Sinks for Plant Surplus Carbon Explain Several Ecological Phenomena

Plants engage in many processes and relationships that appear to be wasteful of the high-energy compounds that they produce through carbon fixation and photosynthesis. For example, living trees keep leafless tree stumps alive (i.e. respiring) and support shaded understory trees by sharing carbohydrates through root grafts or mycorrhizal fungal networks. Plants exude a variety of organic compounds from their roots and leaves, which support abundant rhizosphere and phyllosphere microbiomes. Some plants release substantial amounts of sugar via extra-floral nectaries, which enrich throughfall and alter lichen communities beneath the canopy. Large amounts of photosynthetically fixed carbon are transferred to root associates such as mycorrhizal fungi and N-fixing micro-organisms. In roots, some fixed C is respired through an alternative non-phosphorylating pathway that oxidizes excess sugar. Each of these processes is most prevalent when plants are growing under mild-to-moderate deficiencies or nutrients or water, or under high light or elevated atmospheric CO2. Under these conditions, plants produce more fixed carbon than they can use for primary metabolism and growth, and so have ‘surplus carbon’. To prevent cellular damage, these compounds must be transformed into other compounds or removed from the leaf. Each of the above phenomena represents a potential sink for these surplus carbohydrates. The fundamental ‘purpose’ of these phenomena may therefore be to alleviate the plant of surplus fixed C.

Assessing Forest Level Response to the Death of a Dominant Tree within a Premontane Tropical Rainforest

Small-scale treefall gaps are among the most important forms of forest disturbance in tropical forests. These gaps expose surrounding trees to more light, promoting rapid growth of understory plants. However, the effects of such small-scale disturbances on the distribution of plant water use across tree canopy levels are less known. To address this, we explored plant transpiration response to the death of a large emergent tree, Mortoniodendron anisophyllum Standl. & Steyerm (DBH > 220 cm; height ~40 m). Three suppressed, four mid-story, and two subdominant trees were selected within a 50 × 44 m premontane tropical forest plot at the Texas A&M Soltis Center for Research and Education located in Costa Rica. We compared water use rates of the selected trees before (2015) and after (2019) the tree gap using thermal dissipation sap flow sensors. Hemispherical photography indicated a 40% increase in gap fraction as a result of changes in canopy structure after the treefall gap. Micrometeorological differences (e.g., air temperature, relative humidity, and vapor pressure deficit (VPD)) could not explain the observed trends. Rather, light penetration, as measured by sensors within the canopy, increased significantly in 2019. One year after the tree fell, the water usage of trees across all canopy levels increased modestly (15%). Moreover, average water usage by understory trees increased by 36%, possibly as a result of the treefall gap, exceeding even that of overstory trees. These observations suggest the possible reallocation of water use between overstory and understory trees in response to the emergent tree death. With increasing global temperatures and shifting rainfall patterns increasing the likelihood of tree mortality in tropical forests, there is a greater need to enhance our understanding of treefall disturbances that have the potential to redistribute resources within forests.

Seven Ways a Warming Climate Can Kill the Southern Boreal Forest

The southern boreal forests of North America are susceptible to large changes in composition as temperate forests or grasslands may replace them as the climate warms. A number of mechanisms for this have been shown to occur in recent years: (1) Gradual replacement of boreal trees by temperate trees through gap dynamics; (2) Sudden replacement of boreal overstory trees after gradual understory invasion by temperate tree species; (3) Trophic cascades causing delayed invasion by temperate species, followed by moderately sudden change from boreal to temperate forest; (4) Wind and/or hail storms removing large swaths of boreal forest and suddenly releasing temperate understory trees; (4) Compound disturbances: wind and fire combination; (5) Long, warm summers and increased drought stress; (6) Insect infestation due to lack of extreme winter cold; (7) Phenological disturbance, due to early springs, that has the potential to kill enormous swaths of coniferous boreal forest within a few years. Although most models project gradual change from boreal forest to temperate forest or savanna, most of these mechanisms have the capability to transform large swaths (size range tens to millions of square kilometers) of boreal forest to other vegetation types during the 21st century. Therefore, many surprises are likely to occur in the southern boreal forest over the next century, with major impacts on forest productivity, ecosystem services, and wildlife habitat.

Phylogeny and Systematics of Crescentieae (Bignoniaceae), a Neotropical Clade of Cauliflorous and Bat-Pollinated Trees

Abstract—The tribe Crescentieae includes Amphitecna (21 species), Crescentia (six species), and Parmentiera (10 species), three genera of understory trees with a center of diversity in Central America and a small number of species in the Antilles and northern South America. Species in Crescentieae are united by their fleshy, indehiscent fruit and cauliflorous, bat-pollinated flowers. To lay a foundation for examining morphological, ecological, and biogeographic patterns within the tribe, we inferred the phylogeny for Crescentieae using both chloroplast (ndhF, trnL-F) and nuclear markers (PepC, ITS). The most recent circumscription of Crescentieae, containing Amphitecna, Crescentia, and Parmentiera is supported by our phylogenetic results. Likewise, the sister relationship between Crescentieae and the Antillean-endemic Spirotecoma is also corroborated by our findings. This relationship implies the evolution of fleshy and indehiscent fruits from dry and dehiscent ones, as well as the evolution of bat pollination from insect pollination. Fruits and seeds from species in Crescentieae are consumed by humans, ungulates, birds, and fish.

Arbuscular Mycorrhizal Community in Roots and Nitrogen Uptake Patterns of Understory Trees Beneath Ectomycorrhizal and Non-ectomycorrhizal Overstory Trees

Nitrogen (N) is an essential plant nutrient, and plants can take up N from several sources, including via mycorrhizal fungal associations. The N uptake patterns of understory plants may vary beneath different types of overstory trees, especially through the difference in their type of mycorrhizal association (arbuscular mycorrhizal, AM; or ectomycorrhizal, ECM), because soil mycorrhizal community and N availability differ beneath AM (non-ECM) and ECM overstory trees (e.g., relatively low nitrate content beneath ECM overstory trees). To test this hypothesis, we examined six co-existing AM-symbiotic understory tree species common beneath both AM-symbiotic black locust (non-ECM) and ECM-symbiotic oak trees of dryland forests in China. We measured AM fungal community composition of roots and natural abundance stable isotopic composition of N (δ15N) in plant leaves, roots, and soils. The root mycorrhizal community composition of understory trees did not significantly differ between beneath non-ECM and ECM overstory trees, although some OTUs more frequently appeared beneath non-ECM trees. Understory trees beneath non-ECM overstory trees had similar δ15N values in leaves and soil nitrate, suggesting that they took up most of their nitrogen as nitrate. Beneath ECM overstory trees, understory trees had consistently lower leaf than root δ15N, suggesting they depended on mycorrhizal fungi for N acquisition since mycorrhizal fungi transfer isotopically light N to host plants. Additionally, leaf N concentrations in the understory trees were lower beneath ECM than the non-ECM overstory trees. Our results show that, without large differences in root mycorrhizal community, the N uptake patterns of understory trees vary between beneath different overstory trees.

Integrating high resolution drone imagery and forest inventory to distinguish canopy and understory trees and quantify their contributions to forest structure and dynamics

Tree growth and survival differ strongly between canopy trees (those directly exposed to overhead light), and understory trees. However, the structural complexity of many tropical forests makes it difficult to determine canopy positions. The integration of remote sensing and ground-based data enables this determination and measurements of how canopy and understory trees differ in structure and dynamics. Here we analyzed 2 cm resolution RGB imagery collected by a Remotely Piloted Aircraft System (RPAS), also known as drone, together with two decades of bi-annual tree censuses for 2 ha of old growth forest in the Central Amazon. We delineated all crowns visible in the imagery and linked each crown to a tagged stem through field work. Canopy trees constituted 40% of the 1244 inventoried trees with diameter at breast height (DBH) > 10 cm, and accounted for ~70% of aboveground carbon stocks and wood productivity. The probability of being in the canopy increased logistically with tree diameter, passing through 50% at 23.5 cm DBH. Diameter growth was on average twice as large in canopy trees as in understory trees. Growth rates were unrelated to diameter in canopy trees and positively related to diameter in understory trees, consistent with the idea that light availability increases with diameter in the understory but not the canopy. The whole stand size distribution was best fit by a Weibull distribution, whereas the separate size distributions of understory trees or canopy trees > 25 cm DBH were equally well fit by exponential and Weibull distributions, consistent with mechanistic forest models. The identification and field mapping of crowns seen in a high resolution orthomosaic revealed new patterns in the structure and dynamics of trees of canopy vs. understory at this site, demonstrating the value of traditional tree censuses with drone remote sensing.

Forest Compositional Changes after a Decade of Emerald Ash Borer

Emerald ash borer is an invasive pest in North American forests. Ecological impacts of ash mortality from emerald ash borer are wide-ranging, including shifts in insect communities and wildlife behavior. Additionally, loss of ash from forests may have important implications regarding plant succession. Surveys of overstory, midstory, and understory trees within forests in northeastern Indiana, Lower Peninsula of Michigan, and northwestern Ohio were conducted to quantify the change in forest composition over a 10 year period. Interpolation of ash dominance illustrated inversion of live and dead ash values between 2007 and 2017. Even though more than 83% of overstory live ash basal area was lost across the study area, green ash was the most abundant midstory and understory species representing regeneration. Additionally, loss of ash from many of the sites resulted in compositional changes that were greater than merely the subtraction of ash. Due to the relatively large number of forest types with which ash species are associated, loss of ash will have broad ecological consequences, including on community composition.

Comparison of Allometric Equation and Destructive Measurement of Carbon Storage of Naturally Regenerated Understory in a Pinus rigida Plantation in South Korea

The forest understory plays an important role in the carbon and nutrient cycling and forest stability, but cost-efficient quantification of its biomass remains challenging. Most of the existing biomass allometric equations have been developed and designed only for mature forest trees (i.e., Diameter at breast height (DBH) ≥ 10 cm), and those for trees with DBH less than 10 cm are not readily available. In this study, we compared the biomass by plant component (i.e., foliage, branch, and stem) measured by a destructive method with those estimated by the existing biomass allometric equations for understory trees with DBH less than 10 cm in a Pinus rigida plantation. We also developed an allometric biomass equation for the identified understory tree species, namely, Quercus variabilis, Quercus acutissima, Quercus mongolica, Quercus serrata, and Carpinus laxiflora. The estimated biomass using allometric equations for foliage, branch, and stem was lower than the values obtained using the destructive method by 64%, 41%, and 18%, respectively. The biomass allometric equations developed in this study showed high coefficients of determination (mean R2 = 0.970) but varied depending on species and tree part (range: 0.824–0.984 for foliage, 0.825–0.952 for branch, and 0.884–0.999 for the stem, respectively). The computed biomass of the understory vegetation was 22.9 Mg ha−1, representing 12.0% of the total biomass of the P. rigida plantation. The present study demonstrates that understory trees with DBH less than 10 cm account for a considerable portion of carbon stock in forest ecosystems, and therefore suggests that more biomass allometric equations should be optimized for small-DBH trees to improve forest carbon stock estimation.

A Multi-Threshold Segmentation for Tree-Level Parameter Extraction in a Deciduous Forest Using Small-Footprint Airborne LiDAR Data

The development of new approaches to tree-level parameter extraction for forest resource inventory and management is an important area of ongoing research, which puts forward high requirements for the capabilities of single-tree segmentation and detection methods. Conventional methods implement segmenting routine with same resolution threshold for overstory and understory, ignoring that their lidar point densities are different, which leads to over-segmentation of the understory trees. To improve the segmentation accuracy of understory trees, this paper presents a multi-threshold segmentation approach for tree-level parameter extraction using small-footprint airborne LiDAR (Light Detection And Ranging) data. First, the point clouds are pre-processed and encoded to canopy layers according to the lidar return number, and multi-threshold segmentation using DSM-based (Digital Surface Model) method is implemented for each layer; tree segments are then combined across layers by merging criteria. Finally, individual trees are delineated, and tree parameters are extracted. The novelty of this method lies in its application of multi-resolution threshold segmentation strategy according to the variation of LiDAR point density in different canopy layers. We applied this approach to 271 permanent sample plots of the University of Kentucky’s Robinson Forest, a deciduous canopy-closed forest with complex terrain and vegetation conditions. Experimental results show that a combination of multi-resolution threshold segmentation based on stratification and cross-layer tree segments merging method can provide a significant performance improvement in individual tree-level forest measurement. Compared with DSM-based method, the proposed multi-threshold segmentation approach strongly improved the average detection rate (from 52.3% to 73.4%) and average overall accuracy (from 65.2% to 76.9%) for understory trees. The overall accuracy increased from 75.1% to 82.6% for all trees, with an increase of the coefficient of determination R2 by 20 percentage points. The improvement of tree detection method brings the estimation of structural parameters for single trees up to an accuracy level: For tree height, R2 increased by 5.0 percentage points from 90% to 95%; and for tree location, the mean difference decreased by 23 cm from 105 cm to 82 cm.