Effect of leaf flutter on the light environment of poplars

The dynamics of the canopy light environment for two poplar species (Populus tremuloides Michx., and P. fremontii Wats.) were characterized with an array of photocells in fixed positions within the canopy or attached directly to leaves and using a data logger that recorded photon flux density (PFD) at frequencies from 1 to 20 Hz. The majority of sunflecks were short in duration (<1 s) with a similar short interval between sunflecks. Sunflecks contribute as much as 90% of the total daily PFD in the lower canopy. Leaf flutter may cause high frequency (3 to 5 Hz) variations of PFD in poplar canopies. The amount of light intercepted by a fluttering leaf at the top of the canopy decreased with increasing flutter, whereas a fluttering lower canopy leaf showed no such trend. When leaves fluttered at the top of the canopy the understory light environment showed an increased number of shorter sunflecks. Leaf flutter may increase mean PFD for understory leaves. It also creates a canopy light environment that is more dynamic temporally and more evenly distributed spatially. The potential benefits of these changes in light dynamics are discussed.     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

VAM association in the shrub Myrica cerifera on a Virginia,USA barrier island

A combined laboratory and field study examined the potential for a symbiotic association between the actinorhizal shrub Myrica cerifera and vesicular arbuscular mycorrhizal (VAM) fungi on a Virginia barrier island. M. cerifera seedlings and two test species, Zea mays and Strophostyles umbellata, were grown in an environmental chamber on soils collected from four sites differing in soil age (< 5 to over 130 years), salinity (1–35 g/g total soil chloride), and edaphic characteristics. Seedling root infection was significantly lower for all three species in the youngest soils from the beach where salinity was highest. Stained M. cerifera roots revealed all the components for a functional VAM association; however, there were significantly fewer arbuscules and vesicles relative to the test species. Among field-collected M. cerifera, infection was not detected in mature shrubs from the bay side of the island, where M. cerifera thickets were in a state of degeneration. Infection was highest in soils from the young, developing thickets, and in the most stable thickets of the island interior. Despite the dynamic nature of the barrier island environment, VAM associations with M. cerifera appear to be present, especially in seedlings and developing shrub thickets.     

Ecophysiology of exotic and native shrubs in Southern Wisconsin

Summary We compared seasonal trends in photosynthesis of two naturalized exotic shrubs (Rhamnus cathartica and Lonicera X bella) and two native shrubs (Cornus racemosa and Prunus serotina) in open and understory habitats in southern Wisconsin. We examined the relationships between resource availability and leaf photosynthetic performance in these four species. All four species had similar relationships between leaf nitrogen (N) content and photosynthetic rate, but the species differed in absolute leaf N content and therefore in photosynthetic rates. Maximum daily photosynthetic rates of all species were significantly correlated with leaf N content in the open habitat, but not in the understory, where low light availability was the major limitation to photosynthesis. Extended leaf longevity was important in the forest understory because it allowed shrubs to take advantage of high light availability at times when the overstory canopy was leafless. Early leaf emergence was more important than late senescence: from 27% to 35% of the annual carbon gain of P. serotina, R. cathartica, and L. X bella occurred prior to leaf emergence of C. racemosa, the species with the shortest leaf life span. Extended leaf longevity of exotic shrubs may help explain their persistence in the understory habitat, but it contributed relatively less to their annual carbon gain in the open habitat.     

Light limitation creates patchy distribution of an invasive grass in eastern deciduous forests

Species interactions and their indirect effects on the availability and distribution of resources have been considered strong determinants of community structure in many different ecological systems. In deciduous forests, the presence of overstory trees and shrubs creates a shifting mosaic of resources for understory plants, with implications for their distribution and abundance. Determination of the ultimate resource constraints on understory vegetation may aid management of these systems that have become increasingly susceptible to invasions by non-native plants. Microstegium vimineum (Japanese grass) is an invasive annual grass that has spread rapidly throughout the understory of forests across the eastern United States since it was first observed in Tennessee in 1919. M. vimineum occurs as extensive, dense patches in the understory of eastern deciduous forests, yet these patches often exhibit sharp boundaries and distinct gaps in cover. One example of this distributional pattern was observed relative to the native midstory tree Asimina triloba (pawpaw), whereby dense M. vimineum cover stopped abruptly at the drip line of the A. triloba patch and was absent beneath the A. triloba canopy. We conducted field and greenhouse experiments to test several hypotheses regarding the causes of this observed pattern of M. vimineum distribution, including allelopathy, seed dispersal, light limitations, and soil moisture, texture, and nutrient content. We concluded that light reduction by the A. triloba canopy was the environmental constraint that prevented establishment of M. vimineum beneath this tree. Whereas overstory tree canopy apparently facilitates the establishment of this shade-tolerant grass, the interaction of overstory canopy with midstory canopy interferes with M. vimineum by reducing the availability of sunflecks at the ground layer. It is likely that other midstory species influence the distribution and abundance of other herb-layer species, with implications for management of understory invasive plant species.     

Seasonal variation of temperature response of respiration in invasive <Emphasis Type="Italic">Berberis thunbergii</Emphasis> (Japanese barberry) and two co-occurring native understory shrubs in a northeastern US deciduous forest

In the understory of a closed forest, plant growth is limited by light availability, and early leafing is proposed to be an important mechanism of plant invasion by providing a spring C “subsidy” when high light is available. However, studies on respiration, another important process determining plant net C gain, are rare in understory invasive plants. In this study, leaf properties and the temperature response of leaf respiration were compared between invasive Berberis thunbergii, an early leafing understory shrub, and two native shrubs, Kalmia latifolia, a broadleaf evergreen and Vaccinium corymbosum, a late-leafing deciduous species, in an oak-dominated deciduous forest. The seasonal trend of the basal respiration rates (R ₀) and the temperature response coefficient (E ₀), were different among the three shrubs and species-specific negative correlations were observed between R ₀ and E ₀. All three shrubs showed significant correlation between respiration rate on an area basis (20°C) and leaf N on an area basis. The relationship was attributed to the variation of both leaf N on a mass basis and leaf mass per area (LMA) in B. thunbergii, but to LMA only in K. latifolia and V. corymbosum. After modeling leaf respiration throughout 2004, B. thunbergii displayed much higher annual leaf respiration (mass based) than the two native shrubs, indicating a higher cost per unit of biomass investment. Thus, respiratory properties alone were not likely to lead to C balance advantage of B. thunbergii. Future studies on whole plant C budgets and leaf construction cost are needed to address the C balance advantage in early leafing understory shrubs like B. thunbergii.     

Light environment under Rhododendron maximum thickets and estimated carbon gain of regenerating forest tree seedlings

Canopy tree recruitment is inhibited by evergreen shrubs in many forests. In the southern Appalachian mountains of the USA, thickets of Rhododendron maximum L. restrict dominant canopy tree seedling survival and persistence. Using R. maximum as a model system, we examined available light under the thickets and the photosynthetic responses of seedlings of canopy tree species. We tested the hypothesis that the additional shading from under R. maximum drives carbon gain in seedlings below the threshold for growth and survival. A reduction in light under the thicket was found where canopy openness (derived from canopy photographs) under R. maximum was half the amount measured in forest without R. maximum. R.␣maximum also reduced direct radiation by 50% and diffuse radiation by 12–29% compared to forest without the shrub layer. Mean mid-day PPFD (photosynthetically active photon flux density between 1000 and 1400 h) under R. maximum (obtained from quantum sensors) was below 10 mol m⁻² s⁻¹ on both clear and overcast days and the amount of sunflecks greater than 10 mol m⁻² s⁻¹ PPFD was only 0–20 min per day. In contrast, forest without R. maximum received a mean PPFD of 18–25 mol m⁻² s⁻¹ on clear days and a cumulative sunfleck duration of 100–220 min per day in all sky conditions. Consistent with light availability between the sites, daily carbon gain in Quercus rubra L. seedlings was lower in forest with R. maximum compared to forest where the shrub was absent. The presence of the shrub layer also significantly suppressed average mid-day photosynthesis of both Q. rubra and Prunus serotina Ehrt. seedlings on 8 out of 11 measurement dates. However, parameters derived from light response curves between seedlings growing in forest sites with or without a thicket of R. maximum was significantly different only in A _max (maximum photosynthetic rate), indicating a lack of further acclimation to the deeper shade under R. maximum. While the additional shade cast by R. maximum is sufficient to prevent the regeneration of tree seedlings under this shrub, there was sufficient heterogeneity in light under the thicket to imply that deep shade only partially explains the complete inhibition of regenerating canopy trees under R. maximum.     

Short-Term Effects of Fire and Other Fuel Reduction Treatments on Breeding Birds in a Southern Appalachian Upland Hardwood Forest

Abstract: We compared the effects of 3 fuel reduction techniques and a control on breeding birds during 2001-2005 using 50-m point counts. Four experimental units, each >14 ha, were contained within each of 3 replicate blocks at the Green River Game Land, Polk County, North Carolina, USA. Treatments were 1) prescribed burn, 2) mechanical understory reduction (chainsaw-felling of shrubs and small trees), 3) mechanical + burn, and 4) controls. We conducted mechanical treatments in winter 2001-2002 and prescribed burns in spring 2003. Tall shrub cover was substantially reduced in all treatments compared to controls. Tree mortality and canopy openness was highest in the mechanical + burn treatment after burning, likely due to higher fuel loading and hotter burns; tree mortality increased with time. Many bird species did not detectably decrease or increase in response to treatments. Species richness, total bird density, and some species, including indigo buntings (Passerina cyanea) and eastern bluebirds (Sialia sialis), increased in the mechanical + burn treatment after a 1-year to 2-year delay; eastern woodpewees (Contopus virens) increased immediately after treatment. Hooded warblers (Wilsonia citrina), black-and-white warblers (Mniotilta varia), and worm-eating warblers (Helmitheros vermivorus) declined temporarily in some or all treatments, likely in response to understory and (or) leaf litter depth reductions. Densities of most species affected by treatments varied with shrub cover, tree or snag density, or leaf litter depth. High snag availability, open conditions, and a higher density of flying insects in the mechanical + burn treatment likely contributed to increased bird density and species richness. In our study, fuel reduction treatments that left the canopy intact, such as low-intensity prescribed fire or mechanical understory removal, had few detectable effects on breeding birds compared to the mechanical + burn treatment. High-intensity burning with heavy tree-kill, as occurred in our mechanical + burn treatment, can be used as a management tool to increase densities of birds associated with open habitat while retaining many forest and generalist species, but may have short-term adverse effects on some species that are associated with the ground- or shrub-strata for nesting and foraging.     

Leaf δ13C in Pinus resinosa trees and understory plants: variation associated with light and CO2 gradients

 Our objective was to evaluate the relative importance of gradients in light intensity and the isotopic composition of atmospheric CO₂ for variation in leaf carbon isotope ratios within a Pinus resinosa forest. In addition, we measured photosynthetic gas exchange and leaf carbon isotope ratios on four understory species (Dryopteris carthusiana, Epipactus helleborine, Hieracium floribundum, Rhamnus frangula), in order to estimate the consequence of the variation in the understory light microclimate for carbon gain in these plants. During midday, CO₂ concentration was relatively constant at vertical positions ranging from 15 m to 3 m above ground. Only at positions below 3 m was CO₂ concentration significantly elevated above that measured at 15 m. Based on the strong linear relationship between changes in CO₂ concentration and δ¹³C values for air samples collected during a diurnal cycle, we calculated the expected vertical profile for the carbon isotope ratio of atmospheric CO₂ within the forest. These calculations indicated that leaves at 3 m height and above were exposed to CO₂ of approximately the same isotopic composition during daylight periods. There was no significant difference between the daily mean δ¹³C values at 15 m (–7.77‰) and 3 m (–7.89‰), but atmospheric CO₂ was significantly depleted in ¹³C closer to the ground surface, with daily average δ¹³C values of –8.85‰ at 5 cm above ground. The light intensity gradient in the forest was substantial, with average photosynthetically active radiation (PAR) on the forest floor approximately 6% of that received at the top of the canopy. In contrast, there were only minor changes in air temperature, and so it is likely that the leaf-air vapour pressure difference was relatively constant from the top of the canopy to the forest floor. For red pine and elm tree samples, there was a significant correlation between leaf δ¹³C value and the height at which the leaf sample was collected. Leaf tissue sampled near the forest floor, on average, had lower δ¹³C values than samples collected near the top of the canopy. We suggest that the average light intensity gradient through the canopy was the major factor influencing vertical changes in tree leaf δ¹³C values. In addition, there was a wide range of variation (greater than 4‰) among the four understory plant species for average leaf δ¹³C values. Measurements of leaf gas exchange, under natural light conditions and with supplemental light, were used to estimate the influence of the light microclimate on the observed variation in leaf carbon isotope ratios in the understory plants. Our data suggest that one species, Epipactus helleborine, gained a substantial fraction of carbon during sunflecks. Received: 21 March 1996 / Accepted: 13 August 1996     

Response of Understory Species to Gap Formation and Soil Disturbance m Lonicera maackii Thickets

We studied the utility of gap formation and soil disturbance as methods to enhance establishment of plant species in the understory of a northern Kentucky forest where Lonicera maackii (Amur honeysuckle) produced dense thickets. In May 1994, gaps (5 m diameter) were cut in the shrub thicket. In adjacent areas, the shrub canopy remained intact. Subplots were established where soil was either turned with a spade to a depth of 15 cm or not disturbed. We monitored plant establishment for three growing seasons (1994, 1995, and 1996). Shrub removal increased light availability to about 10% of full sun. Gap formation had a significant (p < 0.05) and positive influence on total plant density (exclusive of L. maackii), and soil disturbance did not (p > 0.05). After three growing seasons, the most important species were L. maackii, Alliaria petiolata, Parthenocissus quinquefolia, Vitis vulpina, and Acer negundo. Of these species, only V. vulpina showed significantly (p < 0.05) higher densities in gaps. Other less important species such as Phytolacca americana, Campsis radicans, and Eupatorium rugosum occurred almost exclusively in gaps. Of the 44 taxa observed in this study, most were generalist species that also occur in early successional habitats. Long-term dominance of the understory by L. maackii has likely modified system attributes with corresponding effects on community development. Shrub removal provides a window of establishment for various plant species, but successful restoration may require further management species availability and to control new invaders.     

Trade-offs between light interception and leaf water shedding: a comparison of shade- and sun-adapted species in a subtropical rainforest

Species in high-rainfall regions have two major alternative approaches to quickly drain off water, i.e., increasing leaf inclination angles relative to the horizontal plane, or developing long leaf drip tips. We hypothesized that shade-adapted species will have more pronounced leaf drip tips but not greater inclination angles (which can reduce the ability to intercept light) compared to sun-adapted species and that length of leaf drip tips will be negatively correlated with photosynthetic capacity [characterized by light-saturated net photosynthetic rates (A _max), associated light compensation points (LCP), and light saturation points (LSP)]. We tested this hypothesis by measuring morphological and physiological traits that are associated with light-interception and water shedding for seven shade-adapted shrub species, ten sun-adapted understory shrub species, and 15 sun-adapted tree species in a subtropical Chinese rainforest, where mean annual precipitation is around 1,600 mm. Shade-adapted understory species had lower LMA, A _max, LSP, and LCP compared to understory or canopy sun-adapted species; their leaf and twig inclination angles were significantly smaller and leaf drip tips were significantly longer than those in sun-adapted species. This suggests that shade-adapted understory species tend to develop pronounced leaf drip tips but not large leaf inclination angles to shed water. The length of leaf drip tips was negatively correlated with leaf inclination angles and photosynthetic capacity. These relationships were consistent between ordinary regression and phylogenetic generalized least squares analyses. Our study illustrates the trade-offs between light interception and leaf water shedding and indicates that length of leaf drip tips can be used as an indicator of adaptation to shady conditions and overall photosynthetic performance of shrub species in subtropical rainforests.     

How tree species identity and diversity affect light transmittance to the understory in mature temperate forests

Light is a key resource for plant growth and is of particular importance in forest ecosystems, because of the strong vertical structure leading to successive light interception from canopy to forest floor. Tree species differ in the quantity and heterogeneity of light they transmit. We expect decreases in both the quantity and spatial heterogeneity of light transmittance in mixed stands relative to monocultures, due to complementarity effects and niche filling. We tested the degree to which tree species identity and diversity affected, via differences in tree and shrub cover, the spatiotemporal variation in light availability before, during, and after leaf expansion. Plots with different combinations of three tree species with contrasting light transmittance were selected to obtain a diversity gradient from monocultures to three species mixtures. Light transmittance to the forest floor was measured with hemispherical photography. Increased tree diversity led to increased canopy packing and decreased spatial light heterogeneity at the forest floor in all of the time periods. During leaf expansion, light transmittance did differ between the different tree species and timing of leaf expansion might thus be an important source of variation in light regimes for understory plant species. Although light transmittance at the canopy level after leaf expansion was not measured directly, it most likely differed between tree species and decreased in mixtures due to canopy packing. A complementary shrub layer led, however, to similar light levels at the forest floor in all species combinations in our plots. Synthesis. We find that a complementary shrub layer exploits the higher light availability in particular tree species combinations. Resources at the forest floor are thus ultimately determined by the combined effect of the tree and shrub layer. Mixing species led to less heterogeneity in the amount of light, reducing abiotic niche variability.     

山西灵空山天然松栎混交林群落特征与冠层结构

以2011年建设的山西灵空山4 hm²天然松栎混交林森林动态监测样地为研究平台,以400个10 m×10 m样方为测量单元,于2016年进行群落特征研究,采用半球面影像法(DHP)分析冠层结构和林下光照特征.结果表明: 样地内共有乔木5558株,共计25种,分属于10科15属.冠层开阔度(CO)集中在15.0%～25.0%,叶面积指数(LAI)集中在1.5～2.5,林下光环境参数集中在10.0%～30.0%.建群种在样地内的分布对冠层结构和林下光环境影响显著;冠层结构对林下光环境所有参数的影响方向一致,其中采用叶面积指数评价冠层结构动态的效果更佳;冠层开阔度和叶面积指数对林下光环境产生相反的影响,且均对散射光入射率影响程度最大.温性松栎混交林的林冠层整体较为均匀,林下光分布较为集中,林分树种组成与冠层结构对林下光照影响显著.     

From endogenous to exogenous pattern formation: Invasive plant species changes the spatial distribution of a native ant

Invasive species are a significant threat to global biodiversity, but our understanding of how invasive species impact native communities across space and time remains limited. Based on observations in an old field in Southeast Michigan spanning 35 years, our study documents significant impacts of habitat change, likely driven by the invasion of the shrub, Elaeagnus umbellata, on the nest distribution patterns and population demographics of a native ant species, Formica obscuripes. Landcover change in aerial photographs indicates that E. umbellata expanded aggressively, transforming a large proportion of the original open field into dense shrubland. By comparing the ant's landcover preferences before and after the invasion, we demonstrate that this species experienced a significant unfavorable change in its foraging areas. We also find that shrub landcover significantly moderates aggression between nests, suggesting nests are more related where there is more E. umbellata. This may represent a shift in reproductive strategy from queen flights, reported in the past, to asexual nest budding. Our results suggest that E. umbellata may affect the spatial distribution of F. obscuripes by shifting the drivers of nest pattern formation from an endogenous process (queen flights), which led to a uniform pattern, to a process that is both endogenous (nest budding) and exogenous (loss of preferred habitat), resulting in a significantly different clustered pattern. The number and sizes of F. obscuripes nests in our study site are projected to decrease in the next 40 years, although further study of this population's colony structures is needed to understand the extent of this decrease. Elaeagnus umbellata is a common invasive shrub, and similar impacts on native species might occur in its invasive range, or in areas with similar shrub invasions.     

Optical properties of juniper and lentisk canopies in a coastal Mediterranean macchia shrubland

Light absorbance and reflectance were measured on representative shrubs of two dominant shrub species (Pistacia lentiscus and Juniperus phoenicea) of coastal Mediterranean macchia ecosystems in Sardinia (Italy). An array of 64 calibrated photodiodes was adopted for both calculations of leaf area index (LAI) and canopy transmittance of photosynthetic active radiation (PAR). PAR and NIR reflectance of canopies having similar LAI allowed us to establish the particular optical properties of each species. Lentisk reflected twice as much as juniper in the 700-1,100 spectral region. Water indexes of the two species were also different in relation to leaf water content and red edge amplitude appeared to be related to chlorophyll content. Epidermis and mesophyll structures of both species are shown.     

伏牛山自然保护区森林冠层结构对林下植被特征的影响

在伏牛山自然保护区典型地段设立样方,测定了森林生态系统内几种典型群落类型的冠层结构、光环境特征,调查了林下植被的特征,分析了它们之间的相互关系.结果显示:各群落的冠层结构和光环境有一定的差异,单因素方差分析表明,部分群落间的差异性达到显著水平;各群落灌木层物种丰富度、多样性和均匀度均高于草本层,而优势度正相反;线性拟合的结果表明,草本层的物种丰富度、多样性与冠下光合量子通量密度间呈极显著负相关,优势度与冠下光合量子通量密度间呈显著正相关,灌木层各参数与冠层结构特征间相关性不显著.研究表明,冠层结构的变化对草本层(包括更新幼苗)的影响显著高于灌木层.林隙/林窗或林中空地的出现可能对草本物种或其他阳性及先锋物种具有促进作用,而对优势种幼苗的萌发和定植产生负效应.推测在典型的落叶阔叶林生态系统演替进程中,林下光照强度可能不是最主要的限制因素,优势种种子的扩散、萌发和定植限制可能更重要.     

Forest Canopy Properties and Variation in Aboveground Net Primary Production over Upper Great Lakes Landscapes

Relationships among aboveground net primary production (ANPP) and forest canopy properties were investigated in secondary successional forests of similar age and disturbance history in northern Lower Michigan, USA. Aboveground biomass, ANPP, canopy leaf area index (LAI), and several canopy nitrogen (N) measures were estimated from 12 stands representing major landform-level ecosystems and vegetation associations. Stand single-date and growing season average normalized difference vegetation indices (NDVI) were derived from Landsat TM. ANPP correlated most strongly with total canopy N content (r ² = 0.81, P < 0.001), followed by LAI (r ² = 0.73, P < 0.001) and area-based canopy-average leaf N concentration (r ² = 0.37, P < 0.05). No significant relationship was detected between ANPP and mass-based canopy-average leaf N concentration. Stand ANPP correlated positively with both total canopy N content (r ² = 0.62, P < 0.05) and mass-based leaf N concentration (r ² = 0.53, P < 0.05) of commonly dominant Populus spp. Relatively higher ANPP, total canopy N content and LAI corresponded to simultaneous presence of shade-intolerant P. grandidentata with shade-tolerant species. Both forms of NDVI were significantly related to ANPP, and more strongly to total canopy N content and LAI; relationships were stronger for seasonally averaged (r ² ≥ 0.75, P < 0.001) than for single-date NDVI (r ² ≥ 0.52, P < 0.01). Results indicate that on the transitioning study landscapes, ANPP was more closely related to canopy N content than to LAI, seasonally averaged NDVI was a more reliable predictor of ANPP and canopy properties than the single-date index, whereas measured canopy characteristics varied significantly between major landform-level ecosystems. The ongoing decline of P. grandidentata is likely to alter aboveground carbon and pools and fluxes in the course of succession.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

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.     

Microclimate and fluxes of water vapour, sensible heat and carbon dioxide in structurally differing subalpine plant communities in the Central Caucasus

Effects of canopy structure on microclimate, energy budget and CO₂ exchange were analysed in a pasture, two hay meadows, a tall herb community and a dwarf shrub community in the subalpine belt of the Central Caucasus. The results show that canopy structure exerts a marked influence on the distribution of photon flux density, temperature and canopy photosynthesis A_c. Three canopy types were distinguished. Type 1 (pasture) has a small LAI (leaf area index) and more than two-thirds of the phytomass is concentrated in the lowest few cm of the canopy, mainly as planophile leaves. This results in (1) a low degree of utilization of photosynthetic photon flux density (PPFD) by assimilatory plant components, (2) high leaf temperatures and a high soil heat flux during the phase of incoming radiation, and (3) a relatively low A_c/ LAI ratio. Type 2 (meadows), in spite of its erect leaves, which at high solar elevations permit light to penetrate to the lower canopy layers, is characterized by (1) marked effects of mutual shading in the lower canopy layers for most of the day, and thus (2) only slight variations in air and leaf temperatures and (3) a comparatively low A_c/LAI ratio. In canopies of type 3 (tall herb and dwarf shrub communities), there is a concentration of flat leaves in the upper layers. This results in (1) very good utilization of PPFD; (2) no strong fluctuations in canopy temperature as the flat leaves act as a buffer, reducing the amounts of incoming and outgoing radiation in lower canopy layers, and (3) high values of the A_c/LAI ratio. The energy budgets of the canopies investigated are governed not so much by their spatial structure, but rather indirectly by LAI and the degree of coupling of the canopy with the atmosphere.