Classifying the Neotropical savannas of Belize using remote
Transcription
Classifying the Neotropical savannas of Belize using remote
Journal of Biogeography (J. Biogeogr.) (2006) 33, 476–490 SPECIAL ISSUE Classifying the Neotropical savannas of Belize using remote sensing and ground survey Neil Stuart1*, Timothy Barratt2 and Christopher Place1 1 Institute of Geography, School of Geosciences, University of Edinburgh, Edinburgh, UK and 2Tweed Forum, South Court, Drygrange Steading, Melrose, Roxburghshire, UK ABSTRACT Aim This paper evaluates a method of combining data from GPS ground survey with classifications of medium spatial resolution LANDSAT imagery to distinguish variations within Neotropical savannas and to characterize the boundaries between savanna areas and the associated gallery forests, seasonally dry forests and wetland communities. Location Rio Bravo Conservation Area, Orange Walk District, Belize, Central America. Methods Dry season LANDSAT data for 10 April 1993 and 9 March 2001 covering a conservation area of 240,000 acres (97,459 ha), were rectified to subpixel accuracy using ground control points positioned by GPS ground survey. The 1993 image was used to assess the accuracy with which the boundaries between the savanna matrix and gallery forests, high forests, wetlands and water bodies could be discriminated. The image was classified by a maximum likelihood (ML) classifier and the shapes and areas of forest and wetland classes were compared with an interpretation of these land cover types from 1 : 24,000 aerial photography, mapped at 1 : 50,000 scale in 1993. The 2001 image was used to assess whether different subtypes of savanna could be distinguished from LANDSAT data. This required the creation of a reference (‘ground truth’) data set for testing classifications of the image. One hundred and sixty sample patches (650 ha, distributed over an area of 7000 ha) of ten sub-types of savanna vegetation and associates identified using a physiognomic classification scheme, were delineated on the ground by GPS and divided into two subsets for training and testing. Continuous classifications of LANDSAT data covering the savannas were developed that estimated potential contributions from up to five sub-types of land cover (grassland, wetland, pine woodland, gallery forest and palmetto). The accuracy of each classification was assessed by comparison against ground data. An ML classification was also produced for the 2001 image using the same areas for training. This allowed a comparison of the relative accuracy of both continuous and Boolean ML methods for classifying savanna areas. *Correspondence: Neil Stuart, Institute of Geography, School of Geosciences, University of Edinburgh, Edinburgh, EH8 9XP, UK. E-mail: [email protected] 476 Results The boundary between savannas and evergreen forests, gallery forests and open water in the study region could be delineated by the ML classifier to within 2 pixels (60 m) using LANDSAT imagery. However, the constituent subtypes within the savanna were poorly discriminated. Whilst the shape and extent of closed canopy forest, gallery forest, wetlands and water bodies agreed closely with the distributions interpreted from aerial photography, classes such as ‘open pine savanna’ or ‘grassland’ were only 45–65% accurate when tested against ground data. A continuous classification, estimating the proportions of three savanna vegetation subtypes (grassland, marshland and woodland) present in each pixel, correctly classified more of the ground data for these cover types than doi:10.1111/j.1365-2699.2005.01436.x ª 2006 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi Classifying Neotropical savannas the comparable ML result. Proportional mixtures of the land cover estimated by the continuous classifier also compared realistically with the vegetation formations observed along ground transects. Main conclusions By using GPS, a ground survey of vegetation cover was accurately matched to remotely sensed imagery and the accuracy of delineating boundaries and classifying areas of savanna was assessed directly. This showed that ML classification techniques can reliably delineate the boundaries of savannas, but continuous classifiers more accurately and realistically represent the distribution of the subtypes comprising savanna land cover. By combining these ground survey and image classification methods, medium spatial resolution satellite sensor data can provide an affordable means for land managers to assess the nature, extent and distribution of savanna formations. Over time, using the archives of LANDSAT (and SPOT) data together with marker sites surveyed in the field, quantitative changes in the extents and boundaries of savannas in response to both natural (e.g. fire, hurricane and drought) and anthropogenic (e.g. cutting and disturbance) factors can be assessed. Keywords Belize, continuous classification, GPS survey, land cover classification, LANDSAT, remote sensing, savanna. INTRODUCTION The last 15–20 years has witnessed a resurgence of interest in the biogeography and ecology of savannas. The greater awareness of savanna biodiversity, the increasing number of endangered and endemic species being identified within savannas and an emerging consensus that savannas worldwide may represent a larger sink for carbon than Boreal forests (Scurlock & Hall, 1998) are among the factors raising awareness that this globally extensive land cover type may have been undervalued in the past and afforded until now a low priority for protection or conservation (Ratter et al., 1997). With recent studies identifying the potential for significant biomass to be stored within savanna woodlands, there is a growing need to map reliably the extent of savanna land cover, including all its component sub-types and associates, even (and perhaps, especially) where some of these are dominated by trees (Furley, 2004). Although the savannas of Central America have fewer woody species than the cerrado of South America, a phytogeographic analysis by Lenthall et al. (1999) of over forty savanna sites from southern USA to subtropical South America identified the savannas of Belize, the Yucatán and the Petén as a unique subgroup. The woody component of these savannas is defined by Pinus caribaea Morelet and in some locations, Quercus oleoides Schltdl and Cham. (Oak), as well as the characteristic shrubs Byrsonimia crassifolia (L. Kunth) (craboo), Curatella americana (L.) (Yaha) and the palmetto palm Aceolorraphe wrightii H.Wendl which is prevalent in frequently inundated savannas. Laughlin (2002), citing Furley (1999), asserts that ‘these pine savannas are unique amid the variety of Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd neotropical savannas’, whilst Bridgewater et al. (2002) emphasize the phytogeographical and ecological importance of this most northerly instance in Central America of humid, tropical lowland savanna. The study of related tree species by Brewer et al. (2003) also emphasized the biogeographical significance of this area, challenging the conventional view that the flora of Belize is simply derived from emigrant species from South America at the time of the Great American Interchange. They suggest that the hyperseasonality (Sarmiento, 1984) of the Belizean savannas and the shallow, oligotrophic soils give rise to a unique diversity of species. Thus, previous work has demonstrated a distinctive character for the Central American savannas, which may result from their distance from the richer, core areas in the Neotropics, from hyperseasonality and a distinctive flora adapted to the substrates and ground water conditions. The problem of delineating subtypes within the savanna biome remains as acute in Central as for South American counterparts, however, and this is critical in attempts to quantify the changes occurring to savanna associations at regional and global scales as a result of both natural and anthropogenic factors. Mapping savannas by remote sensing As savanna areas are fragmented by land development, their biodiversity and capacity to function as a sink for atmospheric carbon is reduced (Alho & Martins, 1995). Yet the extent to which Neotropical savanna areas are being cleared in different countries and regions is often unmeasured. Comprehensive mapping of savanna areas and their 477 N. Stuart, T. Barratt and C. Place constituent subtypes is therefore needed to evaluate their present condition and distribution and to assist in identifying priority areas for active management and conservation (Dougill & Trodd, 1999; M.D. Bitencourt et al., unpublished). The widespread distribution of Neotropical savannas, covering over 2 million km2 of South and Central America (Mistry, 2000) and their predominance in developing countries has meant that mapping efforts have generally been undertaken using remote sensing. Two recently completed projects are the Central American Ecosystems Maps funded mainly by the World Bank [in Belize using LANDSAT Thematic Mapper (TM) to delineate land cover at a map scale of 1 : 250,000; Meerman & Sabido, 2001] and the 1-km resolution Land Cover of North (and Central) America published in 2004 by the Canadian Center for Remote Sensing (CCRS) and the United States Geological Survey (USGS) as part of the Global Land Cover Mapping Project (Latifovic et al., 2002). Both projects used medium spatial resolution remote sensing [LANDSAT TM or SPOT VEGETATION (VGT)] to map land cover and monitor change at regional to continental scales in a cost-effective manner. In addition to these continental-scale mapping activities, some studies have explored the use of medium spatial resolution remote sensing to map the overall extent of savanna at more regional scales. In Brazil, Valeriano & Bitencourt (1988) used the Normalized Difference Vegetation Index (NDVI) to differentiate savanna areas based on their lower above-ground biomass than closed forest areas. A savanna land cover inventory was produced from an interpretation of 66 LANDSAT TM images, to identify changes in the areal extents and to assess the viability of savanna fragments remaining in Sao Paulo state (Kronka et al., 1997). This provided a useful quantification of overall loss of savanna, but as it did not differentiate between subtypes, it was not possible to ascertain from remote sensing whether clearance was greater for certain types of savanna (e.g. areas dominated by grassland or woodland). Also in Brazil, Almeida-Filho & Shimabukuro (2002) used the now substantial archives of LANDSAT sensor products to undertake a time-series analysis from 1987 to 1999 to monitor the gross changes in cerrado vegetation within Roraima state. Monitoring of the extent and nature of the savannas in the Neotropics has borrowed ideas from the approaches of workers in other parts of the world, such as Cahoon et al. (1992) who showed the potential of medium spatial resolution imagery to map the seasonal distribution of fires across parts of the African savanna. Trigg & Flasse (2000) developed means for wide-area monitoring of recently burnt savanna throughout Namibia using the SPOT VEGETATION (VGT) sensor and the Moderate Resolution Imaging Spectroradiometer (MODIS), while Stroppiana et al. (2003) showed how a time-series of SPOT VGT images could be used to detect recently burned areas in the savannas of the Australian Kakadu National Park. More comprehensive and current data about savanna land cover from remote sensing creates the potential for ecological 478 studies using digital methods of analysis at a variety of scales. As well as the possibility to monitor changes in the forest– savanna boundary over time, the associations theorized by biogeographers and ecologists such as Cole (1986), Solbrig (1991), Archer (1995), Kellman & Meave (1997) and Ludwig et al. (1999) between the internal savanna mosaics and factors such as topography, water availability, drainage, herbivory and fire history could, for example, be investigated more rigorously over time and space using a sequence of satellite sensor images, a digital elevation model (DEM) and other layers of data, such as the history of land management for an area that may be stored in a geographic information system (GIS). A pre-requisite for such mapping and monitoring is a reliable assessment of the accuracy with which the boundaries between and within savanna areas can be determined from remotely sensed imagery and in particular, from medium spatial resolution optical data, which is presently the most affordable for developing nations, where the majority of savannas are found. King (1994) has argued from experience of land cover mapping in the humid Neotropics, that when using LANDSAT data, only the broadest land cover types can often be discriminated accurately by conventional maximum likelihood (ML) classifiers and only when the optical imagery is sufficiently free from cloud and haze. Whilst for some investigations, detailed structural information about the composition of savannas is needed, which is unlikely to be extracted from medium spatial resolution sensor data (see, for example, Brown et al. (2004), we believe that recent improvements in GPS and in techniques of digital image classification offer the potential to extract further information from imagery such as LANDSAT about the characteristics of savanna boundaries and about the variations within savannas. Given the familiarity of land managers with LANDSAT data, the relative consistency of the Thematic Mapper sensor data and the continuity of global coverage since 1984, any method that improves the extraction of information about vegetation boundaries from these sensor data has significant practical value for the monitoring and reporting of present conditions and past changes. In this paper, we evaluate the potential for extracting information about both boundaries and internal conditions of savannas, choosing a test site in a developing nation where the local agencies have an urgent need to report on the distribution, extent and condition of the savanna under their conservation. LOCATION AND CONTEXT Belize lies approximately between 1552¢ and 1830¢ N and between 8728¢ to 8913¢ W. The land area of Belize is estimated by Meerman & Sabido (2001) at 22,963 km2, including some 1000 offshore cays. The climate is subtropical to tropical, with mean monthly minimum/maximum temperature ranging from 16/28 C in winter to 24/33 C in summer. There is a marked rainfall gradient from an annual average of 1200 mm in the north to 4000 mm in the south. The climate of Northern Belize is ‘tropical wet-dry’, following Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd Classifying Neotropical savannas km Figure 1 Extent of savanna based on the Ecosystems Map for Belize (Meerman & Sabido, 2001) and the location of the Hillbank (HB) and Booth River (BR) tracts within the Rio Bravo Conservation and Management Area. km Koeppen’s climatic classification, and a marked dry season occurs between February and May (Wright et al., 1959). Two main groups of savannas can be identified in Belize. These are the upland savannas of Mountain Pine Ridge and the lowland savannas of the northern and coastal plains. Both groups are found in oligotrophic conditions over coarse textured soils of low fertility, but the upland savannas are nearly always well-drained, whereas the lowland savannas frequently suffer from alternate seasonal flooding and desiccation, leading them to be classified as ‘hyperseasonal’ savannas (Sarmiento, 1984). Study site The study was conducted in the Rio Bravo Conservation and Management Area (RBCMA) in Orange Walk District of north-west Belize. This conservation area of 97,549 ha is almost entirely covered with natural forest, savanna and marsh. The RBCMA is one of the largest conservation areas in the country, accounting for over 4% of the land area and containing the largest lowland savanna under protection. Brokaw and Mallory estimated that lowland savanna covered 2.8% of the Rio Bravo (c. 2300 ha) in 1993. For management purposes, savanna areas in the eastern part of the reserve provide a ‘buffer zone’ that protects natural forests in the more inaccessible western part close to the Guatemalan border, from the increasing pressures outside the conservation area to clear woodland for subsistence agriculture and for cash-crop production (Programme for Belize, 2000). Figure 1 shows that, in Belize, savannas occur mostly on the gently rolling Northern and Coastal Plains. This area is geologically part of the Yucatán limestone platform which, during the Pliocene (2–13 Myr BP), was a shallow sea into which white marls, reddish clays, sands and gravels eroded from the granitic rocks of the Maya Mountains were deposited and accumulated to form the present low-lying (< 20 m) plain (Standley & Record, 1936). The lowland savannas are mostly found on lobes of these outwashed sands and gravels that are often nutrient poor and very well drained in the dry season but subject to varying durations of inundation during the wet season. This is illustrated by Fig. 2, based on observations by Wright et al. (1959) and re-drawn from Brokaw (2001). On these outwash deposits, undulating relief gives rise to locally varying drainage conditions and a repeating sequence of savanna subtypes is often observed, ranging from the savanna orchards in the lowest topographic positions that experience the longest periods of seasonal inundation, to the freely drained sandy ridges that are dominated by pine and oak. As Figure 2 Position of lowland savannas in Belize according to topography and substrate (after Brokaw, 2001). Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd 479 N. Stuart, T. Barratt and C. Place suggested by Fig. 2, the savannas are often bounded by wetlands and water bodies at low elevations, and at higher elevations by denser evergreen forests that develop on outcropping limestone parent material. The savannas may also be dissected internally by linear gallery forests that follow the lines of local water courses that cross these outwash deposits. The two main areas of savanna within the RBCMA are located to the east of the Booth River marsh (marked BR in Fig. 1) and to the east of the New River Lagoon between the old logging camp of Hillbank and the East Gate of the conservation area (marked HB). Whilst the remote sensing imagery we analysed had been geometrically rectified to allow mapping for the entire Rio Bravo area, in this paper we focus our analysis on the Hillbank savanna (nominal centre 1734¢ N, 8841¢ W) as the greatest amount of data has been collected for the vegetation of this area from a series of botanical and geographical surveys over the last 10 years. Previous mapping of Belizean savannas Although Lundell (1934, cited in Standley & Record, 1936) devised the first classification scheme for the regional vegetation of the Yucatán, the natural vegetation of Belize was mapped in detail for the first time by Wright et al. (1959). Wright’s classification interpreted soils, drainage and geomorphology to map the agricultural potential of the land, rather than the actual land use at the time. As such the 262,000 ha attributed to savanna and related pine forests (c. 11.4% land area) probably provides an upper estimate of upland and lowland savanna in the country. A new vegetation classification and mapping of vegetation cover in Belize was produced by Iremonger & Brokaw (1995); this was based on Wright’s classification but greater detail was provided by a visual interpretation of satellite sensor data. Intended for conservation planning, this map showed actual vegetation cover interpreted from SPOT panchromatic imagery from 1993 at a scale of 1 : 250,000. A total of 51 vegetation types were identified in this national vegetation classification, including a class for savanna. Field checking by the author revealed, however, that many savanna woodlands were classified within ‘lowland needle-leaf moist open forests over poor soils’. At the other end of the ecotone, seasonally inundated savanna orchards were included within ‘non-saline seasonally waterlogged scrubs’. This underscores the difficulty in estimating changes in savanna extents based on previous mapping, when the land cover type has either not been accorded a dedicated class, or the full extent of savanna is likely to have been underestimated. In 2001, the Ecosytems Map for Belize was completed as part of the Central American Ecosystems Map series. This map is in fact a revision of the Iremonger & Brokaw (1995) map, with the nomenclature adapted to conform to the UNESCO classification adopted for all Central American ecosystems maps and with the class definitions extended to include an altitudinal component (Meerman & Sabido, 2001). The revised 480 map depicts 78 terrestrial ecosystems at a scale of 1 : 250,000. The altitudinal component of the classification allowed lowland savanna to be identified as a major land cover type. Two subtypes were defined; both were types of short-grass land cover, one dominated by trees (V.A.2.a), and one by shrubs (V.A.2.b). Despite a re-interpretation of the original LANDSAT imagery to resolve greater spatial detail and further ground checking, the classification scheme still attributed denser areas of pine and oak woodland to a class (I.A.2.a) for tropical evergreen seasonal forest, although such formations are increasingly being included as ‘savanna woodlands’ in classifications that focus on savannas (e.g. Furley, 1999). The mapping of savanna land cover by remote sensing therefore needs to address a series of challenges concerning, among others, the definition of the land cover that will be considered and mapped as sub-types of savanna, the recognition of these classes in the field and the identification of their reflectance spectra in a satellite image. With such high levels of both endemism and variety of species in savannas in different regions, there is an emerging consensus that savanna land cover classes should be identified primarily on the basis of physiognomy (vegetation structures that can be recognized in the field) rather than their floristics (Eiten, 1982; Cole, 1986). Recent mapping of savannas as part of the Central American Ecosystems Maps and the Global Land Cover Mapping Project followed this approach, adopting the UNESCO system for physiognomic–ecological classification of plant formations, first drafted by Mueller-Dombois & Ellenberg (1974) and the FAO Land Cover Classification System (Di Gregorio & Jansen, 2000), respectively. The clarity with which land cover classes can be identified both in the field and at corresponding locations on a satellite sensor image is partly an issue of locational accuracy and of sensor resolution. Mismatch between the image and the ground survey data can be caused by uncertainty in locating ground control points (GCPs). This mismatch is a source of error that causes uncertainty in both the training and testing of land cover classifications from remote sensing imagery. When LANDSAT TM data had been corrected using GCPs obtained from topographic maps, Combs et al. (1996) found the maps derived from the imagery contained significant errors when locations, areas and distances were measured. Cook & Pinder (1996) improved the locational accuracy of mapping from remote sensing by using GPS to supply control points directly by ground survey. Stuart & Moss (2000) used GPS to supply the ground training and testing data, as well as the GCPs for detailed mapping of the Rio Bravo reserve from LANDSAT data in 2002. Once the imagery has been corrected using these higher accuracy GCPs and because selective availability was removed in 2000, field scientists can use single frequency handheld GPS receivers to position field sites accurately for re-visiting on maps derived from remote sensing imagery (McCormick, 1999). Two further difficulties that were reported when undertaking the ecosystems mapping project in Belize were the limited availability of LANDSAT imagery between 1996 and 1998 that Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd Classifying Neotropical savannas was sufficiently free of cloud to allow ground control points to be selected for image rectification, and problems of classifying the short grass savanna subtypes from the satellite sensor data (Meerman & Sabido, 2001). This illustrates the continuing difficulty of devising land cover classifications of the subtypes of savanna vegetation that can be used with a spectral classification from remotely sensed imagery. Using the classes of the revised Ecosystems map, Meerman & Sabido (2001) estimated that 8.8% of Belize (c. 200,000 ha) was covered by lowland savanna at the time of remote sensing data acquisition in 1993. The relatively small difference between this and Wright’s estimate in 1959 of 11.4% covered by upland and lowland savannas supports the general view that wholesale conversion of savanna to other land uses has not yet occurred in Belize and reinforces the opportunity for land management to conserve substantial, unfragmented areas of savanna. NGOs such as the Programme for Belize, which manages the RBCMA, are investing in technology such as geographic information systems (GIS) to aid in the management of their land area. Classifications from imagery such as LANDSAT TM are considered a vital and often the only source of contemporary, comprehensive information about the state of the land resources that can be obtained at an affordable price. The accurate reporting of land cover types, extents and condition can be a requirement placed on NGOs before they may claim grants from conservation agencies for the upkeep of conservation areas. In the RBCMA, for example, quantitative assessment and regular reporting of the extent of pine woodland savanna is required to secure continued funding for a carbon sequestration pilot project and for various bird conservation projects in this habitat (Programme for Belize, 2000). In sum, this growing need at both an international and local level to analyse the distribution of savannas using digital methods makes it timely to assess the quality of data that can be collected about the boundaries and internal conditions of savannas using affordable, medium spatial resolution remote sensing imagery. The extent to which the different savanna classes can be recognized from each other and from associated vegetation such as evergreen and deciduous forests, gallery forests and wetlands using LANDSAT TM data is the main question addressed in the remainder of this paper. DATA AND METHODS Physiognomic classification of savanna subtypes from vegetation survey The subtypes of savanna to be classified by remote sensing were defined from a survey of savanna vegetation in Belize by Bridgewater et al. (2002). Whilst the primary output of that survey was a detailed floristic inventory to assist local planning for savanna conservation, a physiognomic classification of savanna vegetation subtypes was developed as a by-product from the botanical data. The classification allowed the Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd distribution of different savanna subtypes to be studied in relation to variations in edaphic and drainage conditions across a much more extensive area of the RBCMA by using remote sensing (Furley et al., 2001). From botanical collecting conducted in the savanna and associated wetlands and gallery forests of the RBCMA from July to September 1996, with supplementary visits in March 2000 and March 2001, a total of 258 savanna species were recorded. Of these, 148 were characteristic of the drier savanna ecosystem, 47 were associated with seasonally inundated areas and 57 were woody species, of which 15 could be classified as trees (Bridgewater et al., 2002). For the purposes of land cover mapping, broad descriptions were then made of all the subtypes of savanna vegetation encountered. The dominant or defining species and general physiognomy of each subtype were recorded. Where possible, sub-types were defined to conform to land cover classes previously established by Brokaw & Mallory (1993) for the trees of the RBCMA and the generalized land cover types of the national vegetation classification by Iremonger & Brokaw (1995). When further subdivision was required, new subtypes of savanna were defined that complemented terminology already used locally, were easily distinguishable on the ground and had significance for management purposes. The vegetation was grouped broadly into three main categories of ‘savanna’, ‘wetland’ and ‘savanna–forest transition’, reflecting the geographic and topographic position of the savannas between forests and wetlands in this area. The savanna–forest transition class included all situations where savanna graded into forest. This included both denser pockets of forest within the savanna matrix and also gallery forests that fringed water courses within savanna areas. Because of the management priority for information about savanna areas, Bridgewater et al. (2002) did not subdivide the savanna–forest transition class, which had been previously studied in Belize by Wright & Baillie (1995), but focused on six subtypes of savanna and five differing types of associated wetland communities (Table 1). Remote sensing data A dry season LANDSAT 5 TM image of north-west Belize (Path 19, Row 48, nominal centre 1720¢ N, 8840¢ W) acquired on 10 April 1993 was used to investigate how reliably Table 1 Savanna and wetland vegetation classification for the RBCMA (Bridgewater et al., 2002) 1.0 Savanna 2.0 Wetland 1.1 Grassland and scrub grassland 1.2 Pine–palmetto savanna 1.3 Palmetto thicket 1.4 Savanna orchard 1.5 Woodland and Pine Ridge 1.6 Oak thicket 2.1 Fringing riverine red mangrove 2.2 2.3 2.4 2.5 Cutting grass marsh Marl flat Sedge marshland Eleocharis–calabash marsh 481 N. Stuart, T. Barratt and C. Place savanna boundaries could be delineated using the sensor data. A maximum likelihood classification of the 1993 image was undertaken using ERDAS Imagine (Leica Geosystems, Norcross, GA, USA) and the location of boundaries between savanna areas and gallery forests, open water and seasonally inundated marshlands was compared to a delineation of these three land cover types as interpreted from 1 : 24,000 aerial photography and mapped on the 1 : 50,000 topographic mapping of Belize. As this photo-interpretation was undertaken as a training exercise by the UK Military Survey, the topographic maps contained an unusual richness of detail about land cover types, with areas of high forest, marshland and open water all delineated. By accepting the boundaries and shape of the land cover types on this mapping as an independent source of higher accuracy, the relative accuracy with which savanna boundaries could be discriminated from LANDSAT data was assessed by extensive visual comparison. Although the topographic mapping provided a high quality source of ground truth for testing the classification of the land cover types forming boundaries with savanna in the 1993 LANDSAT data, the mapping could not be used to assess the accuracy of classification by LANDSAT within the savanna areas because subtypes of savanna had not been interpreted from the aerial photography. It was therefore necessary to collect independent reference data by a ground survey of vegetation types within the savanna and to obtain an image that was contemporaneous with the completion of this ground survey in Spring 2001. A relatively cloud-free LANDSAT TM image was acquired dated 9 March 2001. A sub-scene was extracted covering an area of c. 96 km east-west and 68 km north-south, including the RBCMA together with areas outside this reserve where there were clearly defined features (bridges, road junctions, airstrips, etc.) that could be used for GCPs. GPS survey of ground control and vegetation patches Coarse acquistion (C/A) code GPS receivers with postprocessed differential correction, were used to position the GCPs for rectifying the two satellite images. For the 1993 image, 48 control points were used to rectify the image into NAD27/UTM(16) projection datum and map projection for Belize. The accuracy of the rectification had an RMSE of 0.6 pixels (20 m). For the 2001 image the RMS error was 0.37 pixels (11 m). In both cases, further visual assessment was used to confirm that sub-pixel accuracy had been achieved in the image rectification. This was necessary to ensure correct matching between the locations of the field reference data and the corresponding pixels in the satellite image. The main sub-types of savanna vegetation and associates to be mapped were then defined. Several examples of each type were visited under the supervision of field botanists and land managers to ensure correct identification on the ground. Representative patches of each savanna subtype were then surveyed using a GPS receiver mounted on an all-terrain vehicle. Since this ground reference data would be matched to 482 image pixels of a minimum size of 30 m, vegetation patches were chosen that were each as homogenous as possible and typically of 1–2 km2 in extent. Remaining areas of vegetation that could not be encircled in this way were encircled on foot with a GPS receiver. Using this approach, 160 patches of vegetation, covering a total of 631 ha were encircled with c. 15 person-days of fieldwork. These areas of reference vegetation data were then divided into two subsets, with one used to train and the other to test classifications of the satellite data. Further details of the field survey methods, software and equipment used are described in Moss et al. (2003). Classifying savanna vegetation by remote sensing Given the difficulties anticipated in mapping a land cover type that is, by definition, an ecotone between the forest and grassland biomes (Longman & Jenik, 1992), we explored a two-tiered approach to classification of savanna land cover from remotely sensed imagery. The first stage used a conventional Boolean maximumlikelihood (ML) classifier to distinguish areas of high forest, gallery forest, wetlands and open water. The areas remaining to be classified within the study region were then primarily the savanna tracts. The savanna areas were then classified by linear spectral mixture analysis (LSMA) (Settle & Drake, 1993). An ML classification of the savanna areas was also undertaken to allow a comparison with this more widely used technique. Both classification methods made use of the same ground reference data to assign classes to image pixels and to evaluate the accuracy of the resulting classification. The main characteristic of the ML technique pertinent to this study is that the resulting classification is Boolean, meaning that each pixel is allocated to one and only one class – that to which it is estimated to have the greatest probability of membership. In contrast, LSMA creates a continuous classification in which each pixel has partial membership to many land cover classes. In the same way that the texture of a soil can be represented as its constituent proportions of sand, silt and clay on a triangular diagram, so mixture modelling seeks to ascertain the proportional contributions of certain vegetation ‘end-members’ into the reflectance observed for a given pixel in a remote sensing image. The rationale for using LSMA to ‘unmix’ the constituents of savanna land cover rests upon an acceptance that a continuous classification is more able to represent the range and diversity of vegetation subtypes that collectively create savanna formations. For example, several workers including Furley (1999) have identified at least five sub-formations to the savannas (cerrados) of Brazil, ranging from open grasslands (campo limpo) through to dense savanna woodlands (cerradão), which can contain trees of up to 20 m in height. An inclusive mapping of savannas should encompass all these subtypes. Compared to conventional Boolean classification, a continuous classification is less likely to misallocate pixels that inherently exhibit a mixed response arbitrarily into one or another discrete land cover classes. Continuous classification Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd Classifying Neotropical savannas may be more suitable to a range of natural landscapes for which the medium spatial resolution of the LANDSAT TM sensor may not be high enough to resolve natural spatial variability fully (Adams et al., 1995; Zhang & Stuart, 2001). Evidence from using ML techniques to classify the savannas of Belize suggests this may be the case (Waltham et al., 2002). In this study, ML classification was applied to the 1993 image primarily for the purpose of discriminating the high forest, gallery forest, wetlands and water bodies that border the savanna areas. A total of 97 vegetation patches surveyed in the field were used as training data. Characteristic spectra were generated for each of 10 land cover types for which training data were collected. These 10 cover types mostly accord directly with those listed in Table 1, although a larger amount of training data for the pine woodlands allowed two sub-types to be defined according to tree density. Training data were not collected for areas of inundated marl and mangrove as these areas could not be accessed on the ground. Separability of each of the class spectra was evaluated using the Jefferies–Matusita (JM) index of divergence, using various combinations of spectral bands (Waltham et al., 2002). Despite evidence that some of the savanna vegetation sub-types had overlapping spectral properties in the image, a classification by ML of the entire area was initially undertaken and the accuracy of the resulting map was evaluated using the remaining groundsurveyed patches of vegetation as independent test data. Following the ideas of Settle & Drake (1993), a series of LSMA models were then formulated to unmix the extract of the 2001 LANDSAT imagery of the savanna tract south of the Hillbank station (Fig. 1). Contributions from between three to five end-members were modelled. Five is the maximum number that may be defined when using a LANDSAT image with six spectral bands. The end-members defined were grass, marsh, woodland, palmetto and evergreen forest. This definition was based on the main physiognomic types of savanna and associated vegetation identified and for which ground training areas had already been surveyed. Other end-members could have been defined, such as photosynthetic, non-photosynthetic vegetation, bare ground or cloud shadow. Such generic end-members might have had more separable individual spectra and allowed a greater understanding of the contributions from different surface types to overall scene reflectance. End-members based on the savanna subtypes were used in this study because we wished to explore if the proportional contributions estimated by LSMA showed any agreement with the vegetation formations identified by the eco-botanical survey as comprising the savanna in this region. We also wished to use the same ground data for developing the training spectra for both the ML and continuous classifiers, to allow a direct comparison between the two techniques. A preliminary assessment of the image extract showed that cloud, shadow and bare ground accounted for only a small number of pixels. By overlaying the polygonal areas of vegetation types digitized from the field survey onto the LANDSAT image, pixels falling within these polygons were selected and used to compute the mean reflectance of each end-member in each of Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd six spectral bands. These mean reflectance values were then entered into the MIXMOD module of the public-domain mips software of Mather (1999). Spectral mixture analyses were conducted using different combinations of the savanna end-members. Each analysis produced a set of gridded images, one for each end-member used, giving the proportional contribution of that end-member (vegetation type) to each image pixel. For ease of interpretation, the proportions were constrained to sum to 100%. The accuracy of these estimated proportions was evaluated by inspecting the pixel values in areas where the actual land cover was known from the field survey to be dominated by a single vegetation type. These ‘purer’ patches of land cover should be estimated by LSMA to have a high proportion of only one end-member. The LSMA using the three end-members (grass, marsh and wood) was found to reproduce the degree of homogeneity of land cover observed in the tested areas most accurately. As well as testing this classification in areas where the land cover was known to be dominated by one vegetation type (e.g. an area of predominantly woodland), we also investigated if the variations in the end-member contributions estimated by LSMA agreed with observations of changes in the sub-types of savanna vegetation made along transects surveyed in the field. This was assessed by overlaying the co-ordinates of the transects onto the image and comparing whether changes in the proportions of grass, marsh and forest estimated for the image pixels conformed with any observed changes in savanna subtypes recorded along the ground. RESULTS AND DISCUSSION Classification of savanna boundaries and subtypes by maximum likelihood methods The MLC technique reliably discriminated the gallery forest, forest transition areas and standing water bodies found in the study area. This was confirmed by a systematic comparison of the results for extracts of the classified 1993 LANDSAT image (such as Fig. 3a) against reference data for the high forest, gallery forest and marsh land cover types interpreted from 1 : 24,000 aerial photography from 1993 and mapped at 1 : 50,000 (such as Fig. 3b). The location and shape of the boundaries of the gallery forests, forest thickets and open water bodies classified in the satellite sensor image were found in most cases to agree to within 1 or 2 pixels (30–60 m) with the boundaries interpreted from the aerial photography. To illustrate, comparison of Fig. 3(a) and 3(b) shows close agreement as to the shape of a distinct boundary between the savanna and the high evergreen forest in the south-east of this extract. The gallery forest following Big Pond Creek is clearly resolved by LANDSAT data and many smaller pockets of forest that were delineated from the photography as outliers within the savanna matrix are also correctly classified as isolated forest patches in the imagery. Many of the smaller seasonal ponds that appear on the aerial photography are also classified correctly by the LANDSAT data in Fig. 3(a). In contrast to the sharp boundary with evergreen forest, the 483 N. Stuart, T. Barratt and C. Place Figure 3 Part of the study area, showing (a) a Maximum Likelihood (ML) classification of the land cover and (b) topographic map of the same area as (a), interpreted from 1 : 24,000 scale air photography. Crown copyright 1993. Crown copyright map reproduced with the permission of the Controller of Her Majesty’s Stationery Office. boundary between savanna and wetlands is much less distinct in this study area. Whilst the extents of the Marsh and Savanna Orchard classes from LANDSAT correspond in general to the land interpreted from the photography as subject to inundation (blue stipple on the western side of Fig. 3b), the satellite sensor also detects thickets of at least 20 pixels (> 2 ha) within these seasonally inundated areas where there is a definite woody response (brown hues interspersed within the light blue areas on Fig. 3a). 484 Whilst the sharper savanna boundaries could clearly be discriminated from LANDSAT data using the MLC technique, classification of the sub-types of vegetation occurring within the savanna tracts was generally less successful. An assessment of the accuracy with which this area could be completely classified into Bridgewater et al.’s (2002) savanna vegetation subtypes and associates by maximum likelihood methods is reported in Table 2. The producer’s accuracy statistic indicates the percentage of pixels covering areas of known reference Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd Classifying Neotropical savannas Table 2 Accuracy assessment for classification of the savanna areas corresponding to Bridgewater’s classes by maximum likelihood method (after Waltham et al., 2002) Class name Unclassified (incl. 2.1, 2.3) Cutting grass marsh (2.2) Eleocharis marsh (2.5) Grassland (1.1) Oak–pine woodland (1.6) Palmetto thicket (1.3) Pine savanna (1.2) Pine wood – dense(1.5) Pine wood – open (1.5) Savanna orchard (1.4) Sedge marsh (2.4) Totals Reference Totals Classified Totals 0 45 54 188 164 43 248 299 26 276 37 1380 178 42 67 241 157 140 196 177 12 152 18 1380 Number Producers Users Correct Accuracy (%) Accuracy (%) 0 0 1 124 91 2 16 91 12 30 0 367 – 0 2 66 55 5 6 30 46 11 0 – 0 2 51 58 2 8 51 100 20 0 Overall classification accuracy ¼ 26.6%. vegetation type that were correctly allocated to that class. Whilst grasslands and some of the oak and pine woodlands were classified with 45–65% accuracy, the wetland, savanna orchard and palmetto classes were poorly separated by the sensor and as a result these classes were poorly distinguished. The conclusions are that, whilst a classification of LANDSAT data by conventional MLC techniques is suitable for delineating the overall extents of savannas and their boundaries with associated vegetation types, it does not reliably map the distribution of the component vegetation formations within savanna areas. With many conservation agencies using LANDSAT data because of its relative affordability and its long archive of data, these findings demonstrate the potential to use these data to monitor even relatively localized changes in some forest–savanna boundaries over the last 20 years. However, mapping and monitoring conditions within savanna areas will require either different sensor data, or a different way of classifying LANDSAT data. Continuous classification of savanna boundaries and subtypes Four classifications of the land cover within the savanna areas were produced using LSMA. Each classification used different numbers of end-members and estimated their contributions to the reflectance of each pixel. Table 3 reports the accuracy of these continuous classifications of savanna land cover. To illustrate, the classification using the three end-members (grass, marsh and wood) was tested against 281 pixels where the land cover was known from the GPS field survey to be predominantly open grassland. The reflectance of 191 pixels was estimated by LSMA to be constituted at least 50% by the grass end-member (i.e. the response was more like grassland than anything else). The accuracy for classifying pixels constituting > 50% grass is therefore 191/281 ¼ 68%. Since the classification estimates the proportional membership of a pixel to each of the land cover classes on a continuous scale (from 0% to 100%), it is necessary to specify the proportional membership at which the accuracy of a class is to be assessed. Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd For example, using the most stringent criteria that > 80% of a pixel response should be constituted from the grass endmember, only 98/281 pixels meet this condition, yielding a class accuracy for ‘>80% grass’ of 35%. For the same 281 test pixels, the number correctly classified as ‘grassland’ by the ML classification was 158 (56%). Adding further end-members into the LSMA did not improve the accuracy with which the test areas were represented in the resulting classifications. Model 4a, which included a fourth end-member for high forest, led to a slightly improved classification of marshlands but then underestimated the extent of woodlands. Model 4b included a fourth endmember for palmetto palm but achieved only 35% accuracy in discriminating palmetto and reduced the areas correctly classified as wood or marshland. Model 5, which included all the above sub-types, had the lowest classification accuracy for all classes, suggesting that LANDSAT data do not have this much dimensionality of information. This supports the conclusion reported by others, including King (1994), that intercorrelation of the visible red, blue and green bands of LANDSAT data effectively reduces the dimensionality to 3 or 4. Based on this assessment, the three end-member LSMA was accepted as offering the more accurate continuous classification. The accuracy of this classification was then investigated in greater detail. Table 3 shows that, by choosing a modest level of pixel purity, such as 40%, the LSMA using the three end-members correctly classified 77% of grasslands, 73% of marshlands and 64% of woodlands. The percentage of pixels allocated to their correct class declines gradually from 70% to 30% for the grassland and woodland classes as the required proportion of the one cover type contributing to the pixel is increased to an upper limit of 80%. Areas dominated by grassland were the most reliably distinguished of all savanna subtypes by LANDSAT data. This illustrates how the continuous classification contains information about both the extents and the relative certainty with which pixels have been allocated into a given class, providing an indication of the robustness of the classification. The final column of Table 3 presents comparable 485 N. Stuart, T. Barratt and C. Place Table 3 Accuracy of four linear spectral mixture analyses using different combinations of end-members, and a maximum-likelihood (ML) classification of the same test areas Table 4 Average (± 1 SE) proportions of each of the three endmembers estimated by LSMA for pixels in areas where the land cover is known from ground survey to be predominantly the ground cover type in the first column Percentage of test pixels classified by each set of end-members End-member proportion (%) Grass > 40 > 50 > 60 > 70 > 80 Marsh > 40 > 50 > 60 > 70 > 80 Wood > 40 > 50 > 60 > 70 > 80 Forest > 40 > 50 > 60 > 70 > 80 Palmetto > 40 > 50 > 60 > 70 > 80 Pixels in test areas 3 4a 4b 5 ML 281 77 68 63 52 35 49 27 20 15 9 86 78 67 52 34 26 15 11 7 4 56 486 73 21 16 12 11 99 28 22 16 12 51 23 18 14 9 55 27 21 15 11 42 1206 64 44 38 31 22 19 11 8 7 5 41 31 26 21 15 12 6 5 4 3 48 29 22 20 17 13 N/A 25 11 10 8 4 26 638 178 51 44 39 31 22 35 11 8 4 3 End-member combinations were – 3: grass, marsh, woodland; 4a: grass, marsh, woodland, evergreen forest; 4b: grass, marsh, woodland, palmetto; 5: grass, marsh, woodland, evergreen forest, palmetto. ML is the percentage of pixels in the test areas that were assigned to the correct ground class by the maximum-likelihood classifier. accuracies for an ML classification based on the same 10-class classification used by Waltham et al. (2002). With class accuracy evaluated using the same test pixels, the figures were 56%, 42% and 48% for grass, marsh and woodland, respectively. The output of the ML classifier does not give any indication, however, about the probability level at which a pixel was assigned to a given class. It is quite possible, for example, in areas where it is difficult to discriminate between classes, for even the most likely class to have a probability < 0.5 (50%). Whilst the classification of grass and woodlands appears robust, only 21% of pixel values in Table 3 for known wetland areas had more than a 50% contribution from the marsh end-member. This suggests that many wetlands in the 486 Mean (1 ± SE) proportion of cover type estimated by LSMA Ground cover Wood (%) Grass (%) Marsh (%) Wood Grass Marsh 84.0 (1.8) 17.0 (1.0) 23.8 (0.9) 13.5 (1.5) 75.8 (0.9) 24.0 (0.7) 2.4 (0.5) 6.8 (0.3) 53.9 (0.9) area have a response that includes a significant, minor contribution from grass and woody vegetation. Further investigation of individual class accuracy is provided in Table 4. For all the test pixels where the ground cover was observed to be mainly a single type, we present the mean proportion of that same cover type as estimated by the LSMA. The results suggest that classification accuracy is high for the woodland and grassland categories, with the mean proportion estimated for these test pixels being 84% and 76%, respectively. Marshland areas were estimated to contain, on average, 24% contributions from both woody and grass components. This prediction is consistent with observations made by the field botanists working in this area, who found some significant woody elements even in the seasonally inundated areas. For example, extensive ‘orchards’ of Crescentia cujete (calabash), Haematoxylon campechianum (logwood) and Bucida buceras (bullet tree) were found with understoreys of grass and sedges in several of the savanna areas that have been observed to be inundated for several months during the wet season (Bridgewater et al., 2002). A significant ‘woody’ response was also interpreted from the MLC of the LANDSAT data in the areas identified as subject to inundation from the aerial photography (Fig. 3a). Savanna orchards that are subject to prolonged seasonal inundation do not necessarily have fewer woody species than more welldrained savanna woodlands and the two subtypes may not therefore be simply differentiated when mapping from remote sensing by assuming a consistently lower biomass (lower NDVI values) throughout the orchard subtype. Figure 4 presents the results of the LSMA of the Hillbank savanna as a map. Variations throughout the savanna in the estimated proportions of the wood, grass and marsh vegetation subtypes are shown by the degree of red, green and blue, respectively, in each part of the map. A circular legend (in the style of a colour wheel) is used to assist with interpreting the contributions from each of the three primary colours (the three subtypes) at each location. Areas of land outside the savanna tracts were classified separately using the ML classification reported earlier in the paper. This was to meet a practical requirement of the land managers for the land cover to be mapped over the whole study area. Figure 4 therefore illustrates the two-stage approach to classifying land cover, where a conventional MLC technique is used for mapping the Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd Classifying Neotropical savannas Figure 4 Mapping of results from the three end-member mixture analysis. Variations across the savanna in the estimated proportions of wood, grass and marsh vegetation subtypes are shown by differing contributions to the image of red, green and blue, respectively. Note that the legend is circular. For example, a turquoise pixel represents a high proportion of marsh (blue) and grass (green) and a low proportion of wood (red) estimated to be contributing to the reflectance at that location. Points mark locations along a transect where the vegetation subtype was recorded by field survey. Refer to Fig. 1 for the general location of the Hillbank savanna. Table 5 Savanna vegetation types observed along transect line (marked on Fig. 4) and the proportions of each of the three endmembers estimated in the image at the corresponding location Pixel proportions from LSMA Transect point no. Field description Grass (%) Marsh (%) Wood (%) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Cutting grass marsh by bridge Cutting grass marsh Cutting grass marsh Seasonally dry marshland Seasonally wet Calabash orchard Bullet tree/Calabash orchard Calabash orchard Grassland with scrub bushes Open grassland Open grassland Open grassland Start of pine ridge End of pine fi open grassland Grassland (open, few trees) Scrubland Marshy depression in grassland Pine, palmetto savanna Grassland 13 53 47 48 5 0 20 45 93 100 100 8 82 71 76 78 60 87 55 36 43 52 35 16 23 23 7 0 0 0 1 0 0 22 0 13 32 11 10 0 60 84 57 32 0 0 0 92 17 29 24 0 40 0 widespread high forest biome, and the continuous classification based on LSMA is applied only for mapping within the savanna ecotone. Table 5 summarizes a series of observations of vegetation cover made at points along one of the field transects, which passes through a range of savanna ecotones. The transect is marked on Fig. 4 for reference. The contributions estimated by LSMA for grass, wood and marsh vegetation types were compared with field observations about the dominant vegetation at each location. Points 1–3, for example, are in low-lying wetlands; the continuous classification of the LANDSAT Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd sensor image at these locations estimates high proportions of marsh and grassland. Points 4–7 pass through savanna orchards where the duration of seasonal inundation reduces with the gradually increasing elevation; for these pixels in the image, the contribution estimated from marshland reduces and is replaced by a more woody response. Points 8–11 traverse open grasslands with few trees, leading up to a freely drained sandy ridge of Pinus Caribaea at point 12. In the image, a grassland response at points 10 and 11 is replaced by a strong woodland response at point 12. Points 13–18 pass through a more heterogeneous area of grass savanna with 487 N. Stuart, T. Barratt and C. Place scattered pine, palmetto and bushes. In the image, the response is dominated by the signal for grassland, with an intermittent smaller contribution from woodland. At locations where two or more vegetation subtypes are both estimated to make a significant contribution to the pixel response, the continuous classification allows the relative proportions to be examined. For example, point 17 is described in field notes as an area of ‘pine/palmetto savanna’. Conventional Boolean classifiers such as ML would allocate this to the grassland class, without retaining information about the significant (40%) woody component to the vegetation estimated by the sensor at this location. Comparison of Table 5 and Fig. 4 leads to the conclusion that the proportions estimated by the continuous classification generally agree with field observations of ground cover type. CONCLUDING REMARKS Previous attempts to create resource mapping from medium spatial resolution LANDSAT and SPOT sensor data have typically been at map scales of 1 : 250,000. At this scale, savannas have often been mapped as a single land cover type or their land area has been divided amongst the classes for evergreen forests and scrub grasslands. It was generally not expected that forest–savanna boundaries could be accurately delimited, or any of the subtypes within the savanna matrix resolved, using medium-resolution satellite data (King et al., 1992). Consequently there has been little work specifically analysing savanna vegetation boundaries and subtypes with LANDSAT sensor data, even though it is the most affordable source of remote sensing data for many developing nations. By using GPS to provide greater accuracy and flexibility in collecting ground control and vegetation data for training and testing the classifications, the method reduced mismatch between image data and ground survey observations. With sub-pixel accuracy of image rectification, we have shown that, in the study area, many of the savanna boundaries with wetland, gallery forest and closed canopy evergreen forests could be clearly delineated to within 60 m using LANDSAT TM data. Isolated forest thickets that are similar in composition to evergreen forests could also be discriminated from the savanna areas surrounding them. These findings suggest that with improved ground control from GPS, LANDSAT TM data might be more usable than previously suggested for quantifying changes in forest–savanna boundaries and the growth or decay of forest patches within savanna areas. Whilst maximum-likelihood (ML) techniques are commonly available in software for processing LANDSAT data, we have shown that even using a physiognomic classification developed locally for the land cover in Belize, many of the savanna subtypes exhibited subtle variations that led to poor classification accuracy by the ML method. A similar difficulty was reported by Meerman & Sabido (2001) for classifications of savanna subtypes using LANDSAT data. The finding that several of the savanna cover types have similar spectral 488 properties and cannot consistently be separated, supports the premise that savanna areas produce a mixed response in satellite imagery depending upon factors such as the amounts of woody canopy, grass understorey, degree of seasonal inundation and soil characteristics. This led us to develop a continuous classification of the savanna that estimated contributions to each pixel from three vegetation types (grass, marsh and woodland) and produced a more accurate classification of the test areas than an MLC using similar classes and training data. Interestingly for ecological and botanical investigations, a continuous classification of the LANDSAT image estimated spatial variations in the proportions of woody, herbaceous and wetland vegetation across the study area in a manner that agreed well with field observations made at corresponding locations along ground transect lines. The study has therefore provided some evidence that further information about the nature and condition of savannas may be extracted from LANDSAT data by using more novel types of image classification. The reducing cost of medium resolution satellite data and GPS receivers now enables biogeographers and land managers to test the applicability of this method for classifying savannas using remote sensing and ground survey in many locations world-wide. The accuracy of the ground reference data collected with this method is sufficient to rectify and test classifications of savanna with higher spatial resolution optical imagery, such as 4-m multispectral bands of IKONOS imagery (Jiao et al., 2001) and with airborne or spaceborne radar data, when these become more affordable. With the loss of savanna areas only being recognized relatively recently as a cause of concern (Ratter et al., 1997), and with relatively few remote sensing studies having focused on savannas, there appears to be considerable scope for both a retrospective analysis of changes in savanna areas, and for their future monitoring using these methods. ACKNOWLEDGEMENTS The physiognomic classification of savanna subtypes of the RBCMA was developed by Sam Bridgewater and colleagues at the Royal Botanical Gardens, Edinburgh. We thank the British Ecological Society for funding additional field survey and collecting work by Dr Bridgewater during 2000 to refine the classification for use with remote sensing data. We thank Duncan Moss for his work in collecting the ground control and ground reference data used to rectify and test the classifications of the LANDSAT images. The support of the Royal Society of London and the RICS Education Trust in funding the collection of field data is gratefully acknowledged. We also thank Duncan Moss and Amanda Waltham, whose earlier classifications of the Belizean savannas laid the foundation for this study. The Programme for Belize provided access to the RBCMA and the logistical support of their Ranger Service during the fieldwork was much appreciated. 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BIOSKETCHES Neil Stuart lectures on the Masters programme in Geographical Information Science (GIS) in the Institute of Geography at Edinburgh University. Since leading components of an ecological survey of the savannas of northern Belize in 1996, he has continued to investigate the extent to which savannas can be mapped using remotely sensed imagery that is affordable in developing countries. Tim Barratt specialized in image processing and linear unmixing of LANDSAT TM data for his Masters thesis, working with imagery and field data collected from previous field seasons in Belize. Tim now works for the Tweed Forum as a field ecologist, mapping and controlling the spread of invasive species. Chris Place is a computing officer in the Institute of Geography and has twenty years of experience in specialist processing of earth observation imagery. His work includes remote sensing studies of the savannas of Belize, Brazil and Ethiopia, and land-cover mapping projects in southern Mexico and along the mangrove coastline of Belize. Editors: Peter Furley, John Grace and Patrick Meir This paper is part of the Special Issue, Tropical savannas and seasonally dry forests: vegetation and environment, which owes its origins to a meeting held at the Royal Botanic Garden, Edinburgh, in September 2003. Journal of Biogeography 33, 476–490, ª 2006 Blackwell Publishing Ltd
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