Forest Ecology and Management 326 (2014) 125–132 Contents lists available at ScienceDirect Forest Ecology and Management journal homepage: www.elsevier.com/locate/foreco An allometric model for estimating DBH of isolated and clustered Eucalyptus trees from measurements of crown projection area Niva Kiran Verma a,b, David W. Lamb a,b,⇑, Nick Reid a,c, Brian Wilson a,c a Cooperative Research Centre for Spatial Information (CRCSI), University of New England, Armidale, NSW 2351, Australia Precision Agriculture Research Group and School of Science and Technology, University of New England, Armidale, NSW 2351, Australia c School of Environmental and Rural Science, University of New England, Armidale, NSW 2351, Australia b a r t i c l e i n f o Article history: Received 26 March 2014 Received in revised form 3 April 2014 Accepted 4 April 2014 Keywords: Diameter at breast height Crown projection area Scattered trees Tree clusters Eucalyptus Farm land a b s t r a c t Owing to its relevance to remotely-sensed imagery of landscapes, this paper investigates the ability to infer diameter at breast height (DBH) for five species of Australian native Eucalyptus from measurements of tree height and crown projection area. In this study regression models were developed for both single trees and clusters from 2 to 27 stems (maximum density 536 stems per ha) of Eucalyptus bridgesiana, Eucalyptus caliginosa, Eucalyptus blakelyi, Eucalyptus viminalis, and Eucalyptus melliodora. Crown projection area and tree height were strongly correlated for single trees, and the log-transformed crown projection area explained the most variance in DBH (R2 = 0.68, mean prediction error ±16 cm). Including tree height as a descriptor did not significantly alter the model performance and is a viable alternative to using crown projection area. The total crown projection area of tree clusters explained only 34% of the variance in the total (sum of) the DBH within the clusters. However average crown projection area per stem of entire tree clusters explained 67% of the variance in the average (per stem) DBH of the constituent trees with a mean prediction error ±8 cm. Both the single tree and tree cluster models were statistically similar and a combined model to predict average stem DBH yielded R2 = 0.71 with a mean prediction error (average DBH per stem) of ±13 cm within the range of 0.28–0.84 m. A single model to infer DBH for both single trees and clusters comprising up to 27 stems offers a pathway for using remote sensing to infer DBH provided a means of determining the number of stems within cluster boundaries is included. Ó 2014 Elsevier B.V. All rights reserved. 1. Introduction Much of eastern Australia is characterised by diverse farming landscapes, or ‘farmscapes’ containing a range of land-use systems including crops, native and sown pasture, remnant vegetation and trees at various densities and configurations. Native eucalyptus trees, both individual and in clusters are an important feature of Australian farmscapes, and provide shade and shelter for livestock (Bird et al., 1992). They also have considerable value through their influence on soils (Wilson, 2002; Goh et al., 1996; Barnes et al., 2011a), biodiversity (Oliver et al., 2006) and their indirect role in pasture quantity and quality (Barnes et al., 2011b). Scattered trees outside the forest (STOF) contribute significantly to above and below-ground carbon stocks in these landscapes (Baffetta et al., 2011; Soto-Pinto et al., 2010). A summary of the role of scattered ⇑ Corresponding author at: Cooperative Research Centre for Spatial Information (CRCSI), University of New England, Armidale, NSW 2351, Australia. Tel.: +61 2 6773 3565. E-mail address: dlamb@une.edu.au (D.W. Lamb). http://dx.doi.org/10.1016/j.foreco.2014.04.003 0378-1127/Ó 2014 Elsevier B.V. All rights reserved. trees in landscapes is given in Manning et al. (2006). The assessment of biomass in eucalypt systems has, to date, been largely restricted to plantation forestry systems. However there is also a growing need to assess carbon and biomass stocks across our farmscapes in order to fully quantify carbon storage change in response to management and provide evidence-based support for carbon inventory and ultimately carbon trading. Such assessments must also include scattered native trees. Destructive sampling is considered the most reliable method for determining biomass in grass and shrub vegetation but it is rarely used for agroforest ecosystems (Snowdon et al., 2002). The biomass of large vegetative structures like forests and open woodland is usually estimated by applying ratio or regression methods using empirical equations and allometric models (Snowdon et al., 2002; Houghton, 2005; Makungwa et al., 2013). Trunk diameter, more formally known as diameter at breast height (DBH), is observed to be closely related to tree biomass and carbon stocks (Ter-Mikaelian and Korzukhin, 1997; Snowdon et al., 2002; Sanquetta et al., 2011a; Kuyah et al., 2012; Beets et al., 2012). 126 N.K. Verma et al. / Forest Ecology and Management 326 (2014) 125–132 DBH is an important tree characteristic in its own right and is straight forward to measure on the ground. The relationships between tree canopy characteristics such as diameter, projection area and coverage, and DBH is of considerable interest as DBH can then be used to infer canopy attributes, for example to quantify canopy competition in response to planting (stem) density, growth potential or habitat modelling (Bella, 1971; Cade 1997; Grote 2003). Of course, large-scale collection of DBH data can be time consuming and now satellite or airborne remote sensing, including LIDAR can be used to infer canopy characteristics such as crown projection area (e.g. Franklin and Strahler, 1988; Leckie et al., 2003; Popescu and Wynne, 2004; Lee and Lucas, 2007). Measuring the canopy diameter or crown projection area from the ground involves measuring the crown projection across different angular segments of the canopy. These angular segments can either be based on fixed (pre-set) directions (Röhle and Huber, 1985), or in the case of highly asymmetric canopies the directions can be adapted to adequately characterise the actual tree dimensions (Hemery et al., 2005; Fleck et al., 2011). Based on an investigation involving 161 trees in an old-growth forest (approx. density 392 stems per hectare), Fleck et al. (2011) recommend the 8-point, flexible approach be used to estimate crown projection area in order to minimise errors due to deviations from canopy symmetry. Other workers have concluded that two, orthogonal diameter measurements (4 radii) are suitable for computing crown diameter (Hemery et al., 2005), from which crown projection area can be subsequently derived. The relationship between DBH and crown diameter or projected area has been the subject of numerous papers in recent years, of which a few exemplars will be discussed here. Unsurprisingly, the exact form of the relationship is driven by competition with neighbouring trees and understorey and much of this driven by factors such as shade tolerance. Many of the relationships are ‘almost’ linear, with observed departures from linearity, for example logarithmic or square-root dependence, often at smaller DBH (Hall et al., 1989; Hemery et al., 2005). Arzai and Aliyu (2010) found statistically significant linear relationships between crown diameter and DBH in some Eucalypt and other tree species from the savana zone in Nigeria (R2 0.23–0.82), as did Sanquetta et al. (2011b) for Araucaria (Araucaria angustifolia), Imbuia (Ocotea porosa) and Canelas (Nectandra grandiflora) in the mid southern Parana State of Brazil (R2 0.47–0.78). On the other hand, O’Brien et al. (1995) observed the good species-dependent predictions using log–log transformed data (R2 > 0.86), and Sanquetta et al. (2011b) also improved their prediction, but only for the combined species dataset, by a log–log transformation (R2 from 0.23 to 0.52). Smith (2008) observed a strong linear relationship between canopy area and trunk cross-sectional area (proportional to DBH2) for native pecan trees grown in managed groves (Smith, 2008) and Gill et al. (2000) observed DBH and DBH2, when used as a sole independent variable, to be reasonably good at predicting canopy radius in forest planted conifer trees (R2 > 0.45). Ultimately, models relating crown projection area and DBH are genus dependant, and often species dependent owing to the range of tree architecture between species. Moreover, little is known about the transferability of models derived for single trees (for example in open woodlands) as compared to clusters of trees (i.e., multiple stems) in light of possible competition in growth (e.g. Biging and Dobbertin, 1992; Biging and Dobbertin, 1995; Moeur 1997). Crown projection area could therefore be used to infer DBH. This offers an important pathway to deploy remote sensing tools for the large-scale assessment of above ground or biomass stocks across entire landscapes. From a remote sensing perspective, crown projection area is more easily measured than canopy diameter for the simple reason that the extracted crown diameter parameter is influenced by the direction of the measurement vector. Assuming image pixels, or objects are correctly classified as a particular tree crown, and the tree crown is at the nadir viewpoint (or close to it), it remains to sum the pixels (of known dimensions) within that crown to determine the projected area. This is not to say, however that crown diameter is irrelevant to the remote estimation of DBH. Numerous workers have found strong relationships between crown diameter and DBH (for example Hall et al., 1989; Gering and May, 1995) and given established sampling regimes for determining crown diameter (or canopy extent) are aimed at minimising the effect of crown asymmetry (Fleck et al., 2011), crown diameter could potentially be derived from the remotely sensed canopy projection area measurements by applying appropriate shape parameters such as discussed in the various work of Nelson (1997); Grote (2003) and Fleck et al. (2011). The present study aims to develop an allometric model for predicting DBH of single trees and trees in clusters using the tree/cluster characteristics of crown projection area and tree height. The overall context of this study is remnant, native vegetation that exists in typical Australian farming landscapes. In this study, we examine trees from the genus Eucalyptus, owing to its overall significance in Australian landscapes (Commonwealth of Australia, 1999) and in particular its prevalence in Australian farmscapes (Attiwill and Adams, 1996). In this study we hypothesize that DBH can be predicted using field measurements of tree height and crown projection area for both single trees as well as clusters of trees. We also wish to test the hypothesis that the same model is applicable for both single trees and isolated clusters of trees. The definition of a cluster can be somewhat arbitrary, and proximity of trees to one another will influence more than just competition for growth. For example Barnes et al. (2011b) demonstrated the extent to which the litterfall from an isolated tree will influence soil nutrients beyond the immediate canopy envelope. Moreover, given the context of this particular work in relation to the use of large scale remote sensing tools to possibly infer DBH, consideration should be given to the spatial resolution of such large scale sensing systems. To this end we define a cluster as a group of trees with canopy envelopes within 3 m proximity to each other. 2. Materials and methods 2.1. Study area The study site was located within the University of New England’s, ‘Newholme-Kirby SMART farm’, Armidale, New South Wales, Australia (longitude 151°350 4000 E to 151°370 1200 E and latitude 30°260 0900 S to 30°250 120 0 S) (Fig. 1). The 662 ha site comprises large tracts of natural eucalypt woodland and pasture cover. Approximately one third of the study area is dense eucalypt woodland, one third open woodland, and the remainder native pasture. Most of the study site is unimproved pasture grazed by sheep. The soils within the study site vary from brown and yellow chromosols, and the mean annual rainfall is 780 mm. It has a cool temperate climate with the majority of rain falling in the summer months (National Parks and Wildlife Service, 2003; BoM, 2014). 2.2. Field measurements High spatial resolution (15 cm), colour infrared (CIR) airborne imagery of the study area was first used to identify single trees and tree clusters in the field. A total of 52 tree clusters and 172 individual trees were identified (Fig. 2). Within the tree clusters the number of stems ranged from 2 to 27 with a density ranging from 38 to 536 stems per ha (SD 4.2 trees per cluster, 105 stems per ha). The tree/cluster locations were extracted from the orthorectified, georeferenced imagery for subsequent field visitation, N.K. Verma et al. / Forest Ecology and Management 326 (2014) 125–132 127 Fig. 1. Location map of the study site in north eastern NSW, Australia. Source: open access data. Fig. 2. Tree and tree cluster locations (white circles) overlaid on a grey-scale aerial image of the field site. aided by a DGPS (GPS PathfinderÒ Pro XRS receiver, Ranger TSC2 model, Trimble, California). The horizontal accuracy of GPS was 0.5 m allowing unambiguous identification of target trees/clusters on the ground. On-ground visitation and measurement of selected trees and tree clusters was conducted during the period September–December 2012. The DBH of individual trees and those in clusters was derived from the measured trunk circumference at 1.3 m above local ground level. In the case of individual trees (as delineated by a single root-ball) with multiple stems at 1.3 m above the ground, the DBH of each stem was measured and tree DBH calculated using: DBH ¼ pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi d1 þ d2 þ d3 For individual trees, the height (Ht) was measured using a combination of laser rangefinder and clinometer (MDL LaserAce 300, Measurement Devices Ltd. Scotland, UK). Cluster height was determined by first measuring the height of each tree within the cluster, and the average height (Htav) calculated. In those clusters where the individual tree crowns could not be delineated, Htav was calculated from six measurements of the entire canopy envelope acquired from different azimuthal viewer positions relative to the cluster. Canopy cover is defined as the proportion of ground covered by the vertical projection of the tree crowns (Jennings et al., 1999). In order to measure crown projection area (CA) of both single trees and tree clusters, we adopted a hybrid version of the 8-point crown projection method recommended by Fleck et al. (2011) and that investigated by Röhle and Huber (1985). Firstly a pair of vertical range poles were placed in the ground to delineate the edges of the tree/cluster canopy along a cardinal direction, for example N–S, passing through the tree stem or the estimated centre of the cluster. The crown periphery for locating the pole positions was located using a clinometer set to ‘vertical view’, in effect acting as a crown mirror. The crown diameter (d) along this direction was then measured using a laser range finder positioned at right-angles to, and a known distance well back from, the line between the poles. This measurement avoided errors that would otherwise be incurred by using tapes to measure the straight-line distance between the poles with tree stems in between. This measurement was undertaken for six cardinal directions namely N, ENE, ESE, S, WSW and WNW, respectively and the average diameter, which is effectively the average crown spread, d, calculated (Sumida and Komiyama, 1997). The crown projection area was then calculated using: 2 ð1Þ where d1, d2, d3 were the diameters of each stem (Pretzsch, 2009). The DBH parameters investigated for tree clusters were the average DBH of the individual trees within a cluster (DBHav) and the sum of the DBH values (DBHsum). CA ¼ pd =4 ð2Þ The crown projection area parameters investigated for tree clusters were the average of the individual trees within a cluster (CAav) and the total area of the cluster (CAsum). The information about tree species for each tree/cluster was noted as was the number of tree stems in a cluster. The dimensions 128 N.K. Verma et al. / Forest Ecology and Management 326 (2014) 125–132 Table 1 The allometric models tested for crown projection area (CA, m2), tree height (Ht, m) and DBH (m) for single trees and tree clusters. The subscript ‘av’ denotes the average value per stem within a cluster. The subscript ‘sum’ as applied to DBH is the sum of each value for the individual stems and when applied to CA is the size of the canopy envelope enclosing all the trees within the cluster. Regression models Individual trees Tree clusters DBH = a0 + a1(Ht,CA) ln(DBH) = a0 + a1ln(Ht, CA) ln(DBH) = a0 + a1ln(Ht) + a2ln(CA) DBHav = b0 + b1(Ht, CA)av ln(DBHav) = b0 + b1ln(Ht, CA)av ln(DBHav) = b0 + b1ln(Htav) + b2ln(CAav) DBHsum = b0 + b1CAsum ln(DBHsum) = b0 + b1ln(CAsum) Table 2 Summary statistics for single trees; n is the number of trees used in both the model development and validation. Tree characteristics/species n Min Max Mean SD DBH Apple box Red gum Stringy bark White gum Yellow box 23 5 51 28 65 0.34 0.5 0.36 0.36 0.28 1.65 0.83 1.33 1.92 1.47 0.840 0.690 0.839 0.911 0.807 0.310 0.151 0.221 0.391 0.244 Crown projection area (CA) Apple box Red gum Stringy bark White gum Yellow box 26.26 70.85 42.41 36.83 9.34 750 167 413 732 683 212 136 195 230 222 183.5 41.6 100.4 164.1 132.3 Tree height (Ht) Apple box Red gum Stringy bark White gum Yellow box 9.1 12.7 13.5 11.8 9.5 40.5 23.6 30.4 44.6 42.1 17.6 18.2 21.4 20.7 21.9 7.28 4.12 4.59 8.05 6.42 value per stem within the cluster. Similarly the subscript ‘sum’, as applied to DBHsum is the sum of each value for the individual stems and the parameter CAsum is the total size of the canopy envelope enclosing all the trees within the cluster. In the case of non-normal data distributions (Shapiro–Wilk Test), a logarithmic transformation was first carried out and the transformed data tested for species-specific variations. Linear regression models were evaluated using the statistical software R (Studio Version 0.97.318). In all the models, the interaction terms between the parameters were also tested. The data were tested for normality using Shapiro–Wilk test and influential outliers, if any, were detected by means of Cooks distance statistics of the residuals. Any data that had a residual Cook’s distance value of P2 was cross-checked with the original dataset to validate its precision and impact on the model. To satisfy the assumptions of linear regression analysis, scatter plots of residuals were checked for linearity, homoscedasticity and normality. The strength of the underlying relationship of the predictor and response variables was evaluated by analyzing the regression coefficients of the fitted models. The coefficient of determination (R2) was used to evaluate the level of variance in DBH explained by the variables. For each model, half of the samples, namely 86 for single trees and 26 for tree clusters, respectively, were withheld from the initial calibration for subsequent validation of the model. The prediction performance of each model was quantified using a mean prediction error (MPE) given by MPE ¼ jDBHpredicted DBHactual j. 3. Results and discussions 3.1. Single trees of five different Eucalyptus species, namely Apple Box (AB, Eucalyptus bridgesiana), Stringy Bark (SB, Eucalyptus caliginosa), Red Gum (RG, Eucalyptus blakelyi), White Gum (WG, Eucalyptus viminalis) and Yellow Box (YB, Eucalyptus melliodora) were sampled in this way. 2.3. Model development and validation 1.6 1.6 1.4 1.4 1.2 1.2 DBH (m) DBH (m) The allometric models tested for crown projection area (CA, m2), tree height (Ht, m) and DBH (m) for single trees and tree clusters are listed in Table 1 and follow the forms reviewed and listed in Hall et al. (1989). The subscript ‘av’ denotes measurements for tree clusters where DBHav, CAav and Htav are effectively the average Table 2 lists the descriptive statistics for the single trees. With all the species combined, the dataset was found to be non-normal, in particular the subset comprising White Gum and Yellow Box (Shapiro–Wilk test W = 0.97 p = 0.004). A logarithm transformation was sufficient to normalize the data (W = 0.992, p = 0.389). Scatter plots of DBH versus individual tree height (Ht) and crown projection area (CA) parameters are given in Fig. 3, along with regression curves of the log-transformed models. The regression statistics are summarised in Table 3. When the models were validated against the 86 trees retained from the data for this purpose, the log-transformed models yielded a MPE of 16 cm. A multi-linear regression model involving both log-transformed Ht and CA parameters combined gave a slightly improved prediction of DBH with a MPE of 14 cm. Interestingly, while log-transformed crown projection area on its own explains significantly more variance in log-transformed DBH than tree height (R2 = 0.68 compared to 0.31), both tree height and crown projection area predict DBH with a similar MPE (16 cm). This is a reflection of the strong inter-relationship between crown 1 0.8 0.6 0.4 0.6 0.4 R2 = 0.31 MPE= 0.16 m 0.2 1 0.8 R2 = 0.68 MPE= 0.16 m 0.2 0 0 0 10 20 30 Ht (m) (a) 40 50 0 200 400 600 800 CA (sq m) (b) Fig. 3. Scatter plots of DBH versus (a) tree height (Ht) and (b) crown projection area (CA) for single trees (all species, n = 172). The regression curves are the log-transformed models for each individual parameter (Table 5). 129 N.K. Verma et al. / Forest Ecology and Management 326 (2014) 125–132 Table 3 Derived regression parameters (95% confidence intervals) for single trees (n = 86). The MPE values are derived from a separate validation dataset (n = 86). Table 5 Derived regression parameters (95% confidence intervals) for tree clusters (n = 26). The MPE values are derived from a separate validation dataset (n = 26). Equation R2 Fstat p MPE (m) Equation R2 F-stat p MPE (m) ln(DBH) = 2.10229 + 0.61742 ln(Ht) ln(DBH) = 2.40568 + 0.42616 ln(CA) ln(DBH) = 2.64742 + 0.15142 ln(Ht) + 0.38002 ln(CA) 0.31 0.68 0.59 37.4 181.6 60.1 <0.0001 <0.0001 <0.0001 0.16 0.16 0.14 ln(DBHav) = 1.85617 + 0.42221 ln(Htav) ln(DBHav) = 2.13471 + 0.335344 ln(CAav) ln(DBHav) = 2.5813 + 0.18246 ln(Htav) + 0.31712 ln(CAav) ln(DBHsum) = 2.17648 + 0.523622 ln(CAsum) 0.14 0.67 0.69 3.8 46.2 24.7 0.07 <0.0001 <0.0001 0.10 0.08 0.07 0.34 11.9 <0.01 0.66 800 Apple Box R2 = 0.30 10 Red Gum 600 Stringy Bark 500 White Gum 9 R2 = 0.34 8 MPE= 0.66 m Yellow Box 400 7 DBHsum (m) CA (sq m) 700 300 200 100 6 5 4 3 0 5 15 25 35 45 2 Ht (m) 1 0 Fig. 4. Scatter plot of crown projection area (CA) against tree height (Ht) for single trees (n = 172, 5 species). The solid regression line (and R2) is based on a linear regression between the parameters. Table 4 Summary statistics for tree clusters (all species); n is the total number of trees used for model development and validation. Tree characteristics n Min Max Mean SD DBHav (m) Htav (m) CAav (m2) Number of stems Stem density (/ha) 52 0.28 9.12 18.66 2 37.91 1.10 26.50 443.45 27 535.91 0.56 18.23 111.75 5.71 150.71 0.17 4.10 78.30 4.20 105.11 projection area and tree height. The scatter plot of crown projection area versus tree height for all species (Fig. 4) illustrates this, with tree height explaining 30% of the variance in crown projection area. It can be concluded at this point that either Ht or CA could be used to infer DBH for single trees, although based on the level of variance explained in the DBH by CA, this latter parameter is likely to provide better precision in predicting DBH on a tree by tree basis. 3.2. Tree clusters 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0 0.0 R2 = 0.14 MPE= 0.10 m 5.0 10.0 15.0 20.0 25.0 30.0 400 600 800 1000 1200 CAsum (sq m) Fig. 6. Scatter plot of DBHsum versus the cluster crown projection area (CAsum) for tree clusters (n = 52). The regression curve is the log-transformed model (Table 5). average tree height (Htav) and average crown projection area (CAav) are given in Fig. 5, along with the curves based on the derived regression models. Regression statistics are summarised in Table 5. An assessment of the datasets showed that, again DBHav was not normally distributed (Shapiro–Wilk W = 0.953, p = 0.03147). Once log-transformed, the parameter CAav explained a significantly larger proportion of the variance observed in the average DBH per stem in each cluster as compared to the average tree height within the cluster (R2 = 0.67 compared to 0.14). Unlike the single trees, Htav and CAav were not strongly correlated (R2 = 0.04, data not shown), which suggests that a combination of both parameters in a multi-linear regression model (log-transformed inputs) may be desirable, even though this yields only modest gains in the level of variance explained in DBHav (R2 = 0.69 compared to 0.67) and precision in predicting values of DBHav (0.07 m compared to 0.08 m). The CAav parameter alone was found to yield a MPE of 0.08 m when predicting DBHav in the range from 0.28 to 0.84 m (28.5% and 9.5% error, respectively) The regression statistics for the sum of the DBH values within the clusters (DBHsum) and the total crown projected area (CAsum) of the clusters are also listed in Table 5 and a scatter plot of the DBHav (m) DBHav (m) Table 4 lists the descriptive statistics for the tree clusters. For the tree clusters, scatter plots of average DBH (DBHav) versus 200 0 0.9 0.8 0.7 0.6 0.5 0.4 0.3 0.2 0.1 0 R2 = 0.67 MPE= 0.08 m 0 50 100 150 Ht av (m) CA av (sq m) (a) (b) 200 250 300 Fig. 5. Scatter plots of DBHav versus (a) average tree height (Htav) and (b) crown projection area (CAav) for tree clusters (n = 52). The regression curves are the log-transformed models for each individual parameter (Table 5). 130 N.K. Verma et al. / Forest Ecology and Management 326 (2014) 125–132 the number of stems (R = 0.55). At the same time we observe the stem density (stems/ha) – CAsum relationship (Fig. 7) to take the form: 800 single tree envelope Stem density (/ha) 600 stem density ¼ 6007:1 ðCAsum Þ0:652 stem density = 6007.1 x (CAsum)-0.652 R² = 0.35 400 200 0 0 200 400 600 800 1000 1200 CAsum (sq m) Fig. 7. Scatter plot of stem density (/ha) versus total crown projection area (CAsum, m2) for tree clusters. The dashed curve is the envelope for the single tree data, calculated from the crown projection area (CA). The solid curve is represents the fitted power curve to the cluster data. Table 6 Derived regression parameters (95% confidence intervals) for a combined individual tree–tree cluster model. The parameters DBH and CA incorporate data for both single trees (DBH, CA) and the average values of each tree within the measured clusters (DBHav, CAav). The parameters DBH+ and CA+ incorporate data for both single trees (DBH, CA) and the sum of each tree within the measured clusters (DBHsum, CAsum); n = 112. The MPE values are derived from a separate validation dataset (n = 112). The percentages in brackets indicate the mean relative prediction error. Equation R2 F-stat p MPE (m) ln(DBH) = 2.46441 + 0.426425 ln(CA) ln(DBH+) = 3.73606 + 0.70211 ln(CA+) 0.71 265.0 <0.0001 0.59 157.0 <0.0001 0.13 (17%) 0.43 (31%) of DBHsum versus CAsum is given in Fig. 6. Again a log transformation provided the best regression model, although the total crown projected area only explained 34% of the variance in the total DBH (R2 = 0.34) with a MPE of 0.66 m where the DBHsum ranged from 1.06 to 8.91 m (63% and 7.4% error, respectively). The fact that the net crown projected area for a cluster is only weakly related to the sum of DBH values within that cluster is evidence of competition effects (Bella, 1971; White 1981). Fig. 6 exhibits increasing scatter at higher CAsum. In the clusters investigated in this work, a higher CAsum is correlated with an increase in The larger tree clusters in our farmscape are made up of fewer, older trees, consistent with the self-thinning rule discussed by White (1981). Superimposed on Fig. 7 is the curve generated by the single tree data; the so-called ‘single tree envelope’. Here the stem density (/ha) for the single trees was calculated using the ratio 10,000 m2/CA, effectively assuming the inter-stem distance is limited to the unperturbed canopy envelope of the individual trees themselves; namely zero influence overlap (Bella, 1971). It is evident that in our scattered tree clusters, the self-thinning mechanism is resulting in trees expressing similar canopy dimensions as isolated trees. With these older trees there is expected to be an increase in variability in canopy extent due to effects of weathering, pests and disease (for example Landsberg and Ohmart, 1989; Elliott et al., 1993; Neumann, 1993; Köstner et al., 2002) and this may also explain the increasing variability observed at higher values of CAsum. Summing the DBH in Fig. 6 effectively accumulates the effects of individual tree competition, and the resulting departure from the behaviour of single, isolated trees. The act of taking average (per stem) crown projected area and average DBH in a cluster most likely reduces this accumulating error. 3.3. Combining both single trees and tree cluster datasets A comparison of the single tree and tree cluster regression models based on crown projected area alone (DBH = f(CA); Tables 3 and 5) suggests the parameters derived for single trees, and those derived on a per-tree basis from tree clusters (stem numbers ranging from 2 to 27), are similar. A comparison of the derived regression equations for the individual trees and tree clusters shows the interaction terms to be non-significant (p = 0.79). This implies that the slopes and the intercepts of the two regression models do not differ significantly; statistically the two equations generate similar estimates of DBH. The log-transformed regression models derived using the combined datasets are given in Table 6 and scatter plots of the model-predicted versus actual DBH are given in Fig. 8. Here the models are based on the log-transformed DBH (DBH, DBH+) and the log-transformed crown projection area (CA, CA+) values. The superscript ‘’ denotes the fact that each parameter involves data for both single trees (DBH, CA), and the average values of each tree 1.6 4 single trees tree clusters single trees tree clusters 3.5 DBH+ predicted (m) 1.4 DBH* predicted (m) ð3Þ 1.2 1 0.8 0.6 0.4 R2 = 0.71 MPE= 0.13 m 0.2 3 2.5 2 1.5 1 R2 = 0.59 MPE= 0.43 m 0.5 0 0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 0 0 0.5 1 1.5 2 2.5 DBH* actual (m) DBH+ actual (m) (a) (b) 3 3.5 4 Fig. 8. Scatter plots of (a) measured versus predicted DBH⁄ (comprising DBH of single trees and DBHav of tree clusters) using both single tree CA and tree cluster CAav data in the regression model, and (b) measured versus predicted DBH+ (comprising both DBH of single trees and DBHsum of tree clusters) using both single tree CA and tree cluster CAsum data in the regression model (n = 112). Solid lines indicate 1:1 equivalence between predicted and actual values. N.K. Verma et al. / Forest Ecology and Management 326 (2014) 125–132 within the measured clusters (DBHav, CAav). The superscipt ‘+’ denotes the fact that each parameter involves data for both single trees (DBH, CA), and the net cluster projected canopy area and sum of DBH of each tree within the measured clusters (DBHsum, CAsum). Both models were created using a random selection of half the single and cluster data for calibration (n = 112) and the remaining data withheld for subsequent validation (n = 112). The data for the single tree and tree cluster datasets are shown as separate symbols. The effect of the error in the DBHsum versus CAsum, discussed earlier is again evident in Fig. 8(b), and it is clear that a single model for both individual trees and tree clusters breaks down when the DBHsum values are included in DBH+. In the DBH – CA model that incorporates both DBHav and CAav values of clusters with the data for single trees, the model performs well; it is noteworthy to observe the tree cluster data to be distributed amongst the single tree data points. The MPE in DBH for the tree-averaged data is 0.13 m. While encouraging, this equates to a mean relative prediction error approximately 17% of the DBH values encountered in the sampling. An investigation of the relative prediction error on a sample by sample basis did not show any systematic trend towards increasing prediction error with increasing DBH except for values exceeding 1.2 m. 4. Conclusions Simple regression models involving crown projection area of Eucalyptus trees (six species), both isolated and in clusters of up to 27 stems ranging from 38 to 536 stems per ha), explained 67% and 68%, respectively of the variance in stem DBH. A single model involving both single trees and the tree clusters to predict average stem DBH had similar explanatory power (R2 = 0.71) and yielded a mean prediction error in average DBH per stem of ±13 cm. We conclude that it is sufficient to use crown projection area to infer DBH for these species (and these stem densities and stem numbers). While the results appear encouraging, we acknowledge that the landscape investigated here was only 662 ha, and while encapsulating considerable variation in soils, elevation and aspect (for example as reported in Garraway and Lamb, 2011), it would be expected that the robustness and precision of the model could potentially be enhanced by including other landscape parameters. This is the subject of further work. Nevertheless, the use of larger scale data sources such as airborne or satellite imagery to infer DBH from derived values of crown projection area for both single Eucalypt trees and clusters up to 27 stems (between 22 and 536 stems per ha) appears feasible and worthy of further investigation. Admittedly, the single model developed here does require knowledge of the number of stems within a given tree cluster. However the increasing use of other sensing technologies like LIDAR to both infer total crown projection area and delineate and count the number of stems within a canopy (for example Popescu et al., 2004) potentially offers the means to achieve this. Acknowledgments This work was partially funded by the CRC for Spatial Information (CRCSI), established and supported under the Australian Government Cooperative Research Centres Programme. One of the authors (NKV) wishes to acknowledge the receipt of a Postgraduate ‘Top-up’ Scholarship from the CRCSI. 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