Variation in individual biomass decreases faster than mean biomass with increasing density of bamboo stands

The total biomass of a stand is an indicator of stand productivity and is closely related to the density of plants. According to the self-thinning law, mean individual biomass follows a negative power law with plant density. If the variance of individual biomass is constant, we can expect increased stand productivity with increasing plant density. However, Taylor's power law(TPL) that relates the variance and the mean of many biological measures(e.g. bilateral areal differences of a leaf, plant biomass atdifferent times, developmental rates at different temperatures, population densities on different spatial or temporal scales), affects the estimate of stand productivity when it is defined as the total biomass of large plants in a stand.Because the variance of individual biomass decreases faster than mean individual biomass, differences in individual biomass decline with increasing density, leading to more homogeneous timbers of greater economic value. We tested whether TPL in plant biomass holds for different species and whether the variance of individual biomass changes faster than the mean with increasing stand density.The height, ground diameter and fresh weight of 50 bamboo species were measured in 50 stands ranging from 1 m by 1 m to 30 m by 30 m to ensure more than 150 bamboos in every stand. We separately examined TPL in height,ground diameter, and weight, and found that TPL holds for all three biological measures, with the relationship strongest for weight. Using analysis of covariance to compare the regression slopes of logarithmic mean and variance against the logarithm of density, we found that the variance in individual biomass declined faster than the mean with increasing density. This suggests that dense planting reduced mean individual biomass but homogenized individual biomass. Thus, there exists a trade-off between effective stand productivity and stand density for optimal forest management. Sparse planting leads to large variation in individual biomass, whereas dense planting reduces mean individual biomass. Consequently, stand density for a plantation should be set based on this trade-off with reference to market demands.     

Species-specific allometric equations for improving aboveground biomass estimates of dry deciduous woodland ecosystems

Allometric equations are important for quantifying biomass and carbon storage in terrestrial forest ecosystems.However,equations for dry deciduous woodland ecosystems,an important carbon sink in the lowland areas of Ethiopia have not as yet been developed.This study attempts to develop and evaluate species-specific allometric equations for predicting aboveground biomass(AGB)of dominant woody species based on data from destructive sampling for Combretum collinum,Combretum molle,Combretum harotomannianum,Terminalia laxiflora and mixed-species.Diameter at breast height ranged from 5 to 30 cm.Two empirical equations were developed using DBH(Eq.1)and height(Eq.2).Equation 2 gave better AGB estimations than Eq.1.The inclusion of both DBH and H were the best estimate biometric variables for AGB.Further,the equations were evaluated and compared with common generic allometric equations.The result showed that our allometric equations are appropriate for estimating AGB.The development and application of empirical species-specific allometric equations is crucial to improve biomass and carbon stock estimation for dry woodland ecosystems.     

Grazing by wild giant pandas does not affect the regeneration of Arundinaria spanostachya

Arundinaria spanostachya is the main bamboo species on which giant pandas forage.Information on its regeneration capacity to meet the foraging demands of pandas is important to ensure the long-term survival of pandas.In 2014,2 plot types(grazed versus control)were selected in the winter habitat used by one wild giant panda.Various morphological and regeneration characteristics of the bamboo were assessed in the respective plots during 2014 and 2015.The panda avoided feeding on bamboo with a basal diameter of<6 mm,preferentially feeding on intermediate-sized bamboo.The density of new shoots and the recruitment rate of shoots were significantly higher in the grazed plots compared to the control plots.The annual recruitment rate of shoots was markedly higher than the annual mortality rate in the same plot type.After grazing,the basal diameter and height of new bamboo were similar between the 2 plots.In the grazed plots,the basal diameter of new bamboo was similar to that of the stumps.Innutrition and foraging by insects were the main factors influencing the survival of new shoots.Our results showed that grazing by giant pandas helped the clonal regeneration of A.spanostachya populations,with the basal diameter of new bamboo shoots meeting the foraging demand of pandas.However,information on the carrying capacity of A.spanostachya is necessary to ensure sufficient forage matter is available for reintroduced giant pandas.Furthermore,fertilization and insect control measures should be implemented during the shooting period to optimize A.spanostachya resources.     

The coppice-with-standards silvicultural system as applied to Eucalyptus plantations — a review

We review the management of Eucalyptus species under a coppice-with-standards （CWS） silvicultural system. CWS management results in product diversification, permitting production of small and large scale timber from the same stand. Eucalyptus species are suitable candidates for CWS management because： there are large worldwide plantation areas, sprouting capacity is high, and eucalypts are multipur- pose species. We discuss （1） short rotation Eucalyptus coppice manage- ment for energy and pulping and （2） Eucalyptus seedling management for solid wood products. We review the literature and discuss experi- ences with Eucalyptus managed under the CWS system. We also assess projects dealing with Eucalyptus coppice management, stand density regulation, pruning, and stand and wood quality. The growth environ- ment of the standard trees （heavy competition up to the first harvest, free growth afterwards） coupled with long rotations （〉20 years） results in high quality logs for solid wood products. Early pruning should be ap- plied to enhance wood quality. We propose a system for the silvicultural management of Eucalyptus under the CWS system, elaborating on the consequences of initial planting density, site productivity, and standard tree densities as well as timing of basic silvicultural applications.     

Carbon stock and rate of carbon sequestration in Dipterocarpus forests of Manipur,Northeast India

We examined the carbon stock and rate of carbon sequestration in a tropical deciduous forest dominated by Dipterocarpus tuberculatus in Manipur,North East India.Estimation of aboveground biomass was determined by harvest method and multiplied with density of tree species.The aboveground biomass was between18.27–21.922 t ha-1and the carbon stock ranged from9.13 to 10.96 t C ha-1across forest stands.Aboveground biomass and carbon stock increased with the increase in tree girth.The rate of carbon sequestration varied from1.4722 to 4.64136 t ha-1year-1among the dominant tree species in forest stands in tropical deciduous forest area.The rate of carbon sequestration depends on species composition,the density of large trees in different girth classes,and anthropogenic disturbances in the present forest ecosystem.Further work is required to identify tree species having the highest potential to sequester CO2 from the atmosphere,which could lead to recommendations for tree plantations in a degraded ecosystem.     

Allometric relationship for estimating above-ground biomass of Aegialitis rotundifolia Roxb. of Sundarbans mangrove forest, in Bangladesh

Tree biomass plays a key role in sustainable management by providing different aspects of ecosystem. Estimation of above ground biomass by non-destructive means requires the development of allometric equations. Most researchers used DBH (diameter at breast height) and TH (total height) to develop allometric equation for a tree. Very few species-specific allometric equations are currently available for shrubs to estimate of biomass from measured plant attributes. Therefore, we used some of readily measurable variables to develop allometric equations such as girth at collar-height (GCH) and height of girth measuring point (GMH) with total height (TH) for A. rotundifolia, a mangrove species of Sundarbans of Bangladesh, as it is too dwarf to take DBH and too irregular in base to take Girth at a fixed height. Linear, non-linear and logarithmic regression techniques were tried to determine the best regression model to estimate the above-ground biomass of stem, branch and leaf. A total of 186 regression equations were generated from the combination of independent variables. Best fit regression equations were determined by examining co-efficient of determination (R2), co-efficient of variation (CV), mean-square of the error (MSerror), residual mean error (Rsme), and F-value. Multiple linear regression models showed more efficient over other types of regression equation. The performance of regression equations was increased by inclusion of GMH as an independent variable along with total height and GCH.     

Seasonal variations in species diversity, dry matter and net primary productivity of herb layer of Quercus leucotrichophora-Pinus roxburghii mixed forest in Kumaun Himalaya, India

Plant biomass, species diversity and net primary productivity are presented for herb layer of banj oak (Quercus leucotrichophora A. Camus)-chir pine (Pinus roxburghii Sarg.) mixed forest in Kumaun, central Himalaya, India. The species diversity declined from a maximum (3.56) in September to a minimum (2.11) in December. The monthly live shoots biomass exhibited a single peak growth pattern with highest live shoot biomass of 185 g·m-2 in August. The seasonal pattern showed that the maximum above-ground production (131 g·m-2) occurred during the rainy season and the minimum (1 g·m-2) during winter season. The below-ground production was maximum during winter season (84 g·m-2) and minimum during summer season (34 g·m-2). The annual net shoot production was 171 g·m-2 and total below-ground production was 165 g·m-2. Of the total input 61% was channeled to above-ground parts and 39% to below-ground parts. Transfer of live shoots to dead shoots compartments and that of dead shoots to litter compartments was 61% and 66%, respectively. The total dry matter disappearance was 61% of the total input within annual cycle. The herb layer showed a net accumulation of organic matter, indicating the seral nature of the community.     

Conservation of an important montane bamboo Thamnocalamus falconeri,Hook.f.ex Munro through axillary bud proliferation

Thamnocalamus falconeri, Hook.f. ex Munro.,an important bamboo species belonging to the family Poaceae, locally known as Ringal, occurs in the hills of Uttarakhand, India. This species has been traditionally exploited by local communities to support their livelihoods.Increasing needs of the hill villages impose unsustainable pressure on natural stands of Ringal in the Uttarakhand hills and forests have been degraded. The long history of excessive cutting of Ringal from natural forests and the lack of replanting threaten villager livelihoods. Replanting is required to conserve the species. We propose a protocol for generation of planting material through axillary bud proliferation for multiplication and conservation of this species. We collected offsets/rhizomes from a natural stand of T. falconeri in the Chopta Mandal areas(Chamoli district, India). These were planted at sites of varied elevation and fresh single nodal segments were collected from them as explants. Different sterilization treatments were assessed to combat contamination. Among these, treatment of 0.1 %Hg Cl2 followed by 5 % Na OCl, proved best. Among two cytokinin treatments, viz. BAP and Kinetin, singly or in combination, BAP alone(5 mg L-1) proved superior and resulted in 100 % bud break. BAP-supplemented MS media yielded maximum vigorous shoot formation(90 %)and maximum number of shoots(8.9). Subculturing of shoots on the same medium with similar BAP treatment(5 mg L-1BAP) enabled continuous production of healthy shoots at similar frequency. Maximum rooting(100 %)was recorded on half-strength MS medium supplemented with 5 mg L-1IBA. Micropropagated plants were hardened and acclimatized in soil mixture(2:1:1) and then transplanted to field sites(Magra, Uttarakhand, 1,834 m).Eight to ten months after field transplantation we recorded100 % survival of transplanted material. This micropropagation protocol could be used successfully for raising a stock of genetically homogenous plant material in bulk for field plantations and for conservation of the species.     

Aboveground biomass and carbon stock in the largest sacred grove of Manipur,Northeast India

Aboveground biomass and carbon stock in the largest sacred grove of Manipur was estimated for trees with diameter [10 cm at 1.37 m height.The aboveground biomass,carbon stock,tree density and basal area of the sacred grove ranged from 962.94 to 1130.79 Mg ha~(-1),481.47 to 565.40 Mg ha~(-1) C,1240 to 1320 stem ha~(-1) and79.43 to 90.64 m~2 ha~(-1),respectively.Trees in diameter class of 30–40 cm contributed the highest proportion of aboveground biomass(22.50–33.73%).The aboveground biomass and carbon stock in research area were higher than reported for many tropical and temperate forests,suggesting a role of spiritual forest conservation for carbon sink management.     

Fuel properties and combustion characteristics of some promising bamboo species in India

We investigated the fuel characteristics of five important bamboo species viz., Dendrocalamus strictus, D. brandisii, D. stocksii, Bambusa bambos and B. balcooa. The selected species cover more than 85% of the total growing stock of bamboo in India. Basic density varied from 0.48 to 0.78 g?cm-3 among the bamboo species studied. Ash content, volatile matter content and fixed carbon content ranged between 1.4%-3.0%, 77.2%-80.8% and 17.6%-21.1%, respectively. Variation in calorific value(18.7-19.6 MJ?kg-1) was marginal. Fuel value index varied widely(586-2120) among bamboo species. The highest calorific value(19.6 MJ?kg-1) and fuel value index(2120) were found in B. balcooa. Ash elemental analysis revealed that silica and potassium are the major ash forming minerals in bamboo biomass. Silica content ranged from 8.7% to 49.0%, while potassium ranged from 20.6% to 69.8%. We studied combustion characteristics under oxidizing atmosphere. Burning profiles of the samples were derived by applying the derivative thermogravimetric technique which is discussed in detail. The five bamboo species were different in their combustion behaviour, mainly due to differences in physical and chemical properties. We compare fuel properties, ash elemental analysis and combustion characteristics of bamboo biomass with wood biomass of Eucalyptus hybrid(Eucalyptus tereticornis × Eucalyptus camaldulensis).     

Patterns of tree buttressing at Lawachara National Park, Bangladesh

We describe patterns of buttress formation and development in eleven tree species at Lawachara National Park, Bangladesh. Fortyfive percent of trees of these 11 species had buttresses. Artocarpus chaplasha Roxb. showed maximum (87%) buttress formation, whereas Alstonia scholaris (L.) R. Br. did not show any buttress. Buttresses were recorded in 20% 40% of trees of six species and 40% 60% of trees in three species. Mean length and height of buttress varied among the species and ranged from 0.37 1.37 m and 0.71 2.13 m, respectively. Buttress height, mean buttress length, total buttress length, and total length pluslength of secondaries increased with DBH (diameter at breast height) and tree height. Buttress number did not increase with DBH or tree height.Under-storey and mid-canopy trees produced less developed buttresses than did emergent trees (p<0.01). Wood density showed moderate effects on buttress characters (p<0.05), while the slope of the land did not. Canopy category was a primary regulating factor for tree buttressing, suggesting that buttresses are mechanical adaptations of trees to counter physical stresses.     

Estimated biomass carbon in thinned Cunninghamia lanceolate plantations at different stand-ages

Chinese fir(Cunninghamia lanceolate [Lamb.]Hook.) is a fast-growing species which is not only important as a timber-supplier,but also as an available sink for carbon(C) storage in biomass.Stand age and density are two critical factors that can determine tree C sequestration as interrelated drivers through natural self-thinning.C.lanceolate were planted using 1-year-old bare-root seedlings at the initial density of 1800 stems ha~(-1) in a 15-ha montane area of Hunan Province,China in 1987.The plantation was thinned twice 10 and 20 years after planting to leave trees of437.5±26.6,675.0±155.2 and 895.8±60.1 stems ha~(-1) as low,medium,and high densities,respectively.Tree height and diameter at breast height(DBH) were measured every2 years beginning from 23 years(2009) to 31 years(2018)after establishment,timber volume(TV) and biomass C were estimated accordingly.We did not find any interactive effect of age and density on any variables except for height.Both TV and biomass C increased with stand age or decreased in higher densities.The allometric heightDBH relationship can be fitted by an exponential risingto-maximum model with higher maximum value over time.The decline of biomass C along density fit with the inverse first-order polynomial model which indicated that at least1300-1500 stems ha~(-1) may be needed to maximize TV and biomass C for a longer term over 20 years.Therefore,to control the density to a reasonable level,over 1300 stems ha~(-1) in a rotation over 20 years old will be practical for tree biomass C in Chinese fir plantations.     

Biomass production of hybrid aspen growing on former farm land in Sweden

We construct dry weight equations for hybrid aspen growing on former farmland in Sweden. Dry weight equations for fractions of hybrid aspen trees were also made. We estimated biomass production in 24 stands. The stands were located in Sweden at latitudes ranging from 55 to 60o N. The mean age was 18 years (range 15-23), the mean stand density 1090 stems·ha-1 (range 378 2374), and the mean diameter at breast height (over bark) 178 mm (range 85 244 mm). Soil types in the hybrid aspen stands were mainly clay (21 stands), tills (2 stands) and other (1 stand). The mean total standing dry weight above stump level (≈ 200 mm) for the hybrid aspen stands was 135±53 t·ha-1 with a range of 42 219 t·ha-1 . In addition to estimating conventional dry weights of trees and tree components, basic density, specific leaf area (SLA), projected leaf area (PLA) and leaf area index (LAI) were estimated and were in agreement with published figures.     

Yield-density effects on growth and biomass partitioning in Leucaena leucocephala seedlings

Experiments were conducted to study the effects of density on growth and biomass partitioning of Leucaena leucocephala seedlings.Four plantations with densities of 10,000,20,000,40,000,and 80,000 seedlings ha^-1 were evaluated only from 15 to 25 months after planting.At 15 months,crown height and width decreased with increasing density.Seedling height/dbh ratios increased with increasing density.Biomass increased with greater density according to the yield–density effect equation,which was evident for all densities.With increasing age,biomass division to branches and leaves increased,whereas partitioning to roots decreased in the 10,000 and 20,000 seedlings ha-1 plantings.Partitioning to branches and leaves remained relatively steady,while partitioning to roots increased in the 40,000 and 80,000 seedlings ha^-1 plantings.Biomass division into stem and bark components remained relatively steady in all densities.Yield–density and organ yield–density curves shifted upward with increasing seedling age on a log–log graph throughout the experimental period.     

Longitudinal growth strains in five clones of Eucalyptus tereticornis Sm.

We studied the variability in longitudinal growth strains and wood basic density in five-year old trees from five clones (one tree per clone) of Eucalyptus tereticornis. Mean longitudinal growth strain in clones ranged from 466 to 876 μm. There was a significant difference between clones in growth strains and wood basic density. Clone 10 exhibited maximum growth strains and basic density, whereas clone 3 and clone 7 exhibited minimum growth strains and basic density, respectively. Within a tree, the growth strain variation with tree height was high but statistically insignificant while within tree variation in basic density was very small. There was no specific trend in variation in either strain or density within a tree. There was 5% 200% difference in growth strain on opposite sides of the logs. However two strains showed a strong positive correlation. There was a moderate positive association of wood basic density and mean growth strains in logs. The variation around the periphery emphasize the need to measure strain more than one, preferably on opposite sides at the same height, on a tree to know the mean strain level for the purpose of selection of clones.     

Effect of auxins on axillary and de novo shoot regeneration from in vitro shoot cultures derived from forced epicormic buds of teak (Tectona grandis L.)

In this presentation,we report on de novo and axillary shoot regeneration and rooting of shoots maintained over a long term,from cultures of Tectona grandis L.Shoot-tips of teak shoots forced from epicormic buds were used as the starting material for axenic shoot-culture establishment.Long term maintenance of such axenic shoot cultures was carried out by regular sub-culturing on MS media supplemented with N 6-benzyleadenine (BA,8.8 μmol L-1) and indole-3-butyric acid (IBA,2 μmol L-1) for 24 months.Vigorously growing shoot tips (2 3 cm long) were inoculated on the MS basal medium supplemented with different concentrations (0,1,2,4,6,8 or 10 μmol L-1) of either IBA or α-naphthaleneacetic acid (NAA) for rooting.Axillary and de novo shoots were developed from axillary and cut basal ends of shoots,respectively.Shoots growing on auxins were further sub-cultured (every 15 days) and maintained for 45 days.The greatest number of de novo (5.06) as well as axillary shoots (2.85) was observed on the MS medium supplemented with 10 μmol L-1 NAA or 8 μmol L-1 IBA,respectively,after 45 days.The combinations of both IBA (μmol L-1) + NAA (μmol L-1) were tested at different concentrations (4 + 4,6 + 6,8 + 8) supplemented to a half strength MS basal medium with 0.1% activated charcoal for rooting of decapitated and non-decapitated de novo and axillary shoots.Rooting from non-decapitated de novo shoots was highest (93.33%) with a mean number of roots of 4.61 on this medium,supplemented with 6 μmol L-1 IBA + 6 μmol L-1 NAA,after 36 days of initial culture.Individual auxin,however,was not effective for root induction.Rooted shoots were acclimatized in a green house and after four weeks plantlets were transferred to the field.     

Modeling of the height–diameter relationship using an allometric equation model:a case study of stands of Phyllostachys edulis

Understanding the relationship between tree height(H) and diameter at breast height(D) is vital to forest design, monitoring and biomass estimation. We developed an allometric equation model and tested its applicability for unevenly aged stands of moso bamboo forest at a regional scale. Field data were collected for 21 plots. Based on these data, we identified two strong power relationships: a correlation between the mean bamboo height(Hm) and the upper mean H(Hu), and a correlation between the mean D(Dm) and the upper mean D(Du). Simulation results derived from the allometric equation model were in good agreement with observed culms derived from the field data for the 21 stands,with a root-mean-square error and relative root-mean-square error of 1.40 m and 13.41 %, respectively. These results demonstrate that the allometric equation model had a strong predictive power in the unevenly aged stands at a regional scale. In addition, the estimated average height–diameter(H–D) model for South Anhui Province was used to predict H for the same type of bamboo in Hunan Province based on the measured D, and the results were highly similar. The allometric equation model has multiple uses at the regional scale, including the evaluation of the variation in the H–D relationship among regions. The model describes the average H–D relationship without considering the effects caused by variation in site conditions, tree density and other factors.     

Influences of antibiotics on plantlet regeneration via organogenesis in Ioblolly pine （Pinus taeda L.）

Three antibiotics ampicillin, carbenicillin, and cefotaxime were evaluated for their effects on induction, growth, and differentiation of organogenic calli, as well as rooting of regenerated shoots of three Ioblolly pine (Pinus taeda L.) genotypes. Of the antibiotics administered, cefotaxime maximally increased the frequency of callus formation and growth rate of organogeni ccalli, carbenicillin maximally increased the frequency of shoot regeneration and the average number of adventitious shoots per piece of organogenic callus, ampicUlin maximally decreased the rooting frequency of regenerated shoots and mean number of roots per regenerated shoot, in comparison with antibiotic-free media. Compared with the control, ampicillin minimally increased the frequency of callus formation, cefotaxime minimally increased the frequency of shoot regeneration, and carbenicillin minimally decreased the rooting frequency of regenerated shoots in three Ioblolly pine genotypes tested. All three antibiotics increased the frequencies of callus formation and shoot regeneration, and reduced the rooting frequency ot regenerated shoots suggested that the establishment of an efficient Agrobacterium tumefaciens-mediated transformation protocol for stable integration of foreign genes into Ioblolly pine need to select a suitable antibiotic. This investigation could be useful for optimizing genetic transformation of conifers.     

Changes in gas exchange,root growth,and biomass accumulation of Platycladus orientalis seedlings colonized by Serendipita indica

Serendipita indica(formerly known as Piriformospora indica),a root endophytic fungus,exhibits multiple functions in some agricultural,horticultural,and medicinal plant species.We studied colonization of the roots of Platycladus orientalis,a forest tree species,by S.indica to improve the quality of the seedlings in seedbeds and survival rates in sylviculture.At 20 days after inoculation,S.indica colonized the root cortex of P.orientalis seedlings.Root colonization by S.indica significantly increased net CO2 assimilation,light use efficiency,and biomass accumulation by both roots and shoots,whereas it did not affect the biomass allocation between roots and shoots.In addition,the symbiosis significantly increased root total length,surface area,and volume.In view of the two specific traits of S.indica,i.e.,axenic culture and wide colonization in plants,the fungus might be used for improving quality of P.orientalis seedlings and increasing their survival after transplanting.     

Assessing current stocks and future sequestration potential of forest biomass carbon in Daqing Mountain Nature Reserve of Inner Mongolia,China

Assessment of regional forest carbon stocks and underlying controls is critical for guiding forest management in the context of carbon sequestration. We investigated the variations in tree biomass carbon stocks relating to forest types, and estimated the total tree biomass carbon stocks and projected gains through natural stand development by 2020 and 2050 in the Daqing Mountain Nature Reserve based on Category II data of the Forest Inventory of Inner Mongolia for the period ending 2008. Over a total area of 388,577 ha,this nature reserve currently stores an estimated 2221 Gg C in tree aboveground biomass alone, with potential to grow by more than 30 % to reach 2938 Gg C by 2020 and nearly double to 4092 Gg C by 2050 through natural development of the existing forest stands. The tree biomass carbon density and potential gain in tree biomass carbon stocks vary markedly among forest types and with stand development.The variations in the potential change of tree biomass carbon density for the periods 2008–2020 and 2008–2050 among forest types partly reflect the varying relationships of tree biomass carbon density with stand age for different tree species, and partly are attributable to variations in the stand age structure among different forest types. Of the major forest types, the ranking of projected changes in tree biomass carbon density are not consistent with variations in the relationship between tree biomass carbon density and stand age, neither are they explainable by variations in stand age structures, implying the interactive effect between forest type and stand dynamics on temporal changes in tree biomass carbon density. Birch rank highest for future biomass carbon sequestration because of its dominance in cover area and better age structure for potential gain in tree biomass carbon stocks. Poplar and larch were out-performers compared to other forest types given their greater contribution to total tree biomass carbon stocks relative to their distributional areas. Findings in this study illustrate that protection and proper management of under-aged forests can deliver marked gains in biomass carbon sequestration. This is of great importance to policy-makers as well as to scientific communities in seeking effective solutions for adaptive forest management and mitigation of anthropogenic greenhouse gases emissions using forest ecosystems.