• <tr id="yyy80"></tr>
  • <sup id="yyy80"></sup>
  • <tfoot id="yyy80"><noscript id="yyy80"></noscript></tfoot>
  • 99热精品在线国产_美女午夜性视频免费_国产精品国产高清国产av_av欧美777_自拍偷自拍亚洲精品老妇_亚洲熟女精品中文字幕_www日本黄色视频网_国产精品野战在线观看 ?

    Predicting the provisioning potential of forest ecosystem servicesusing airborne laser scanning data and forest resource maps

    2018-09-12 09:14:54JariVauhkonen
    Forest Ecosystems 2018年3期

    Jari Vauhkonen

    Abstract Background:Remote sensing-based mapping of forest Ecosystem Service(ES)indicators has become increasingly popular.The resulting maps may enable to spatially assess the provisioning potential of ESs and prioritize the land use in subsequent decision analyses.However,the mapping is often based on readily available data,such as land cover maps and other publicly available databases,and ignoring the related uncertainties.Methods:Thisstudy tested the potential to improve the robustnessof the decisionsby meansof local model fitting and uncertainty analysis.The quality of forest land use prioritization wasevaluated under two different decision support models:either using the developed modelsdeterministically or in corporation with the uncertaintiesof the models.Results:Prediction modelsbased on Airborne Laser Scanning(ALS)dataexplained the variation in proxiesof the suitability of forest plotsfor maintaining biodiversity,producing timber,storing carbon,or providing recreational uses(berry picking and visual amenity)with RMSEsof 15%–30%,depending on the ES.The RMSEsof the ALS-based predictionswere 47%–97%of those derived from forest resource mapswith a similar resolution.Due to applying a similar field calibration step on both of the data sources,the difference can be attributed to the better ability of ALSto explain the variation in the ESproxies.Conclusions:Despite the different accuracies,proxy valuespredicted by both the datasourcescould be used for a pixel-based prioritization of land use at a resolution of 250 m2,i.e.,in aconsiderably more detailed scale than required by current operational forest management.The uncertainty analysis indicated that maps of the ESprovisioning potential should be prepared separately based on expected and extreme outcomesof the ESproxy models to fully describe the production possibilities of the landscape under the uncertainties in the models.

    Keywords:Forestry decision making,Spatial prioritization,Light detection and ranging(LiDAR),Remote sensing

    Background

    Forestry decision making requires evaluating potential management alternatives with respect to multiple objectives(Kangas et al.2008).A fundamental decision is related to which goods and services to produce:in addition to conventional timber production,the management objectives may be related to maintaining habitats,providing recreational and aesthetic opportunities,and carbon storage or sequestration(e.g.Pukkala 2016).These goods and services are jointly called“multiple uses”(Kangas 1992)or,following Costanza et al.(1997),Daily et al.(1997)and many others,“ecosystem services”of forest. In the following text, I use ESs to abbreviate“Ecosystem Services”, referring most essentially to indicators of forest-related ESs that can be derived from Remote Sensing (RS) or other digital map data as indirect proxies (Andrew et al. 2014). The mapping of these proxies allows spatial prioritization and other spatially explicit analyses of multiple ESs at various scales (e.g. Schr?ter et al. 2014; R?s?nen et al. 2015; Sani et al. 2016; Roces-Díaz et al. 2017). According to reviews(Martínez-Harms and Balvanera 2012; Englund et al.2017) and a collection of case studies (Barredo et al.2015), however, such analyses can be expected to suffer from the lack of standardized terminology, methodology and data. Increased attention should especially be focused on quantifying and communicating the resulting uncertainties to the decision makers in order to make informed decisions(see also Eigenbrod et al.2010;Schulp et al.2014;Foody 2015).Accounting for these aspects,the present study examines the robustness of forest land-use prioritization based on maps of the provisioning potential of forest ESs(Vauhkonen and Ruotsalainen 2017a),i.e.,the fitness of forest patches to provide goods and services typical to the ESs occurring in the studied area,re-considering the methodological and data workflow proposed in the earlier study.

    To result in valid conclusions from RS-based decision analyses,the estimates should be accurate already at the level of individual pixels.The use of active RS such as Light Detection and Ranging(LiDAR)is expected to produce more accurate information compared to passive,optical RS(Lefsky et al.2001;Coops et al.2004;Maltamo et al.2006),especially,when using small pixels(e.g.,200 m2as in N?sset 2002).Forest structure and habitat related inventories in particular benefit from the ability of LiDAR to provide three-dimensional information,when operated as Airborne Laser Scanning(ALS;Maltamo et al.2014).Kankare et al.(2015)evaluated the estimation accuracy of biomass attributes based on two different RSsetups in an area closely resembling to that presently studied.According to their results,pixel-level predictions based on coarse to medium resolution satellite imagery had a Root Mean Squared Error(RMSE)of 47.7%of the total biomass,which could be reduced to 25.7%using ALSand local field reference data.The ALS data used were acquired by the land survey,and the availability of such data is increasing due to large-area acquisitions for terrain elevation modelling.Such data have also been used to map attributes related to habitat(Melin et al.2013,2016;Vauhkonen and Imponen 2016),structural(Valbuena et al.2016b;Vauhkonen and Imponen 2016)and aesthetic(Vauhkonen and Ruotsalainen 2017b)propertiesof the forest.

    Overall,when various forest ESs are categorized according to a typology such as the Common International Classification of Ecosystem Services(CICES)as in Englund et al.(2017),the potential of ALSfor assessing the suitability of forest areas to provide these ESs can be characterized as:

    –Regulation and maintenance services:A very high number of studies indicates that the vegetation height and density profiles produced by ALSare useful for a detailed quantification of variations in above-ground biomass(N?sset and Gobakken 2008;Zolkos et al.2013;Popescu and Hauglin 2014)and,thus,carbon storage(Patenaude et al.2004).Essentially,ALSproduces a three-dimensional description of the forest structure,which can be related to ecological properties such as habitat types(B?ssler et al.2011)or biological diversity in general(Müller and Vierling 2014)and employed to assess suitability of forests to be maintained as habitats for different species(Davies and Asner 2014;Hill et al.2014;Simonson et al.2014).

    –Provisioning services:Several studies carried out especially in boreal forest structures indicate ALS data useful for assessing properties related to wood production.Except that the methods listed in the previous paragraphs can be directly used to assess the production potential of bulk biomass,also more detailed predictions of timber assortments(Korhonen et al.2008;Kotamaa et al.2010;Vauhkonen et al.2014;Hou et al.2016)or wood fiber-related attributes(Hilker et al.2013;Luther et al.2014)are possible.Although the yield studies are mostly related to wood-based biomass,there also are examples of improved assessments of the yield of shrub fruits(Barber et al.2016)or edible fungi(Peura et al.2016)based on ALS.

    –Cultural services:The applicability of ALShighly depends on the cultural service of interest.For example,several archaeological studies indicate the potential to improve the mapping of historical remains in the forest using an ALS-based digital terrain model.Similar techniques to visualize the terrain(Domingo-Santos et al.2011)or trees(L?m?s et al.2015)could potentially be used to assess the aesthetic properties of the forest.To date,the study of Vauhkonen and Ruotsalainen(2017b),which assessed the preferences on the visual amenity of a forest area based on cuttings simulated to triangulated vegetation point clouds,appears to be the only ALS-based attempt towards this direction.

    The use of ALScan thus be motivated by the potential to obtain a better correspondence with forest biophysical attributes and these data may be available for some areas in a similar extent as land cover maps and other publicly available data.Despite the high potential,however,also ALS-based information may yield a high degree of uncertainties,if applied in expert models formulated according to conventionally measured field attributes.For example,the suitability index proposed by Pukkala et al.(2012)to map potential habitats of Siberian jay(Perisoreus infaustus L.)would require estimating the availability of Vaccinium myrtillus(L.)berries and epiphytic lichens for food and nests.Although sub-models to estimate these attributes are presented(Pukkala et al.2012),also those include stand age and site fertility,which are difficult to estimate by ALS.Although some researchers have predicted even understorey-related attributes,the results of Korpela et al.(2012)indicate that direct measures are difficult to obtain due to transmission losses occurring in the upper canopy(see also Maltamo et al.2005)and such estimations would be even more unreliable based on passive optical RS methods.Even the recognition of dominant tree species may be challenging in ALS-based inventories:despite promising results based solely on ALS(?rka et al.2013;Vauhkonen et al.2014),the results of R?ty et al.(2016)suggest difficulties in detecting species,which dominate a minor proportion of an area otherwise homogeneous in terms of the species.

    On the other hand,ALS may allow producing other attributes with more relevance from the forest management point of view.For example,forests with multilayered vertical structure can be distinguished based on the data(Zimble et al.2003;Maltamo et al.2005),which can be further employed in detecting the prevailing silvicultural system(Bottalico et al.2014),management intensity (Sverdrup-Thygeson et al. 2016;Valbuena et al.2016a),or development stage(Valbuena et al.2016b).Even more detailed indices may be developed based on ecological rationale(Listopad et al.2015)or a thorough understanding of the properties affecting the ALS response(Valbuena et al.2013,2014).Earlier studies have suggested that the information in the ALSdata may be condensed to a few metrics(Kane et al.2010;Leiterer et al.2015;Valbuena et al.2017),the partitioning of which will provide a stratification corresponding closely to the structural complexity observed in the field(Pascual et al.2008;Thompson et al.2016;Vauhkonen and Imponen 2016).

    Even though properties related to individual ESs have been actively studied,no studies that show how to support management decisions related to the provisioning of multiple forest ESs based on three-dimensional forest structure description obtained by ALS can currently be found from the literature.Barbosa and Asner(2017)and Rechsteiner et al.(2017)derived information from ALS data to prioritize landscapes for ecological restoration and species conservation planning,respectively.Packalén et al.(2011)used ALSdata and spatial optimization to derive so called dynamic treatment units to guide the management of pulpwood production in a plantation forest.Although a similar approach could be extended to the decision making of other or multiple ESs(Pukkala et al.2014),all ALS-based applications are,to date,focused on single ESs.

    The purpose of this study is to test ALSdata for management prioritization of multiple ESs in a boreal forest landscape.Proxies for pixel-wise provisioning potential of biodiversity,carbon,timber,berries,and recreational amenities were formulated using ALS-based features and compared to information obtained from forest resource maps with a resolution of 16×16 m2.The quality of land use prioritization based on the obtained information was evaluated under two different decision support models:either using the developed models deterministically or in corporation with the uncertainties of the models.

    Methods

    A methodological overview

    Specifically,the ALS data are tested for predicting the provisioning potential of ESs (Vauhkonen and Ruotsalainen 2017a)in a spatial prioritization framework,where land use decisions are based on ranking the set of decision alternatives in the considered location(s)and choosing the best according to the decision makers’preferences(cf.,Malczewski and Rinner 2015).When applied to prioritize forests for single(e.g.Lehtom?ki et al.2015)or multiple uses (e.g.Vauhkonen and Ruotsalainen 2017a)based on ESproxy maps,a simplified workflow for such analyses includes three methodological steps:

    1)Data acquisition,feature extraction and/or expert modelling to derive proxy values for the analyzed ESs.

    2)Scaling and normalization of the proxy values derived from different sources to the same scale.The resulting values can be called ‘priority’,‘benefit’,or ‘utility’value and used in different ways depending on the literature source(see also Pukkala 2008;Pukkala et al.2014;Malczewski and Rinner 2015).

    3)Decision analyses using the normalized data at selected spatial scale(s).

    Because the normalized proxy maps resulting from the previous steps ‘measure’the ESs in a same scale and account for the value range of each ESin the entire landscape,they can be used(a)to mutually rank ESs within a spatial unit to subsequently prioritize management to provide most suitable ESs in each unit;and(b)to identify the most important locations of specific ESs in the landscape to be considered as management hot-spots or cold-spots.Because the spatial prioritization is carried out at a sub-stand-level using pixels or other corresponding map units,it is expected to allow a more efficient use of the production possibilities of the forest(Heinonen et al.2007)and,overall,operationalize the concept of ESs for landscape planning,which is further motivated by de Groot et al.(2010).

    The present study examines whether changes to each of the three steps listed above could improve pixel-wise analyses of the provisioning potential of forest ESs(cf.the discussion section of Vauhkonen and Ruotsalainen 2017a):

    1) What data to use for the expert models of the provisioning potential: A consolidated approach to obtain grid-based, wall-to-wall predictions for the tessellated landscapes would be to use forest resource maps based on generalizing field sample plot measurements to larger areas using coarse to medium resolution RS images and other numeric map data (Tomppo et al. 2008a, 2008b,2014). This approach, referred to as Multi-Source National Forest Inventory (MS-NFI), was used by Vauhkonen and Ruotsalainen (2017a). Even if ALS allows more prediction possibilities, as reviewed above, it is practically reasoned to benchmark the accuracies against the pixel data provided by the MS-NFI approach, because different forest resource maps are readily available in many countries (Tomppo et al.2008b; Roces-Díaz et al. 2017; Vauhkonen and Ruotsalainen, 2017a).

    2) How to scale the ESs originally measured in different units for the joint analyses: Vauhkonen and Ruotsalainen (2017a) used a simple normalization to convert the ES values between 0 and 1:

    where vijis the normalized value and nijis the position of the j:th plot in ascending order of the expert model values for the i:th ecosystem service among altogether N plots.Notably,this normalization produced values in an interval scale,whereas the ratios between the expert model values could also be assumed useful for the priority ranking.An alternative,ratio-scale normalization could be computed as:

    where vijis the value(or priority or benefit or utility,depending on literature source;see above)produced by the i:th ESin plot j.

    3)How to use the obtained information in decision analyses:Vauhkonen and Ruotsalainen(2017a)deterministically prioritized each pixel to the ES with the highest predicted proxy value,but highlighted the need to consider uncertainties around the predictions.If a quantification of the uncertainties is obtained(e.g.,by approximating residual errors of calibration models fitted to the data),the decision analyses can consider distributions of uncertainty in addition to the expected values and produce separate recommendations for different decision makers according to their attitudes towards risk(Pukkala and Kangas 1996).Therefore,in addition to deterministic use of the predicted values,this study considered both the expected and extreme outcomes of the predictions when selecting the most suitable ESfor a pixel.The principal idea of this analysis is illustrated in Fig.1.

    On this background,the present study tested the data source(ALS or MS-NFI),priority value function form(Eqs.1 or 2),uncertainty management approach,and joint implications of these choices to the predictions of the provisioning potential of forest ESs and subsequent management prioritization decisions.Forest ESs considered were selected based on two criteria:likelihood to occur in the studied landscape and existence of expert models to derive proxies for their provisioning potential based on the field measurements(Table 1).The field and MS-NFI data contained estimates of forest attribute that could be directly inserted to the expert models.Using ALS data,regression analyses were employed to estimate predictive relationships between ALS-features and ESproxy values to fully utilize the different properties of these data(cf.,Section“ALS-based models for the priority valuesof the ESs”below).In the absence of independent,wall-to-wall data for validation,both the predictions and validations were carried out at the level of individual forest plots.The evaluation is therefore limited to the local fitness of the ESs for a specific forest patch in a single point in time and without considering their spatial or temporal continuum.No decision maker was assumed in this study and the values obtained from both the Eqs.1 and 2 were therefore treated with equal weights,even if those could additionally be weighted according to the decision makers’preference structure.

    Fig.1 A generic example of selecting the best decision alternative based on different outcomesof modelpredictions(colored curves).The yellow curve yieldsthe highest priority value based on the expected(upper horizontal line)or worst outcome of the model.However,if the decision maker weightsbest possible outcomes,the alternative depicted by the grey curve should be selected asit producesthe highest priority in the right tail accumulation point(the interception of the curve and the lower horizontalline)

    Table 1 The ESs considered in this study and expert models for deriving their reference proxy values

    Study area and experimental data

    The study area is located in Evo,Finland(61.19°N,25.11°E),which belongs to the southern boreal forest zone.The data extended over an area of approximately 3 km×6 km.The forest stands in the area vary from intensively managed to natural forests in terms of their silvicultural status.Approximately 84%of the growing stock in the studied plots is dominated by coniferous tree species Scots pine(Pinus sylvestris L.)and Norway spruce(Picea abies[L.]H.Karst.).Deciduous tree species such as birches(Betula spp.L.),aspen(Populus tremula L.),alders(Alnus spp.P.Mill.),willows(Salix spp.L.),and rowan(Sorbus aucuparia L.)occur in mixed stands and below the dominant canopy.

    Data sets used were compiled from three earlier studies in the same area(Vauhkonen and Imponen 2016;Niemi and Vauhkonen 2016;Vauhkonen and Ruotsalainen 2017a).Vauhkonen and Imponen(2016)downloaded and processed ALSdata acquired by the National Land Survey of Finland to stratify the area according to forest structural properties.The ALSdata were acquired from a flying altitude of 2200 m using Leica ALS 50 scanner on 7 May,2012,to yield a nominal pulse density of 0.8 m?2.Circular sample plots(9 m radius)were placed by clustering the ALSdata with respect to forest structural features,which was found to be an efficient strategy to distribute the sample across the spatial,size,and age distributions of the tree stock(Vauhkonen and Imponen 2016).The field measurements were carried out in June–August, 2014. The species and diameter-at-breast height(DBH)were measured for each tree with a DBH≥5 cm.For each tree species of the plot,a tree with a DBH corresponding to the median tree was measured for height and used to calibrate height curves for predicting the missing tree heights.Plot-level forest attributes were computed from the tree-level measurements using standard equations and methods,which are described in detail in an open-access article by Niemiand Vauhkonen(2016).

    Publicly available MS-NFI data(Natural Resources Institute Finland 2017)were included to provide a benchmark for the ALSdata.The MS-NFI maps are the same used Vauhkonen and Ruotsalainen(2017a)and details on their pre-processing are given in that paper.These raster maps depicted site fertility,growing stock volume and biomass components by tree species,total basal area and mean diameter and height corresponding to those of the(basal area weighted)median tree,and they were produced using a k-nearest neighbor(k-NN)estimation method based on optimized neighbor and feature selection(Tomppo and Halme 2004;Tomppo et al.2008a,2014).The method used various satellite images from 2012 to 2014 and National Forest Inventory(NFI)field plot measurements from 2009 to 2013,which were updated to correspond the situation in mid-2013 using growth models.

    Altogether 102 field plots were covered by both ALS and MS-NFI data and were included in the analyses.The models of Table 1 were applied to produce plot-specific reference values for the provisioning potential of the ESs based on field data.According to an exploratory analysis,the expert models of Table 1,fit with many different data sets,had considerably different value ranges over the landscape.As a result,a direct normalization of the expert function values specifically with Eq.2 resulted to emphasizing one ESin the priority rankings only because of the different shape and scale of initial value distributions,as elaborated upon in Appendix 1.For this reason,the expert function values of all ESs were transformed to follow the normal distribution as closely as possible using the Box-Cox-transformation(Appendix 1)prior to applying Eqs.1 and 2.The forest attribute estimates based on the MS-NFI maps were transformed using the same parameter values as with field data.This transformation did not affect the order of the observations,but produced approximately equally shaped frequency distributions of every ES,as detailed in Appendix 1.

    ALS-based models for the priority values of the ESs

    Prediction models with independent variables extracted from the ALS data were formulated to predict priority function values of the form of Eq.2.Priority function values corresponding to Eq.1 were obtained by ordering the aforementioned predictions,i.e.,no separate models were constructed for the function form of Eq.2.

    As reasoned in the Introduction,the aim was not to model the forest attributes used as the predictors of the expert models,but to identify and quantify such properties of the ALS point clouds that directly explained the variation in the ES proxies.As visualized in Fig.2,the point clouds of the plots with maximum proxy values did not considerably differ between the ESs in terms of the total distributions.However,when height values or proportions were computed separately according to echo categories,ES-specific differences could be pointed out(Fig.2).The features were therefore extracted in echo categories,which were “only echoes”(suffix_only),“first of many echoes”(_first), “l(fā)ast of many echoes”(_last),“first echoes”(_FP),and “l(fā)ast echoes”(_LP),where the last two categories included “first of many”and “l(fā)ast of many”echoes,respectively,with “only echoes”duplicated in both.Fixed height values of 0.5 m,5 m,and below or above an adaptive height value determined as the height of the 60th percentile were used as the thresholds of ground,shrub,and suppressed or dominant canopy,respectively.

    Fig.2 ALSheight profiles and descriptive characteristics of the field plotsconsidered to be most important locationsof the ESsin the data studied(priority value of 1 based on Eq.2).For comparison,the lower right panel shows a plot that had low priority values of the considered ESs.The black,green,and blue symbols indicate only,first-of-many,and last-of-many ALSechoes,respectively.Grey horizontal linesindicate the mean heightsof these echo categoriesand allechoesand are drawn to illustrate the differencesin termsof these metricsbetween the ESs

    The following categories of the features were considered:

    –Canopy height and density,which are the basic predictors used in ALSanalyses(N?sset 2002)and were assumed to discriminate between size-specific attributes of the ESs:the maximum(hmax),the mean(hmean),and the standard deviation(hstd)of the height values above the ground threshold;the 5th,10th,20th,...,90th,and 95th percentiles(hzz,where zz denoted the percentile value);and the corresponding proportional densities(dzz)were computed according to Korhonen et al.(2008,pp.502–503).

    –Proportion of echoes above a given threshold to all echoes,corresponding to a vegetation cover estimate(Korhonen et al.2011).Thisproportion wascomputed in two ways:using echoes of different categories above theground(ccX_ground,where X istheecho category)or first echoesabovetheshrub layer threshold(ccshrub),which correspondsto an attempt to quantify theshrub layer thickness(cf.,Vauhkonen and Imponen 2016).

    –Absolute differences between mean heights of different echo categories.These features were computed without height thresholds and assumed to discriminate between properties related to coniferous-or deciduous-dominated forest in the ALSdata acquired during the leaf-off period(Liang et al.2007).These features are denoted by diffx–y,where suffix x–y refers to the height difference of echo categories FP–LP,only–LP,first–only,or first–last.

    –Proportions of the different echo categories,which were assumed to be affected by the species and size specific ESproperties in the canopy similar to the ALS-intensity features(?rka et al.2012;Vauhkonen et al.2014).These features are denoted by propX/Y_z,where X/Y indicated the ratio of two echo categories X and Y,and z was the height threshold employed for computations.

    –Predictors related to the shrub and understorey layers(Vauhkonen and Imponen 2016):the ratio of the echoes reflected above ground but below the dominant canopy threshold(runderstory);the standard deviation of the height values of echoes reflected above ground but below the dominant canopy threshold(stdunderstory);and the ratio of the echoes reflected from the shrub layer to all echoes(rshrub).

    Features xi,i=1,2,…,143,listed above formed the initial set S1of candidate predictors.To account for useful interactions between the features,the final set S was obtained aswhich resulted to altogether 10,296 candidate features per plot.Separate models for each ES were constructed by inserting features iteratively into a model template:

    where n is the number of observations,andand yiare the predicted and reference values,respectively.The feature that minimized the RMSE was retained in the model template and the iterations were continued until the model included a maximum of four features.However,more criteria were employed to select the model to be used for the prioritization analyses among the models with one to four features:

    1)The final predictor inserted had to improve the RMSEby at least 1%.

    2)The residual errors had to satisfy the null hypothesis that the considered sample came from a normally distributed population,which was examined graphically using scatter,residual and QQ-plots,and numerically using the test statistic proposed by Shapiro and Wilk(1965).

    3)The model had to pass a“sensitivity of convergence”test,in which the model was fit separately for each plot using Leave-One-Out-Cross-Validation(LOOCV),i.e.,not allowing the plot in question to be available in the training data for model fitting.Implications of including this test are further described in the Results section.

    Predicting the priority values of the ESs based on the MS-NFImaps

    Benchmark predictions for those based on ALSwere obtained by inserting the forest attribute estimates from the MS-NFI maps to Eq.2.Priority function values corresponding to Eq.1 were obtained by ordering the aforementioned predictions (cf.,previous section).The MS-NFI maps included estimates of all other independent variables except the number of trees per hectare,which was estimated by dividing the total basal area by the basal area corresponding to the mean diameter,i.e.,assuming that the resulting number of average-sized trees existed in a pixel.To compute plot-wise estimates,the pixels of the forest resource maps intersecting with the plot polygons were identified using a spatial query.The estimates of a plot were obtained from the intersecting pixels as weighted averages with the joint areas of the plots and pixels as the weights.Finally,to see if amending the models based on ALS with the MS-NFI layers improved the models,a similar feature selection as with ALS data was run including all MS-NFI-based ES and forest attribute proxies as additional feature candidates.

    Field calibration and evaluation of the predictions

    Following the method described in the previous section,potential estimation errors in the MS-NFI maps propagate to the predicted priority values,whereas similar error propagation is avoided in the ALS-based analyses due to local model fitting.An additional calibration step was therefore included to eliminate the contribution of the local field sample to the predictions.Calibration models,where yiwas the reference priority value of the i:th ES andits RS-based estimate,of all ESs were fit simultaneously as systems of linear equations.Due to the high inter-correlations(see Additional file 1,Table S1),the models were fit in two steps:first,using Ordinary Least Squares(OLS)to produce model residuals,and second,using Seemingly Unrelated Regression(SUR)to account for the residual error covariance matrices in the final models.The computations were carried out in the LOOCV mode using the systemfit package of R(Henningsen and Hamann 2007).The accuracies of the ALS-and MS-NFI-based predictions were compared using the RMSE and coefficient of determination(R2)computed between the reference values and predictions obtained from the LOOCV models.

    Decision analyses

    The effects of the aforementioned prediction accuracies to the management decisions were evaluated by comparing the priority ranking of the ESs in each individual plot.The ES with the highest priority value,based on Eqs.1 or 2 applied to the field reference data,was assumed to be the most suitable ES for the specific plot.The RS-based decision was considered correct,if the most suitable ES based on the field data and the RS prediction equaled.The degree of incorrect decisions was quantified using two approaches.First,the correctness of every decision was given a numerical score(Gopal and Woodcock 1994):situations where RS and field data resulted in the same decision was given a score of 6;those where the RS-based service was the second best according to the field data a score of 5;and so on,until the situation where the RS-based service was the worst according to field data,which was given a score of 1.The distributions of these “decision scores” were compared between the different data sources.Second,the dispersion in field and RS-data between the services selected as the most suitable for the specific plot was examined using confusion matrices.The priority ranking of the less important ESs was not evaluated.

    In addition to ‘deterministic’decision making described above,the sensitivity of the decisions was examined by incorporating the uncertainties of the models to the analyses(Fig.1).Instead of using the expected values of the priority functions,the ranking was carried out assuming the predictions as realized values of a random variable X~N(E,s2),where E was the expected value and s2was the mean squared error of the model residuals.A similar priority ranking as with the expected values was carried out with predictions that were among the worst and best outcomes of the model,obtained as the values of the 5th and 95th percentiles of the distribution of X of each ES.

    Results

    ALS-based models for the priority values of the ESs

    The ALS features considered as the predictors of the regression models are listed in Table 2.All feature and echo type categories and a wide range of different height values was employed when building the models,for which reason only a few specific observations on the structure of models can be made.All features selected were products of form feature1×feature2,where feature1was often an absolute height value(a mean height,percentile,or height difference)and feature2a proportion (either a canopy cover proxy or proportional density).This combination was especially frequent among the first features selected to the models.In the models of TIMB,all selected predictors were such combinations employing various height values and echo categories.The models of BIOD and CARB used proportion×proportion types of interactions and the ratio of first-of-many to only and first returns(propfirst/FP_ground).The predictors of BIOD(e.g.,propfirst/FP_ground;diffonly–LP;hstdLP)were most diverse in terms of describing the canopy structure with features from different categories.The models of BILB,COWB,and AMEN differed from those mentioned above in employing low percentile values,last pulse proportions and features such aspropfirst/FP_ground, andOverall,the canopy cover proxies were the most frequent feature type,whereas the computing heights and echo categories of all features varied.Although a wide range of different height values was used,a predictor with a percentile value above 70 was selected only once.

    Table 2 The features and performance of ALS-based models for predicting ratio-scaled ESproxy values.W–Shapiro-Wilk test statistic

    The graphical assessment of model residuals(detailed results not shown)was mainly in line with the test on residual normality(Table 2):the QQ-plots showed heavy-tailed residuals especially for the models with statistically significant values of the Shapiro-Wilk test statistic.However,the deviations of normality were typically related to one or two plots with the highest or lowest values,and not considered problematic for the further analyses.When examined in the same data used for constructing the models,the performance of every model could be slightly improved by increasing the number of predictors to the maximum number allowed.However,when the models were re-fit using LOOCV,model parameters could not be solved for at least one of the plots in the data,resulting to NA values for this performance factor in Table 2.Although this effect could probably have been avoided by allowing a slightly wider range of initial parameters when fitting the models,it was also considered as a sensitivity issue reflecting an over-parameterization of the initial model to certain types of forest structures.

    The volume of deciduous trees and the MS-NFI based proxy for BILB would have replaced the last ALS-based features in the models of CARB and BILB,respectively,and in the models of COWB,the corresponding MS-NFI proxy would have been selected as the second feature.However,none of the aforementioned MS-NFI features performed better than the ALS-features of these models in terms of the feature selection criteria.Based on the considerations above,the ALSand MS-NFI data sets were always used separately.Also,a different number of predictors was used in the ALS-based models for the priority ranking:BIOD and COWB were modeled using only one predictor(the one selected first);TIMB,BILB,and AMEN using two predictors(those selected first and second);and CARB using three predictors selected first.

    Comparison of ALSand MS-NFIfor predicting the priority values of the ESs

    The models based on ALS data always outperformed those based on forest attribute estimates derived from the MS-NFI maps.As shown in Figs.3 and 4,the ALS-based models generally explained more variation in the ES proxies.The regression lines of the MS-NFI data based on the SUR calibration models also differed more severely from the 0–1-lines.Using MS-NFI data,TIMB was predicted most accurately with an RMSE of 30.4%.The RMSEs of other ESs were also close(30.6%–33.2%),except AMEN,which had an RMSE of 40.5%and BIOD,which was predicted worst with an RMSE of 41.6%.Using ALS,the ES predicted worst(COWB)had an RMSE of 29.8%,which is 97%of the RMSE of the corresponding MS-NFI prediction.The RMSEs of all other ESs were in order of 21.7%–27.5%(57%–83%of the RMSEs of MS-NFI predictions),except CARB,which was predicted most accurately with an RMSE of 15.1%(47%of the RMSE of MS-NFI prediction).The degree of determination of CARB also improved most due to using ALS instead of MS-NFI,from R2=0.11 to 0.81.The R2-improvements of the other ESs were close to this magnitude,except for BILB and COWB,which had R2values close to each other based on both the data sources.The residual errors of models based on ALSand MS-NFIwere somewhat correlated for BILB and COWB,but not for the other ESs(Figs.3 and 4,right column).

    Fig.3 Predicted(x-axis)versus reference priority values of BIOD(upper row),TIMB(middle row)and CARB(bottom row)based on MS-NFI(left column)or ALS(middle column).The broken and solid lines are the 1:1 line and regression line of the SURcalibration models fit with the local field sample,respectively.The right column shows the residuals of the corresponding predictions based on ALS(x-axis)and MS-NFIdata

    Decision analyses based on different priority functions,data,and model uncertainties

    Compared to the use of interval-scaled priority functions,those based on the ratio scale slightly increased the proportion of decisions that had a perfect agreement in both ALSand field data(Fig.5,left panel).The same observation was made regarding the decisions based on the MS-NFI data,but the differences between the priority function forms were in general minor.When the ratio-scaled priority functions were used in the remaining analyses,the ALS and field data resulted to the same decision in 42%of the plots;the ALS-based ES was at least the second best according to field data in 69%;and among the three best alternatives in 84%of the plots.With MS-NFI data calibrated by the local field sample,the corresponding figures were 46%,61%,and 72%and slightly lower without the calibration,the distribution of the decision scores of all data sources being shown in Fig.5(right panel).Thus,although the calibrated MS-NFI data did better in selecting ESs that matched perfectly with the field data,the proportion of poorest decisions was considerably lower based on the ALS data.The ALS-based models resulted to a better decision in 32%,those based on the MS-NFI data in 27%,and the decision equaled in 41%of the plots.Of the latter group of plots,around 2/3 had either BILB or COWB as the most important ES and the decisions for these plots were generally scored≥4.Except for data sources,the decision score was highly dependent on the priority difference between the best ESs observed from the field plots:in the plots where the decisions equaled between the data sets,the average priority difference between the ESs prioritized as the first and second was 0.12(standard deviation 0.08),whereas the corresponding figures for the other plots were 0.07(0.07).

    Fig.4 Predicted(x-axis)versus reference priority values of BILB(upper row),COWB(middle row)and AMEN(bottom row)based on MS-NFI(left column)or ALS(middle column).The broken and solid lines are the 1:1 line and regression line of the SURcalibration models fit with the local field sample,respectively.The right column shows the residualsof the corresponding predictionsbased on ALS(x-axis)and MS-NFIdata

    Fig. 5 The distribution of decision scores (left:) between interval- (Eq. 1) and ratio-scaled (Eq. 2) priority functions in ALS data, or (right:) between data source in ratio-scaled priority. Situations where RS and field data resulted in the same decision was given a score of 6; those where the RS-based service was the second best according to the field data a score of 5; and so on, until the situation where the RS-based service was the worst according to field data, which was given a score of 1.

    Decision making based on the 5th percentile of the ALS-predicted priority value distributions reduced the proportion of worst decisions(Fig.6,left panel).Otherwise,the decisions based on either the 5th or 95th percentile did not compare favorably with those based on the expected value in either data(Fig.6).The confusion matrices for the most important ESs based on the different data sources and either the expected or extreme outcomes are shown as Appendices 2 and 3,respectively.A comparison of the confusion matrices based on either expected or extreme values indicates that many plots with the most important ES as those predicted with the highest or smallest error rates(e.g.,CARBor COWB,respectively,based on ALSdata)could be prioritized for this ES only by explicitly considering the extreme model outcomes.As a number of plots obtain a different prioritization based on the expected values,it was found interesting to look at the plots for which the prioritization changed depending on the model outcome.Using ALS-based ESproxy models,the prioritization based on the expected,worst,and best outcomes equaled in only 22 plots(in 43 using MS-NFI maps calibrated with the field data).If all decision alternatives based on the three outcomes of the ALS-models were considered,the alternative that matched perfectly with the field data was included in 57%of the plots;an alternative among the two best ones in 85%;and among the three best ones in 93%of the plots.With MS-NFI data,the corresponding figures were 59%,76%,and 88%.Thus,especially the ALS-models were found useful in confining the decision alternatives to those most feasible according to production possibilities,from which the one preferred most preferred according to the stakeholder preferences could be selected based on further MCDA.

    Discussion

    Earlier RS-based decision analyses of multiple ESs have relied on map scales such as 1:25,000(Roces-Díaz et al.2017)or pixel sizes such as 500×500 m2(Schr?ter et al.2014).Also smaller pixel sizes of 60×60 m2(Lehtom?ki et al.2015)or 48×48 m2(Vauhkonen and Ruotsalainen 2017a)have been used in spatial prioritization analyses.Many of the aforementioned scales are coarse considering the need to formulate management prescriptions at the level of operational units(e.g.,forest compartments),which are typically 1.5–2.0 ha in size in Finland(Koivuniemi and Korhonen 2006).Based on this background,the results of the present study are encouraging:the use of ALS in particular allowed deriving accurate predictions for plots of around 250 m2,i.e.,in a considerably more detailed resolution than current operational compartments.The RMSEs of predicting the studied ES indicators by ALSwere 47%–97%of the RMSEs of corresponding predictions based on benchmark forest resource maps derived using coarse satellite images.Due to applying a similar field calibration step on both of the data sources,the difference can be attributed to the better ability of ALS to explain the variation in the ES proxies.In the sub-sections below,the results are discussed from the points of view of using ALSdata to predict various ESindicator proxies(Table 1)and using these predictions in decision analysescompared to other data.

    Fig. 6 The distribution of decision scores based on the use of ratio-scaled priority functions and different outcomes of the prediction models in ALS (left) and calibrated MS-NFI data (right). See the caption of Fig. 5 for the interpretation of decision score values.

    On the ESproxies and their ALS-based modelling

    The expert models(Table 1)express the suitability of forests for the ESs as transformations of forest attributes.Applying the expert models yielded the highest biodiversity conservation values for mature,densely stocked forests.The values were weighted by site fertility such that most fertile sites received a considerable weight compared to poorer sites.An increasing basal area and mean diameter increased the soil expectation value of all the species,but otherwise the model included species-specific interactions between these and operational environment related parameters.The value of carbon storage increased according to an increasing stem volume depending on the species-specific biomass expansion factors.An increasing maturity(measured in terms of mean age and diameter)and decreasing stand density(measured in terms of either basal area or stem number)increased the suitability of a site for berry picking and visual amenity.The latter models also included species-specific terms such that especially the presence of pine trees improved the values.Thus,the response variables of all other ESs except CARB were modelled as functions of multiple forest attributes.

    The models based on the ALSdata(Table 2)explained the differences in the provisioning potential of the different ESs with RMSEs of 15%–30%.The selected features and model performances are well in line with the background given in the previous paragraph.The predictions of CARB had the highest accuracies,because those essentially explained the variation in the total stem volume converted to carbon using biomass expansion factors.The ratio of first-of-many to all first echoes(propfirst/FP_ground)was used as a predictor of CARB in addition to height and density metrics,which are typical to total biomass or carbon models.The aforementioned feature has been found useful for separating species(?rka et al.2012;Vauhkonen et al.2014)and should be studied in a broader forest modeling context.Although the field proxy value for TIMB was computed using species-specific predictors,the model based on the ALS data included only features based on height.The models for the other ESs included ALS-based predictors that were clearly related to forest canopy structure:for example,features indicating the existence or abundance of low vegetation were frequently selected to the models of BIOD or recreation-related ESs,respectively.No clear recommendations for the selection of features can be given,except for using multiple heights and echo types when computing the candidate features.The requirement to estimate a high number of model parameters was likely reduced by including products of all candidate features to account for interactions between them,which is a useful property that has not been reported in earlier ALS studies.The features used here are the most common and easily implementable using available software packages such as R.However,it is acknowledged that not all features presented in the literature were included and it could be possible to improve the results with more experimental ones such as those related to volumetric(Vauhkonen and Ruotsalainen 2017b)or textural(Niemi and Vauhkonen 2016)properties.

    The proxies listed in Table 1 are measured in different units.In order to use the proxies in decision analyses,those need to be normalized to the same scale,which was done in this paper prior to the modeling.Due to the transformations,the prediction accuracies cannot be easily compared with earlier studies,even in relative scale.Even CARB,which is a species-specific transformation of the total volume,cannot be directly compared to previous studies that predict total biomass or carbon due to the Box-Cox-transformation (Appendix 1) applied to equalize the distribution of the response variables for the decision analyses.If a similar model for CARB had been fit without the Box-Cox-transformation,the RMSE would have risen to around 34%.It is higher than(e.g.)the RMSE of 25.7%obtained by Kankare et al.(2015)for total biomass in the same region,but when comparing the figures the differences in the definition of the response variable and a slightly different plot size must also be considered.For more reasoning on the need for the transformations,please see the next section.On the other hand,the modeling task considered above may have been alleviated by the fact that all response variables resulted from models that had been formulated earlier using common forest mensurational attributes.Actual differences between berry yields of two forests could be much more discrete than those predicted as a function of forest attributes.The performance of the models should thus be tested by re-fitting or validating the models against ES-specific indicators that are not based on models but direct observations made in the field(see also Hegetschweiler et al.2017;Kohler et al.2017).

    On the other hand,it may be less feasible or even impossible to predict certain ES properties otherwise than using features quantifying the three-dimensional forest structure.Examples in the Scandinavian boreal forest structures include mammal or bird habitats(e.g.,Melin et al.2013,2016),which could have realistically occurred in the studied area,but could not be included in the present comparison due to the lack of models based on information obtainable from forest resource maps or field plots.The value of ALS data can be seen to lie especially in applications that require identifying sites with high value for e.g.conservation as ALS data coverage is globally increasing and the method has been proven to provide 3D descriptions of vegetation structure that,in turn,have been long known as primary determinants of habitat quality and biodiversity(MacArthur and MacArthur 1961;Dueser and Shugart Jr,1978;Brokaw and Lent,1999).Nevertheless,Vihervaara et al.(2017)identified only four Essential Biodiversity Variables that could benefit from the use of RS data.According to this study,the obtainable improvements are most likely indicator-specific,with magnitude depending on what RS data are available.

    Other aspects of RS-based decision analyses:Data,normalization,and uncertainties

    This study tested normalization resulting to values in either an interval-(Eq.1)or ratio-scale(Eq.2).Even though the priority value functions based on the different scales did not differ considerably,those based on the ratio-scale performed slightly better in the priority ranking,which is logical as also the ALS-based features provide information at a ratio scale.Although the purpose is not to exhaustively discuss the implications of different value function forms,one important practical aspect discovered in the exploratory analysis could be brought up:An incorrect selection of a value function form would result in biased decisions by weighting the rank-orderings of the alternatives to an undesired direction,which is exemplified by a comparison of the transformed and non-transformed value function forms(Appendix 1).This discussion highlights the importance of considering data normalization procedures in detail already at the modeling stage,if the final applications aim at decision analyses where all modeled proxies should be measurable at the same scale.

    The ALS and MS-NFI data sources showed several differences,when predicting the priority value functions.Except having a more deterministic relationship with the forest attributes,one additional improvement of the locally fit ALS models is the ability to predict the 0–1-value ranges directly.When using forest attributes such as those predicted by the MS-NFI,the maximum values used in the normalization should be representative of the entire area.In this study,the minima and maxima predictions based on the MS-NFI were more severely incorrect than with ALS data(Figs.3–4),which partially explains the poorer performance of this data source.Many multivariate predictions from RS data are based on non-parametric nearest neighbor methods,which could have been considered also in the case of ALS data.However,the aforementioned problem would be seriously present also in those types of predictions.

    Regardless of the input data or applied scale or mapping technique,it is well recognized that the resulting ES maps will include uncertainties(Eigenbrod et al.2010;Schulp et al.2014;R?s?nen et al.2015).A few earlier spatial prioritization studies(Lehtom?ki et al. 2015; R?s?nen et al.2015;Vauhkonen and Ruotsalainen 2017a)used forest attribute maps in a similar resolution as considered here,but without a calibration based on field data.Such a practice cannot be recommended due to the observed uncertainties in the uncalibrated MS-NFI data in particular.However,it should be acknowledged that all analyses carried out in the aforementioned studies might not be sensitive to incorrect pixel-level prioritization decisions.Also,each of the aforementioned studies used either aggregated pixels to somewhat account for these effects.It is not possible to address the uncertainty reduction due to the use of aggregated resolution based on the field data of this study,which was collected from fixed-size plots.

    Finally,it should be noted that in the actual decision making,the stakeholder preferences may affect or even dictate the allocation of the ESs over suitability of forest structure.Even if the decision maker had no preferences on the ESs,aggregating individual pixels,i.e.,deviating from their local optima to compose larger treatment units,could be feasible with respect to the implementations of management prescriptions or achieving ecological or economic objectives that are determined over a larger area(Pukkala et al.2014).Importantly,the decisions based on the same data may differ for a risk-avoiding,risk-neutral or risk-seeking decision maker(Pukkala and Kangas 1996).The risk preferences of the decision maker should clearly be incorporated in the decision making based on the ES maps.Here,a similar technique as in Pukkala and Kangas(1996)was used to account for the uncertainties emerging from different accuracies of the ES proxy models.The results reported in the last paragraph of the Results section can be concluded such that separate forest ES maps should be prepared using the expected,worst,and best outcomes of the model predictions to fully describe the production possibilities of the landscape under the uncertainties in the models.Also those related to the decision makers’preferences could be accounted for as additional stochastic distributions in the analyses(e.g.,Kangas et al.2007)and incorporated in analyses as pixel-level production constraints.Overall,the prioritization approach should be tested further involving real decision makers.

    Conclusions

    The provisioning potential of ESs(biodiversity,timber,carbon,berries,and visual amenity)was modeled as expert model-based proxies that express the suitability of forests for the ESs as transformations of forest attributes.The models based on the ALS data explained the variation in these proxies with RMSEs of 15%–30%.The RMSEs of the ALS-based models were 47%–97%of the RMSEs of corresponding predictions based on the MS-NFI forest resource maps.Due to applying a similar field calibration step on both of the data sources,the difference can be attributed to the better ability of ALS to explain the variation in the ES proxies.

    The RMSE-differences did not fully translate to the accuracies of land use decisions:instead,prioritizing the land use for the ESs with the highest provisioning potential could be done with rather similar accuracies based on both data sources and at a resolution of 250 m2,i.e.,in a considerably more detailed scale than current operational forest management units.ALS-based models for the ES proxies can however be recommended based on their better stability regarding the model errors.The results suggest that separate forest ES maps should be prepared using the expected,worst,and best outcomes of the model predictions to fully describe the production possibilities of forest under the uncertainties in the models.

    Additional file

    Additional file 1:Empirical cumulative density and probability density function formsand Pearson correlation coefficientsof different transformations of the expert model predictionsfor the ESsconsidered.(DOCX 1758 kb)

    Appendix 1

    Box-Cox transformation of the reference priority values for the ESs

    An exploratory analysis revealed a practical comparability issue related to using the expert function values for the ESs directly in Eq.2,which can be demonstrated by visualizing the empirical cumulative distribution functions and histograms of the data(Additional file 1:Figure S1-Figure S6).The expert functions of Table 1 were fit with many different data sets having different value ranges and resulting to unequal frequencies for the different ESs,when applied in the present data.Using Eq.2,these differences would have translated directly to the priority values,which was found problematic with respect to their ranking.Look especially at the left-hand columns of Additional file 1:Figure S1-Figure S6 and compare Additional file 1:Figure S6 to the others:the direct use of these values would have resulted to a very high number of plots being prioritized as AMEN only because its distribution more frequently included higher values compared to the distributions of the other ESs,which were more often skewed to the left.For this reason,the expert model values of every ES were transformed to produce frequencies that followed the normal distribution as closely as possible prior to converting the value ranges between 0 and 1.The transformation was obtained as(Box and Cox 1964):

    where yijis the original value of the i:th observation of the j:th ES proxy andλis a parameter.The value ofλwas selected from an interval of?3 to 3 as the value that maximized a test statistic on whether the considered sample came from a normally distributed population(Shapiro and Wilk 1965).This transformation did not affect the order of the observations,but produced approximately equally shaped frequency distributions of every ES.As a result,the ESs could be prioritized using two alternative priority value function forms illustrated in the two rightmost columns of Additional file 1:Figure S1-Figure S6:either according to the interval scale(Eq.1)based only on the order of the observations or according to the ratio scale(Eq.2)preserving the ratios between the observations,but thanks to the Box-Cox-transformation,having approximately equal(close to normal)frequency distributions between the ESs.The function forms and frequency distributions of the priority values are shown in Additional file 1:Figure S1-Figure S6 and the correlations between the priority valuesin Additional file 1:Table S1.

    During the feature selection process,it was noted that exactly the same features would have been selected to the models,regardless of whether the original or Box-Cox-transformed response variables were used.Therefore,although the Box-Cox-transformation has been rarely used in ALS-based studies,it could potentially aid also other applications by normalizing the response variable to a desired form in the model fitting step.

    Appendix 2

    Confusion matrices for the most important ESs based on expected outcomes

    Table 3 Confusion between ESs considered as most important based on the field data(observed)and MS-NFImaps(predicted)using the expected values of the predicted ESproxies

    Table 4 Confusion between ESs considered as most important based on the field data(observed)and MS-NFIdata calibrated with the local field sample(predicted)using the expected values of the predicted ESproxies

    Table 5 Confusion between ESs considered as most important based on the field data(observed)and the expected values of the ALS-based modelsfor ESproxies(predicted)

    Appendix 3

    Confusion matrices for the most important ESs based on extreme outcomes

    Table 6 Confusion between ESs considered as most important based on the field data(observed)and MS-NFIdata calibrated with the local field sample(predicted)using the worst outcomes of the predicted ESproxies

    Table 7 Confusion between ESs considered as most important based on the field data(observed)and the worst outcomes of the ALS-based models for ESproxies(predicted)

    Table 8 Confusion between ESs considered as most important based on the field data(observed)and MS-NFIdata calibrated with the local field sample(predicted)using the best outcomes of the predicted ESproxies

    Table 9 Confusion between ESs considered as most important based on the field data(observed)and the best outcomes of the ALS-based models for ESproxies(predicted)

    Abbreviations

    ALS:Airborne Laser Scanning;AMEN:Visual amenity(one of the ecosystem services considered,see Table 1);BILB:Suitability for bilberry picking(one of the ecosystem services considered,see Table 1);BIOD:Biodiversity(one of the ecosystem services considered,see Table 1);CARB:Carbon storage(one of the ecosystem services considered,see Table 1);COWB:Suitability for cowberry picking(one of the ecosystem servicesconsidered,see Table 1);DBH:Diameter-at-breast-height;ES:Ecosystem service;k-NN:k-Nearest neighbor;LOOCV:Leave one out cross validation;MCDA:Multiple criteria decision analysis;MS-NFI:Multi-Source National Forest Inventory;RMSE:Root mean squared error;RS:Remote sensing;TIMB:Timber production(one of the ecosystem services considered,see Table 1)

    Funding

    The acquisition of the studied data was originally supported by the Research Funds of University of Helsinki.

    Availability of data and materials

    All data and materials can be obtained by requesting from the author.

    Author’s contributions

    JVis the sole author.He carried out all analyses and drafted the manuscript.The author read and approved the final manuscript.

    Ethics approval and consent to participate

    Not applicable.

    Competing interests

    The author declares that he has no competing interests.

    Received:14 March 2018 Accepted:24 May 2018

    91av网站免费观看| 91字幕亚洲| 高清毛片免费观看视频网站| av中文乱码字幕在线| 亚洲自偷自拍图片 自拍| www国产在线视频色| 日日干狠狠操夜夜爽| 欧美黑人欧美精品刺激| 国产精品综合久久久久久久免费 | 久久香蕉国产精品| 97碰自拍视频| 日韩高清综合在线| 亚洲全国av大片| 中文字幕精品免费在线观看视频| 亚洲第一电影网av| 午夜精品在线福利| 88av欧美| 激情视频va一区二区三区| 欧美日韩瑟瑟在线播放| 在线观看免费午夜福利视频| 91精品三级在线观看| 国产精品综合久久久久久久免费 | 欧美一级a爱片免费观看看 | 国产人伦9x9x在线观看| 精品久久久久久久毛片微露脸| 神马国产精品三级电影在线观看 | 欧美激情久久久久久爽电影 | 99国产精品99久久久久| 午夜成年电影在线免费观看| 精品欧美一区二区三区在线| 日本vs欧美在线观看视频| 两个人视频免费观看高清| 亚洲七黄色美女视频| 两个人视频免费观看高清| 日本a在线网址| 亚洲av日韩精品久久久久久密| 亚洲人成77777在线视频| 亚洲成人国产一区在线观看| 91成人精品电影| 亚洲欧美激情在线| 成人精品一区二区免费| 久久久久精品国产欧美久久久| 多毛熟女@视频| 欧美中文综合在线视频| 99久久综合精品五月天人人| 如日韩欧美国产精品一区二区三区| 欧美成人性av电影在线观看| 99国产综合亚洲精品| 美女高潮到喷水免费观看| 长腿黑丝高跟| 国产又爽黄色视频| 少妇 在线观看| 久久久久久久久中文| 精品高清国产在线一区| 国产精品自产拍在线观看55亚洲| 国产精品免费一区二区三区在线| 天天躁夜夜躁狠狠躁躁| √禁漫天堂资源中文www| 国产精品日韩av在线免费观看 | 神马国产精品三级电影在线观看 | 亚洲成av人片免费观看| 亚洲欧美日韩另类电影网站| 日本vs欧美在线观看视频| 日本一区二区免费在线视频| 中文字幕人妻熟女乱码| 可以在线观看的亚洲视频| 国产色视频综合| 国产av又大| 免费看十八禁软件| 99香蕉大伊视频| 热99re8久久精品国产| 麻豆国产av国片精品| 制服诱惑二区| 制服诱惑二区| 亚洲人成电影免费在线| 69精品国产乱码久久久| 欧美人与性动交α欧美精品济南到| 日韩视频一区二区在线观看| 97人妻精品一区二区三区麻豆 | 精品免费久久久久久久清纯| 美女高潮喷水抽搐中文字幕| 国产xxxxx性猛交| 在线观看www视频免费| 久久香蕉精品热| 中文字幕久久专区| 日本一区二区免费在线视频| 黄色视频,在线免费观看| 国产精品久久久久久亚洲av鲁大| 波多野结衣巨乳人妻| 国产又爽黄色视频| 国产成人精品久久二区二区免费| 欧美成人免费av一区二区三区| 日日干狠狠操夜夜爽| 亚洲男人天堂网一区| 久久久水蜜桃国产精品网| 巨乳人妻的诱惑在线观看| 在线视频色国产色| 精品久久久久久,| 好男人电影高清在线观看| 精品国产超薄肉色丝袜足j| 久久久久久免费高清国产稀缺| 国产精品亚洲一级av第二区| 大型黄色视频在线免费观看| 国产一级毛片七仙女欲春2 | 国产区一区二久久| 久久久国产成人免费| 午夜福利在线观看吧| 久久久久久国产a免费观看| 国产精品精品国产色婷婷| 国产欧美日韩精品亚洲av| 免费在线观看完整版高清| 成人国产一区最新在线观看| 欧美亚洲日本最大视频资源| 国产亚洲av高清不卡| 国产一卡二卡三卡精品| 日韩 欧美 亚洲 中文字幕| 在线免费观看的www视频| 国产伦人伦偷精品视频| 亚洲aⅴ乱码一区二区在线播放 | 婷婷六月久久综合丁香| 亚洲 国产 在线| 午夜福利高清视频| 国产欧美日韩一区二区三区在线| 一区二区三区激情视频| 亚洲成人久久性| 色哟哟哟哟哟哟| 91精品国产国语对白视频| 亚洲成av片中文字幕在线观看| 午夜福利欧美成人| 在线视频色国产色| 亚洲成人精品中文字幕电影| 亚洲伊人色综图| 嫩草影视91久久| 国产男靠女视频免费网站| 国产一区在线观看成人免费| 国产精品久久久av美女十八| 一级a爱片免费观看的视频| 午夜a级毛片| 国产91精品成人一区二区三区| 19禁男女啪啪无遮挡网站| 淫秽高清视频在线观看| 久久人妻福利社区极品人妻图片| 久久天堂一区二区三区四区| 在线观看免费视频网站a站| 亚洲专区国产一区二区| 亚洲专区字幕在线| 9191精品国产免费久久| 一级毛片精品| 黑人欧美特级aaaaaa片| 熟女少妇亚洲综合色aaa.| 精品欧美一区二区三区在线| 欧洲精品卡2卡3卡4卡5卡区| 热99re8久久精品国产| bbb黄色大片| 别揉我奶头~嗯~啊~动态视频| 欧美日韩福利视频一区二区| 热99re8久久精品国产| 成人亚洲精品av一区二区| 50天的宝宝边吃奶边哭怎么回事| 女警被强在线播放| 国产精品久久视频播放| 大型av网站在线播放| 性色av乱码一区二区三区2| 制服丝袜大香蕉在线| 香蕉丝袜av| 久久精品人人爽人人爽视色| 丰满的人妻完整版| 亚洲成av片中文字幕在线观看| 久久亚洲精品不卡| 亚洲成人国产一区在线观看| 欧美成人一区二区免费高清观看 | 韩国精品一区二区三区| 欧美国产精品va在线观看不卡| 最新在线观看一区二区三区| 亚洲一卡2卡3卡4卡5卡精品中文| 精品午夜福利视频在线观看一区| 国产精品亚洲av一区麻豆| 美女扒开内裤让男人捅视频| 制服人妻中文乱码| 成人永久免费在线观看视频| 亚洲精品一卡2卡三卡4卡5卡| 亚洲专区中文字幕在线| 一区二区日韩欧美中文字幕| 日韩欧美三级三区| 久久天堂一区二区三区四区| 日韩欧美一区二区三区在线观看| 欧美激情极品国产一区二区三区| 国产亚洲欧美在线一区二区| 精品免费久久久久久久清纯| 一区二区三区精品91| 久久人妻熟女aⅴ| 久9热在线精品视频| 中国美女看黄片| 视频在线观看一区二区三区| 性色av乱码一区二区三区2| 亚洲国产欧美一区二区综合| 18禁裸乳无遮挡免费网站照片 | 国产激情欧美一区二区| 一级毛片女人18水好多| 亚洲情色 制服丝袜| 夜夜夜夜夜久久久久| 大码成人一级视频| 成人18禁高潮啪啪吃奶动态图| 国产精品98久久久久久宅男小说| 欧美av亚洲av综合av国产av| 一区福利在线观看| 久久亚洲精品不卡| 欧美激情高清一区二区三区| 日韩av在线大香蕉| 两个人视频免费观看高清| 这个男人来自地球电影免费观看| 无人区码免费观看不卡| 国产av一区在线观看免费| 国产激情欧美一区二区| av电影中文网址| 又黄又粗又硬又大视频| 一本综合久久免费| 国产精品98久久久久久宅男小说| 村上凉子中文字幕在线| 免费av毛片视频| 久久久精品欧美日韩精品| 一进一出抽搐动态| 久久这里只有精品19| 国产精品久久电影中文字幕| 国产激情欧美一区二区| 老司机在亚洲福利影院| 国产高清视频在线播放一区| 久久狼人影院| 欧美日本中文国产一区发布| 国产av一区二区精品久久| 老汉色av国产亚洲站长工具| 国产精品精品国产色婷婷| 国产亚洲欧美在线一区二区| av有码第一页| 少妇裸体淫交视频免费看高清 | 国产免费男女视频| 日韩免费av在线播放| 精品人妻1区二区| 美女高潮喷水抽搐中文字幕| 亚洲七黄色美女视频| 香蕉国产在线看| 国产97色在线日韩免费| 精品一区二区三区av网在线观看| 色综合婷婷激情| 久久精品亚洲熟妇少妇任你| 精品不卡国产一区二区三区| 久久人人精品亚洲av| 国产av精品麻豆| 亚洲av第一区精品v没综合| 青草久久国产| 久久午夜综合久久蜜桃| 999久久久精品免费观看国产| 久久久久久国产a免费观看| 日本一区二区免费在线视频| 不卡av一区二区三区| 国产免费av片在线观看野外av| 黄片大片在线免费观看| 精品第一国产精品| 男人舔女人的私密视频| av在线天堂中文字幕| 自拍欧美九色日韩亚洲蝌蚪91| 亚洲 欧美一区二区三区| 91字幕亚洲| 亚洲精品美女久久久久99蜜臀| 91国产中文字幕| 成人欧美大片| 999久久久国产精品视频| 69av精品久久久久久| 国产人伦9x9x在线观看| 18禁观看日本| 成人免费观看视频高清| 国产欧美日韩一区二区精品| √禁漫天堂资源中文www| 777久久人妻少妇嫩草av网站| 亚洲少妇的诱惑av| 欧美激情 高清一区二区三区| cao死你这个sao货| 性欧美人与动物交配| 国产免费男女视频| 久久天堂一区二区三区四区| 桃红色精品国产亚洲av| 亚洲狠狠婷婷综合久久图片| 国产亚洲av嫩草精品影院| 伦理电影免费视频| 熟妇人妻久久中文字幕3abv| 夜夜看夜夜爽夜夜摸| 亚洲国产欧美日韩在线播放| 亚洲精品在线观看二区| 日韩 欧美 亚洲 中文字幕| 午夜a级毛片| 久久天堂一区二区三区四区| www.999成人在线观看| 日韩欧美一区视频在线观看| 岛国视频午夜一区免费看| 19禁男女啪啪无遮挡网站| 91麻豆精品激情在线观看国产| 国产在线观看jvid| 日韩精品免费视频一区二区三区| 欧美av亚洲av综合av国产av| 日韩成人在线观看一区二区三区| 自线自在国产av| 午夜精品国产一区二区电影| 女同久久另类99精品国产91| 色老头精品视频在线观看| 精品少妇一区二区三区视频日本电影| 侵犯人妻中文字幕一二三四区| 91九色精品人成在线观看| 色婷婷久久久亚洲欧美| 老熟妇乱子伦视频在线观看| 岛国在线观看网站| 757午夜福利合集在线观看| 手机成人av网站| 亚洲精品一区av在线观看| 亚洲精品美女久久av网站| 麻豆国产av国片精品| 亚洲天堂国产精品一区在线| 亚洲国产精品合色在线| 波多野结衣高清无吗| 免费av毛片视频| 亚洲人成电影观看| 欧美另类亚洲清纯唯美| 亚洲精品av麻豆狂野| 久久香蕉精品热| 欧美在线黄色| 俄罗斯特黄特色一大片| aaaaa片日本免费| 怎么达到女性高潮| 一边摸一边抽搐一进一出视频| 黑人巨大精品欧美一区二区蜜桃| 中文字幕人妻丝袜一区二区| 18禁裸乳无遮挡免费网站照片 | 久久久久久人人人人人| 中文字幕人妻丝袜一区二区| 大香蕉久久成人网| 久久人妻av系列| 亚洲欧美激情在线| 国产欧美日韩精品亚洲av| 三级毛片av免费| 99久久99久久久精品蜜桃| 亚洲av成人不卡在线观看播放网| 变态另类丝袜制服| 一级作爱视频免费观看| 久久人人爽av亚洲精品天堂| 国产精品免费视频内射| 久久国产精品男人的天堂亚洲| 黄频高清免费视频| 久久久国产成人精品二区| 亚洲中文日韩欧美视频| 亚洲专区中文字幕在线| 国产精品爽爽va在线观看网站 | 亚洲国产精品sss在线观看| 成人免费观看视频高清| 欧美日韩一级在线毛片| 国产高清videossex| 午夜激情av网站| 久久婷婷人人爽人人干人人爱 | 丝袜美足系列| 大型黄色视频在线免费观看| 久久久久久国产a免费观看| 色av中文字幕| 久久香蕉精品热| 日本精品一区二区三区蜜桃| 给我免费播放毛片高清在线观看| 人妻丰满熟妇av一区二区三区| 一进一出抽搐gif免费好疼| 色综合站精品国产| 两人在一起打扑克的视频| 精品高清国产在线一区| 女人被躁到高潮嗷嗷叫费观| 国产一区二区激情短视频| 69精品国产乱码久久久| 啦啦啦观看免费观看视频高清 | 久久久久久久久久久久大奶| 熟女少妇亚洲综合色aaa.| 变态另类成人亚洲欧美熟女 | 美女 人体艺术 gogo| 看片在线看免费视频| 国产野战对白在线观看| 亚洲精品中文字幕一二三四区| 欧美中文日本在线观看视频| 久久久久国产精品人妻aⅴ院| 啦啦啦免费观看视频1| 久久人妻福利社区极品人妻图片| 国产单亲对白刺激| 欧美 亚洲 国产 日韩一| 亚洲国产看品久久| 欧美成人性av电影在线观看| 亚洲精品国产一区二区精华液| 国产精品免费视频内射| 欧美激情 高清一区二区三区| 亚洲精华国产精华精| 级片在线观看| 黄网站色视频无遮挡免费观看| 国产精品一区二区三区四区久久 | 999久久久国产精品视频| 欧美日韩中文字幕国产精品一区二区三区 | 99久久久亚洲精品蜜臀av| 亚洲av成人不卡在线观看播放网| 欧美激情高清一区二区三区| 一进一出抽搐动态| 久久精品亚洲精品国产色婷小说| 免费少妇av软件| 99国产精品一区二区蜜桃av| 国产成人一区二区三区免费视频网站| 好看av亚洲va欧美ⅴa在| 国产主播在线观看一区二区| 91国产中文字幕| 久久青草综合色| 女人高潮潮喷娇喘18禁视频| 亚洲av第一区精品v没综合| 国产精品久久电影中文字幕| 亚洲精品一卡2卡三卡4卡5卡| 在线免费观看的www视频| 亚洲人成电影免费在线| 丝袜美足系列| 午夜影院日韩av| 视频在线观看一区二区三区| 亚洲第一电影网av| 日韩视频一区二区在线观看| 亚洲av电影不卡..在线观看| 麻豆成人av在线观看| 国产欧美日韩精品亚洲av| 少妇的丰满在线观看| 亚洲人成网站在线播放欧美日韩| 国产一区二区三区综合在线观看| 欧美日韩福利视频一区二区| 欧美绝顶高潮抽搐喷水| 午夜福利,免费看| 成年女人毛片免费观看观看9| 国产乱人伦免费视频| 91成年电影在线观看| 久久久久久亚洲精品国产蜜桃av| 亚洲一区中文字幕在线| 国产精品久久久人人做人人爽| 亚洲色图 男人天堂 中文字幕| 中文字幕精品免费在线观看视频| 精品人妻在线不人妻| 后天国语完整版免费观看| 香蕉久久夜色| 精品国产美女av久久久久小说| 18禁裸乳无遮挡免费网站照片 | 丝袜美腿诱惑在线| 自拍欧美九色日韩亚洲蝌蚪91| 在线观看www视频免费| av在线天堂中文字幕| 国产精品 欧美亚洲| 12—13女人毛片做爰片一| 91成年电影在线观看| 精品欧美一区二区三区在线| 亚洲午夜精品一区,二区,三区| 人人妻人人澡欧美一区二区 | 两性夫妻黄色片| 亚洲精品国产一区二区精华液| 色综合婷婷激情| 国产一级毛片七仙女欲春2 | 成人精品一区二区免费| 亚洲成人免费电影在线观看| 日韩 欧美 亚洲 中文字幕| 女生性感内裤真人,穿戴方法视频| av网站免费在线观看视频| 国产麻豆69| 精品国产乱子伦一区二区三区| 午夜两性在线视频| 午夜福利欧美成人| 久久中文字幕一级| 老司机午夜十八禁免费视频| 正在播放国产对白刺激| 亚洲午夜精品一区,二区,三区| 国产三级在线视频| 高清黄色对白视频在线免费看| 人妻丰满熟妇av一区二区三区| 天天躁狠狠躁夜夜躁狠狠躁| 夜夜爽天天搞| 99久久99久久久精品蜜桃| 婷婷六月久久综合丁香| 非洲黑人性xxxx精品又粗又长| 777久久人妻少妇嫩草av网站| 国产成年人精品一区二区| 亚洲男人的天堂狠狠| 日韩精品免费视频一区二区三区| 99久久综合精品五月天人人| 99久久久亚洲精品蜜臀av| 搡老妇女老女人老熟妇| 人人澡人人妻人| 97人妻精品一区二区三区麻豆 | www.自偷自拍.com| 久久九九热精品免费| 国产精品99久久99久久久不卡| 俄罗斯特黄特色一大片| 啦啦啦韩国在线观看视频| 日本 av在线| 操美女的视频在线观看| 成人特级黄色片久久久久久久| 黄频高清免费视频| 国产亚洲精品av在线| 国产黄a三级三级三级人| svipshipincom国产片| 美女免费视频网站| 国产精品亚洲av一区麻豆| 午夜精品国产一区二区电影| 禁无遮挡网站| 亚洲人成网站在线播放欧美日韩| 久久人妻熟女aⅴ| 国产成人精品久久二区二区免费| 亚洲激情在线av| 欧美一级毛片孕妇| 韩国精品一区二区三区| 亚洲精品久久国产高清桃花| 超碰成人久久| 天天添夜夜摸| 级片在线观看| 亚洲国产精品999在线| 99香蕉大伊视频| 精品少妇一区二区三区视频日本电影| 免费看美女性在线毛片视频| 亚洲avbb在线观看| 精品人妻1区二区| 热re99久久国产66热| 此物有八面人人有两片| 给我免费播放毛片高清在线观看| 99国产精品一区二区三区| 亚洲人成77777在线视频| 精品久久久久久久久久免费视频| 久久人人精品亚洲av| 日本一区二区免费在线视频| 一边摸一边做爽爽视频免费| 亚洲精品中文字幕一二三四区| 一级毛片高清免费大全| 18禁国产床啪视频网站| 久久伊人香网站| 中文字幕人成人乱码亚洲影| 91麻豆av在线| 精品一品国产午夜福利视频| 色尼玛亚洲综合影院| 精品人妻在线不人妻| 精品久久久久久成人av| 美女午夜性视频免费| 免费不卡黄色视频| 色老头精品视频在线观看| 十八禁网站免费在线| 久久午夜综合久久蜜桃| 在线永久观看黄色视频| 精品久久久久久成人av| 久久国产乱子伦精品免费另类| 久久久久久久久久久久大奶| 午夜免费鲁丝| 国产精品二区激情视频| 少妇的丰满在线观看| 亚洲成人免费电影在线观看| 久久久久国内视频| 麻豆成人av在线观看| 一级片免费观看大全| 久久国产精品影院| 久久九九热精品免费| 每晚都被弄得嗷嗷叫到高潮| 精品熟女少妇八av免费久了| 日本免费一区二区三区高清不卡 | 精品少妇一区二区三区视频日本电影| 一本久久中文字幕| 丝袜在线中文字幕| 午夜福利视频1000在线观看 | 一边摸一边抽搐一进一小说| 亚洲中文字幕日韩| 午夜免费激情av| 亚洲 欧美 日韩 在线 免费| 一区在线观看完整版| 日韩免费av在线播放| 极品人妻少妇av视频| 久久国产乱子伦精品免费另类| 欧美中文日本在线观看视频| 亚洲国产毛片av蜜桃av| svipshipincom国产片| 亚洲av美国av| 少妇的丰满在线观看| 日韩精品免费视频一区二区三区| svipshipincom国产片| 日韩欧美三级三区| 欧美最黄视频在线播放免费| 久久久久久大精品| 欧美乱色亚洲激情| 久久亚洲真实| 久久香蕉精品热| 好男人在线观看高清免费视频 | 97碰自拍视频| 亚洲欧美精品综合久久99| 国产xxxxx性猛交| 久久影院123| 国产亚洲欧美在线一区二区| 色综合婷婷激情| 亚洲欧美一区二区三区黑人| 欧美精品亚洲一区二区| 中国美女看黄片| 亚洲三区欧美一区| 午夜福利视频1000在线观看 | 男女下面插进去视频免费观看| 麻豆成人av在线观看| 好男人在线观看高清免费视频 | 亚洲国产精品sss在线观看| 法律面前人人平等表现在哪些方面| 国产精品久久久人人做人人爽| 人人澡人人妻人| 欧美激情极品国产一区二区三区| 99国产精品99久久久久| 叶爱在线成人免费视频播放| 欧美国产精品va在线观看不卡| 亚洲五月天丁香| 悠悠久久av| 这个男人来自地球电影免费观看| 国产成人欧美| aaaaa片日本免费| 国产一区在线观看成人免费| 日本在线视频免费播放| 宅男免费午夜| 可以在线观看毛片的网站| 一个人免费在线观看的高清视频|