock mortality in the southern Appalachians

Understanding changes in community composition caused by invasive species is critical for predicting effects on ecosystem function, particularly when the invasive threatens a foundation species. Here we focus on dynamics of forest structure, composition and microclimate, and how these interact in southern Appalachian riparian forests following invasion by hemlock woolly adelgid, HWA, Adelges tsugae. We measured and quantified changes in microclimate; canopy mortality; canopy and shrub growth; understory species composition; and the cover and diversity in riparian forests dominated by eastern hemlock Tsuga canadensis over a period of seven years. Treatments manipulated hemlock mortality either through invasion (HWA infested stands) or girdling (GDL) hemlock trees. Mortality was rapid, with 50% hemlock tree mortality occurring after six years of invasion, in contrast to more than 50% mortality in two years following girdling. Although 50% of hemlock trees were still alive five years after infestation, leaf area lost was similar to that of girdled trees. As such, overall responses over time (changes in light transmittance, growth, soil moisture) were identical to girdled stands with 100% mortality. Our results showed different growth responses of the canopy species, shrubs and ground layer, with the latter being substantially influenced by presence of the evergreen shrub, rhododendron Rhododendron maximum. Although ground layer richness in the infested and girdled stands increased by threefold, they did not approach levels recorded in hardwood forests without rhododendron. Increased growth of co‐occurring canopy trees occurred in the first few years following hemlock decline, with similar responses in both treatments. In contrast, growth of rhododendron continued to increase over time. By the end of the study it had a 2.6‐fold higher growth rate than expected, likely taking advantage of increased light available during leaf‐off periods of the deciduous species. Increased growth and dominance of rhododendron may be a major determinant of future responses in southern Appalachian ecosystems; however, our results suggest hemlock will be replaced by a mix of Acer, Betula, Fagus and Quercus canopy genera where establishment is not limited by rhododendron.     

Feedbacks between canopy composition and seedling regeneration in mixed conifer broad‐leaved forests

To address the role of canopy‐seedling feedbacks in the structure and dynamics of mixed conifer broad‐leaved forests in the eastern US, we monitored seedling regeneration patterns and environmental conditions in the understorey of stands dominated by either hemlock (Tsuga canadensis) or red oak (Quercus rubra) for three years. Hemlock seedlings were favoured over other species’ seedlings in hemlock stands (a true positive feedback), due to a combination of high seed inputs, high seedling emergence and relatively high seedling survival during the growing season, which allowed hemlock to remain dominant under its own canopy. Red oak stands favoured a suite of mid‐successional broad‐leaved species over hemlock. A more even age structure of broad‐leaved species in red oak stands revealed that high seedling survival in such stands were driving this feedback. Canopy‐mediated variations in both understorey light availability (1.5% for hemlock vs 3.5% for red oak) and soil pH (3.9 for hemlock vs 4.4 for red oak) were found to be the primary correlates of stand‐level differences in seedling regeneration dynamics. In mixed temperate forests in the eastern US, canopy‐seedling feedbacks could act to slow successional trajectories and contribute to the maintenance of a stable landscape structure over many generations.     

Composition and Dynamics of Functional Groups of Trees During Tropical Forest Succession in Northeastern Costa Rica

We compared the functional type composition of trees ≥10 cm dbh in eight secondary forest monitoring plots with logged and unlogged mature forest plots in lowland wet forests of Northeastern Costa Rica. Five plant functional types were delimited based on diameter growth rates and canopy height of 293 tree species. Mature forests had significantly higher relative abundance of understory trees and slow-growing canopy/emergent trees, but lower relative abundance of fast-growing canopy/emergent trees than secondary forests. Fast-growing subcanopy and canopy trees reached peak densities early in succession. Density of fast-growing canopy/emergent trees increased during the first 20 yr of succession, whereas basal area continued to increase beyond 40 yr. We also assigned canopy tree species to one of three colonization groups, based on the presence of seedlings, saplings, and trees in four secondary forest plots. Among 93 species evaluated, 68 percent were classified as regenerating pioneers (both trees and regeneration present), whereas only 6 percent were classified as nonregenerating pioneers (trees only) and 26 percent as forest colonizers (regeneration only). Slow-growing trees composed 72 percent of the seedling and sapling regeneration for forest colonizers, whereas fast-growing trees composed 63 percent of the seedlings and saplings of regenerating pioneers. Tree stature and growth rates capture much of the functional variation that appears to drive successional dynamics. Results further suggest strong linkages between functional types defined based on adult height and growth rates of large trees and abundance of seedling and sapling regeneration during secondary succession.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp     

Gap-scale disturbance processes in secondary hardwood stands on the Cumberland Plateau,Tennessee, USA

Disturbance regimes in many temperate, old growth forests are characterized by gap-scale events. However, prior to a complex stage of development, canopy gaps may still serve as mechanisms for canopy tree replacement and stand structural changes associated with older forests. We investigated 40 canopy gaps in secondary hardwood stands on the Cumberland Plateau in Tennessee to analyze gap-scale disturbance processes in developing forests. Gap origin, age, land fraction, size, shape, orientation, and gap maker characteristics were documented to investigate gap formation mechanisms and physical gap attributes. We also quantified density and diversity within gaps, gap closure, and gap-phase replacement to examine the influence of localized disturbances on forest development. The majority of canopy gaps were single-treefall events caused by uprooted or snapped stems. The fraction of the forest in canopy gaps was within the range reported from old growth remnants throughout the region. However, gap size was smaller in the developing stands, indicating that secondary forests contain a higher density of smaller gaps. The majority of canopy gaps were projected to close by lateral crown expansion rather than height growth of subcanopy individuals. However, canopy gaps still provided a means for understory trees to recruit to larger size classes. This process may allow overtopped trees to reach intermediate positions, and eventually the canopy, after future disturbance events. Over half of the trees located in true gaps with intermediate crown classifications were Acer saccharum, A. rubrum, or Liriodendron tulipifera. Because the gaps were relatively small and close by lateral branch growth of perimeter trees, the most shade-tolerant A. saccharum has the greatest probability of becoming dominant in the canopy under the current disturbance regime. Half of the gap maker trees removed from the canopy were Quercus; however, Acer species are the most probable replacement trees. These data indicate that canopy gaps are important drivers of forest change prior to a complex stage of development. Even in relatively young forests, gaps provide the mechanisms for stands to develop a complex structure, and may be used to explain patterns of shifting species composition in secondary forests of eastern North America.     

Structural characteristics and aboveground biomass of old‐growth spruce–fir stands in the eastern Carpathian mountains,Ukraine

Abstract

Temperate old‐growth forests are known to have ecological characteristics distinct from younger forests, but these have been poorly described for the remaining old‐growth Picea abies–Abies alba forests in the eastern Carpathian mountains. In addition, recent studies suggest that old‐growth forests may be more significant carbon sinks than previously recognized. This has stimulated interest in quantifying aboveground carbon stocks in primary forest systems. We investigated the structural attributes and aboveground biomass in two remnant old‐growth spruce–fir stands and compared these against a primary (never logged) mature reference stand. Our sites were located in the Gorgany Nature Reserve in western Ukraine. Overstory data were collected using variable radius plots; coarse woody debris was sampled along line intercept transects. Differences among sites were assessed using non‐parametric statistical analyses. Goodness‐of‐fit tests were used to evaluate the form of diameter distributions. The results strongly supported the hypothesis that old‐growth temperate spruce–fir forests have greater structural complexity compared to mature forests, including higher densities of large trees, more complex horizontal structure, and elevated aboveground biomass. The late‐successional sites we sampled exhibited rotated sigmoid diameter distributions; these may reflect natural disturbance dynamics. Old‐growth Carpathian spruce–fir forests store on average approximately 155–165 Mg ha^?1 of carbon in aboveground tree parts alone. This is approximately 50% higher than mature stands. Given the scarcity of primary spruce–fir forests in the Carpathian region, remaining stands have high conservation value, both as habitat for late‐successional species and as carbon storage reservoirs.     

Changing growth response to wildfire in old‐growth ponderosa pine trees in montane forests of north central Idaho

North American fire‐adapted forests are experiencing changes in fire frequency and climate. These novel conditions may alter postwildfire responses of fire‐adapted trees that survive fires, a topic that has received little attention. Historical, frequent, low‐intensity wildfire in many fire‐adapted forests is generally thought to have a positive effect on the growth and vigor of trees that survive fires. Whether such positive effects can persist under current and future climate conditions is not known. Here, we evaluate long‐term responses to recurrent 20th‐century fires in ponderosa pine, a fire‐adapted tree species, in unlogged forests in north central Idaho. We also examine short‐term responses to individual 20th‐century fires and evaluate whether these responses have changed over time and whether potential variability relates to climate variables and time since last fire. Growth responses were assessed by comparing tree‐ring measurements from trees in stands burned repeatedly during the 20th century at roughly the historical fire frequency with trees in paired control stands that had not burned for at least 70 years. Contrary to expectations, only one site showed significant increases in long‐term growth responses in burned stands compared with control stands. Short‐term responses showed a trend of increasing negative effects of wildfire (reduced diameter growth in the burned stand compared with the control stand) in recent years that had drier winters and springs. There was no effect of time since the previous fire on growth responses to fire. The possible relationships of novel climate conditions with negative tree growth responses in trees that survive fire are discussed. A trend of negative growth responses to wildfire in old‐growth forests could have important ramifications for forest productivity and carbon balance under future climate scenarios.     

亚高山云冷杉混交林树木生长释放与干扰分析

对云南碧塔海亚高山云冷杉林内４个样地冠层树木的生长压制和释放的历史,用树木年轮分析方法进行了重建,然后根据生长释放频率推测林冠干扰强度（每１０年冠层树木的死亡百分率）。４个林分（１个中龄林,３个成过熟林）生长释放的平均百分率为４８％～９２％。中龄林内,平均生长释放频率为７１％／１０ａ,成过熟林则为７４％～９５％／１０ａ,在油麦吊云杉〔Ｐｉｃｅａｂｒａｃｈｙｔｙｌａｖａｒ．ｃｏｍｐｌａｎａｔａ（Ｍａｓｔ．）ＣｈｅｎｇｅｘＲｅｈｄ．〕占优势的林分和大果红杉（Ｌａｒｉｘｐｏｔａｎｉｎｉｖａｒ．ｍａｃｒｏｃａｒｐａＬａｗ）油麦吊云杉混交林分内,估测的林冠干扰强度分别为４８％／１０ａ和５９％／１０ａ。     

Comparison of structure and biodiversity in the Rajhenav virgin forest remnant and managed forest in the Dinaric region of Slovenia

Comparisons are made between a virgin forest remnant (primeval forest) and a lightly managed (near‐to‐nature) forest with regard to horizontal forest structures, the structure of forest stands, and the diversity of plant and bird species. In the virgin forest remnant the proportion of canopy gaps is smaller, there are no stands in the developmental phase of a pole stand (10 < cm d.b.h. < 30 cm), and both the growing stock and the proportion of less vigorous trees are considerably greater. In addition, there is a higher percentage of dead trees, a smaller proportion of minor tree species and a considerably lower diversity of plants. The diversity of bird species is similar for each forest type, but rare bird species are confined to the virgin forest remnant. The biological differences between the two types of forest have led to changes in the physical site conditions. The results of the research are valuable in assessing the consequences of near‐to‐nature forest management.     

Conservation management of Pinus palustris ecosystems from a landscape perspective

Question: Do case studies from silvicultural and restoration studies and applied conservation management in second‐growth Pinus palustris stands provide unique insights for conservation models? Methods: A review of management paradigms that conserve the high biological diversity and rare species, drawn from characteristics in both second‐growth and old‐growth stands, is presented for fire‐maintained Pinus palustris (longleaf pine) forests. Results: A common assumption that old‐growth stands provide the primary information for the development of conservation management strategies de‐emphasizes lessons learned from second‐growth and restoration studies. Primary conservation management goals for the Pinus palustris ecosystem include the perpetual regeneration of the fire‐maintained forest and conservation of the characteristically high biological diversity and rare species. Several attributes, such as a sustained population of Picoides borealis (red‐cockaded woodpecker), Aristida stricta (wiregrass)‐dominated ground cover, and undisturbed upland‐wetland ecotones, can predict a diverse and ecologically functional ecosystem. Such indicators are linked to critical structural and functional features of the system and reflect previous land management histories that suggest sustainable approaches. Conclusions: A traditional definition of ‘old‐growth’ relying on overstorey may be limited in describing important features of healthy, diverse Pinus palustris ecosystems. Some characteristics are significantly more important for maintenance of diversity than age of the trees and these features may be present in old‐ or second‐growth forests. We advocate that the management history, structural characteristics and landscape context of stands that harbour desirable conservation attributes (red‐cockaded woodpeckers, wiregrass, gopher tortoises and undisturbed upland‐wetland ecotones) can be used as indicators to identify important conservation and forest management principles.     

Structural segregation and scales of spatial dependency in Abies amabilis forests

Question: Is a mosaic structure apparent in the spatial distribution of trees in old‐growth Abies amabilis forests? Location: Montane forests of the western Cascade Range, Washington, USA. Methods: Maps of tree locations were created for study areas located in two, 300‐year old stands and a single 600‐year old stand. Stand structure parameters were calculated using several subsample quadrats sizes (56.25 ‐ 306.25 m²), which were drawn randomly with replacement at a density of 250 quadrats per ha from the stem maps in the computing environment. Spatial cross‐covariance functions between different canopy strata were estimated using the spline cross‐correlogram. Results: Negative spatial correlation (segregation) between subcanopy tree density and areas of high overstorey occupancy was detected. Understorey and midstorey tree densities were positively spatially correlated. These general trends were apparent across the range of observational scales investigated. Significant spatial correlation between canopy strata was observed at spatial scales of 12 ‐ 44 m and extended to the largest scales in the 600‐year old stand. Conclusion: The observed spatial segregation between canopy strata supports the hypothesis that old A. amabilis forests form fine‐scale structural mosaics. Structural segregation at small scales may be due to competitive interactions as well as exogenous forcing of tree locations (e.g. by mortality due to pathogens or disturbance), however segregation at large scales in the 600‐year old stand is likely due to exogenous factors alone. This study reinforces the idea that horizontal heterogeneity is an emergent property of old‐growth forests.     

A long‐term record of Nothofagus dominance in the southern Andes,Chile

Abstract The general model of regeneration dynamics in Nothofagus forests of southern South America could have value in community ecology if predictive relationships between disturbance history, functional traits and site attributes could be identified. Examined here is the proposal that on favourable sites shade‐intolerant Nothofagus are likely not to survive in competition with shade‐tolerant, broad‐leaved evergreen taxa of temperate rain forests, and persistence, thus, is dependent on periodic coarse‐scale disturbance. Comparison of stand dynamics of three old‐growth Nothofagus forests at different elevations in the southern Andes, Chile where deciduous Nothofagus alpina dominates the upper canopy, and examination of the life history trade‐offs of this variation were made. Stem density of all stems ≥5.0 cm d.b.h. was 233–303 stems per hectare, and basal area was 123.9–171.0 m²ha^?1. Maximum lifespan of N. alpina was found to be greater than ca 640 years, exceeding all previously reported ages for this species in the region. Forests had a stable canopy composition for this long‐term, but some appeared to lack effective regeneration of N. alpina in recent years. Regeneration of N. alpina was generally greater in disturbed stands and higher elevation than in undisturbed stands and at lower elevation. Recruitment emerged to be strongly affected by competitive over‐ and understorey associates. There was a gradient of increasing dependence of N. alpina on disturbance towards the more productive end of the environment gradients, and hence less dependence of N. alpina on disturbance for its regeneration towards higher elevation. The study confirms that changes in forest composition may be explained by processes occurring in accordance with the predictions of the existing model of Nothofagus regeneration dynamics, providing stronger evidence specifically directed at mid‐tolerant N. alpina, and by factoring out regeneration dynamics on favourable sites. Thus, for N. alpina, trait differences probably contribute to the competitive advantage over its associates in productive habitats, and may be linked to small‐to‐intermediate‐sized disturbances which inevitably occur as older trees die, enabling N. alpina to persist in forests and therefore maintain species coexistence for the long‐term.     

Present and past old‐growth forests of the Lake Tahoe Basin,Sierra Nevada,US

Abstract. We described 38 relictual old‐growth stands – with data on the mortality, regeneration, floristic richness, fuel load and disease incidence in our study area in the Tahoe Basin of California and Nevada. The stands are within the lower and upper montane zones (1900–2400 m a.s.l.) and they are rare, occupying < 2% of the land in the Basin's watershed. Correlation matrices and ANOVAs of forest types and conifer species with environmental gradients revealed significant relationships with elevation, distance east of the Sierran crest, slope aspect, annual precipitation, date of complete snow melt, litter depth and degree of soil profile development. Pathogens, parasites and wood‐boring insects were present on 23% of living trees; 16% of all trees were dead. We compared these stands to a reconstruction of pre‐contact Basin forests and to ecologically analogous old‐growth forests of Baja California that have never experienced fire suppression management. Currently, overstorey trees (> 180 yr old) in the Basin stands have ca. 33% cover, 54 m².ha^‐1 basal area and 107 individuals.ha^‐1, values very similar to reconstructions of pre‐contact Basin forests and to modern Baja California forests. Understorey trees (60–180 yr old), however, are several times more dense than historic levels and species composition is strongly dominated by A. concolor, regardless of the overstorey composition. The ratio of Pinus: Abies has increased – and the age structure of extant stands predicts that it will continue to increase – from approximately 1:1 in pre‐contact time to 1:7 within the next century. Disease incidence and mortality in Baja forests were lower. Although we quantitatively defined current Basin old‐growth forests – in terms of stand structure – we realize that our definition will differ from that of both past and future old‐growth forests unless management protocols are changed.     

Structure, floristic composition and natural regeneration in the forests of Cat Tien National Park, Vietnam: an analysis of the successional trends

The forests in Cat Tien National Park, appear as a mosaic of different communities, distinct from each other with respect to their floristic and structural parameters. The objectives of this study are (1) to characterize the different formations occurring in the lowland part and (2) to identify the main successional trends in the area. Understanding forest succession is important for silviculture and restoration of forests and land rehabilitation, as adequate information on the ecological role of local species in the functioning of the forests is not available in Vietnam. Five plots (1 ha each) were established in the lowland part of Cat Tien National Park, where all the trees ≥ 10 cm d.b.h. (diameter at breast height) were located, measured and identified. A systematic sampling was made to assess the regeneration. Three plots (A, C and D) can be considered as secondary forests on the basis of their structural parameters. Plots A and C are dominated by Lagerstrmia calyculata and plot D by Dipterocarpus alatus. The other two plots can be regarded as mature forests. Plot B corresponds to a semideciduous formation dominated by Lagerstrmia calyculata and Fabaceae species, and plot E to an evergreen one dominated by dipterocarp species. The floristic composition of plots A and C will change in the future because dominant canopy species are rare or absent in regeneration. A correspondence analysis performed on the number of trees per species shows two kinds of successional trends: one from A to B on shallow and drier soils, and another from C to E on deeper and wetter soils.     

Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions

Stand density reductions have been proposed as a method by which old‐growth ponderosa pine (Pinus ponderosa) forests of North America can be converted back to pre‐1900 conditions, thereby reducing the danger of catastrophic forest fires and insect attacks while increasing the productivity of the remaining old‐growth individuals. However, the duration of productivity response of individual trees and the physiological mechanisms underlying such a response remain speculative issues, particularly in old trees. Tree‐ring measurements of carbon isotope ratios (δ¹³C) and basal area increment (BAI) were used to assess the response of intrinsic water‐use efficiency (the ratio of photosynthesis, A to stomatal conductance, g) and growth of individual> 250‐year‐old‐ponderosa pine trees to stand density reductions. It was hypothesized that reductions in stand density would increase soil moisture availability, thus decreasing canopy A/g and increasing carbon isotope discrimination (Δ). Cellulose‐δ¹³C of annual tree rings, soil water availability (estimated from pre‐dawn leaf water potential), photosynthetic capacity, stem basal growth and xylem anatomy were measured in individual trees within three pairs of thinned and un‐thinned stands. The thinned stands were treated 7 to 15 years prior to measurement. The values of δ¹³C and BAI were assessed for 20 consecutive years overlapping the date of thinning in a single intensively studied stand, and was measured for 3 years on either side of the date of thinning for the two other stands to assess the generality of the response. After thinning, Δ increased by 0.89‰ (± 0.15‰). The trees in the un‐thinned stands showed no change in Δ (0.00‰ ± 0.04‰). In the intensively studied trees, significant differences were expressed in the first growing season after the thinning took place but it took 6 years before the full 0.89‰ difference was observed. BAI doubled or tripled after disturbance, depending on the stand, and the increased BAI lasted up to 15 years after thinning. In the intensively studied trees, the BAI response did not begin until 3 years after the Δ response, peaked 1 year after the Δ peak, and then BAI and Δ oscillated in unison. The lag between BAI and Δ was not due to slow changes in anatomical properties of the sapwood, because tracheid dimensions and sapwood‐specific conductivity remained unchanged after disturbance. The Δ response of thinned trees indicated that A/g decreased after thinning. Photosynthetic capacity, as indexed by foliar nitrogen ([N]) and by the relationship between photosynthesis and internal CO₂ (A–C_i curves), was unchanged by thinning, confirming our suspicion that the decline in A/g was due to a relatively greater increase in g in comparison with A. Model estimates agreed with this conclusion, predicting that g increased by nearly 25% after thinning relative to a 15% increase in A. Pre‐dawn leaf water potential averaged 0.11 MPa (± 0.03 MPa) less negative for the thinned compared with the un‐thinned trees in all stands, and was strongly correlated with Δ post‐thinning (R² = 0.91). There was a strong relationship between BAI and modelled A, suggesting that changes in water availability and g have a significant effect on carbon assimilation and growth of these old trees. These results confirm that stand density reductions result in increased growth of individual trees via increased stomatal conductance. Furthermore, they show that a physiological response to stand density reductions can last for up to 15 years in old ponderosa pines if stand leaf area is not fully re‐established.     

Identification of Old‐Growth Forest Reference Ecosystems Using Historic Land Surveys,Redwood National Park,California

Old‐growth forests in the American West typically represent fragments of former, more extensive forests that were subjected to nineteenth and twentieth century land‐clearing activities, such as logging. These present‐day forest fragments are thought to be representative of the former landscape, and thus are capable of serving as living references for guiding restoration of logged forests. Yet how do we determine the extent to which existing old‐growth stands represent the former forest, especially when little of the surrounding original vegetation remains? Historic land surveys conducted prior to significant logging can reconstruct the former forest at the stand level, thereby allowing an analysis of old‐growth patches within the larger historic landscape. This study utilized original Public Land Surveys to assess the applicability of old‐growth stands in Redwood National Park as reference ecosystems. A geographic information system (GIS) and statistical analysis of the nineteenth century forest found that vegetation communities, woody species composition, and ratios of dominant canopy species in unlogged patches were highly representative of the forests that were logged. Significance testing (H_o: μ₁ = μ₂) revealed p‐values greater than 0.10000 in all measures of community and species composition, except for the higher abundance of oak in present‐day old‐growth (p‐value = 0.0395). The results of this study suggest that the national park should increase efforts to protect old‐growth reference ecosystems from further human impacts, and minimize ongoing degradation from edge effects by prioritizing restoration of adjoining second‐growth forest.     

Past levels of canopy closure affect the occurrence of veteran trees and flagship saproxylic beetles

Aim

Open woodlands are biologically highly diverse habitats, and veteran (i.e., old, senescent) trees are key structures supporting their biodiversity. Open canopy structure had been maintained by both natural‐ and human‐induced disturbances. In the past two centuries, suppression of such disturbances, together with forestry intensification, has turned most lowland woodlands into closed‐canopy forests. We investigated the effect of increased canopy closure on veteran trees and several threatened beetles associated with them.

Location

Floodplain woodlands along the lower Dyje and Morava rivers, Czech Republic.

Methods

We used an approach combining the study of aerial photographs with on‐ground survey of veteran trees and associated endangered beetles. The aerial images were used to obtain the information on historical (1938) and recent (2009) canopy closure in the area of 146 km², where we mapped large oaks (d.b.h. >70 cm), hollow trees and three associated beetles including the hermit beetle (Osmoderma barnabita), the great capricorn beetle (Cerambyx cerdo) and the jewel beetle Eurythyrea quercus.

Results

The presence of large oaks, hollow trees and their associated beetle species are negatively related to recent high canopy closure, and the historical level of canopy closure matters, as in nowadays closed‐canopy stands, the beetles and veteran trees are more common in places that were rather open in 1938 than in the places with closed canopy already in 1938. Moreover, the health state of veteran trees highly depends on the canopy closure.

Main conclusion

The negative effect of canopy closure on veteran trees and their endangered inhabitants is several decades delayed and may thus often go undetected. In the forests, however, large and hollow trees and their associated biodiversity are relics of the past, more open conditions. The restoration of open woodlands is therefore vital for preventing their further decline. Conservation management planning needs to take this into account wherever, veteran trees and associated biota are concerned.