LandscapeEcol(2011)26:877–890 DOI10.1007/s10980-011-9616-0 RESEARCH ARTICLE Landscapes as continuous entities: forest disturbance and recovery in the Albertine Rift landscape Joel Hartter • Sadie J. Ryan • Jane Southworth • Colin A. Chapman Received:3November2010/Accepted:16May2011/Publishedonline:29May2011 (cid:2)SpringerScience+BusinessMediaB.V.2011 Abstract Kibale National Park, within the Alber- surrounding the park. To explore the effect of these tine Rift, is known for its rich biodiversity. High different levels of forest management and protection humanpopulationdensityandagriculturalconversion over time, we assessed forest change over the in the surrounding landscape have created enormous previous three decades, using both discrete and resource pressure on forest fragments outside the continuous data analyses of satellite imagery. Park park. Kibale presents a complex protected forest boundaries have remained fairly intact and forest landscape comprising intact forest inside the park, cover has been maintained or increased inside the logged areas inside the park, a game corridor with park, while there has been a high level of defores- degradedforest,andforestfragmentsinthelandscape tation in the landscape surrounding the park. While absolutechangesinlandcoverareimportantchanges in vegetation productivity, within land cover classes are often more telling of longer term changes and future directions of change. The park has lower Normalized Difference Vegetation Index (NDVI) J.Hartter(&) values than the forest fragments outside the park DepartmentofGeography,UniversityofNewHampshire, andtheformerlyloggedarea—probablyduetoforest 102HuddlestonHall,73MainStreet,Durham,NH03824, regenerationandearlysuccessionstage.Thecorridor USA region has lower productivity, which is surprising e-mail:[email protected] given this is also a newer regrowth region and so S.J.Ryan(&) should be similar to the logged and forest fragments. DepartmentofEnvironmentalandForestBiology,Illick Overall, concern can be raised for the future trajec- Hall,CollegeofEnvironmentalScienceandForestry, tory of this park. Although forest cover has been StateUniversityofNewYork,(SUNY-ESF),Syracuse, NY13210,USA maintained, forest health may be an issue, which for e-mail:[email protected] futuremanagement,climatechange,biodiversity,and increased human pressure may signify troubling J.Southworth signs. DepartmentofGeography,LandUseandEnvironmental ChangeInstitute(LUECI),POBox117315,3141 TurlingtonHall,Gainesville,FL32611,USA Keywords Normalized Difference Vegetation Index (NDVI) (cid:2) Kibale National Park (cid:2) C.A.Chapman Forest fragments (cid:2) Land cover change (cid:2) McGillSchoolofEnvironment,McGillUniversity, Montreal,QCH3A2T7,Canada Continuous analyses (cid:2) Reforestation (cid:2) Regrowth 123 878 LandscapeEcol(2011)26:877–890 Introduction because they represent reservoirs of land, resources, and economic opportunity for people, and simulta- Tropical deforestation is purported to be a primary neously are often viewed as buffers for PAs by cause of global environmental change, with substan- managers (Schonewald-Cox and Bayless 1986). tial projected impacts on biodiversity and world Since very few PAs represent intact ecosystems, it climate (Geist and Lambin 2002). Protected areas has become increasingly important to consider each (PAs) are the main means by which tropical forests protected area as a functional component of a larger are now protected. However, PAs are also often landscape(ParksandHarcourt2002),whichincludes established after forests have been degraded or non-protected areas. cleared,eitherthroughlogging, thinning,agricultural Despite the abundant use and relative success of conversion, or even creation of plantations. These traditional classification methods used in landscape areas ‘‘in recovery’’ or ‘‘returning’’ to an original or analyses, the use of these methods alone may not modified intact state present us with interesting alwaysbeoptimal.Discreteclassificationapproaches challenges in the realm of conservation and preser- areconceptuallysimple,butamajordrawbackisthat vation of biodiversity. While the purpose of a PA much of the variability within each land-cover type may be to restore original habitat, understanding the has been removed (Southworth et al. 2004). Another dynamics of recovery and what the relative value of methodistoincorporatecontinuousdatatorepresent these areas is, requires serious consideration in order a more constant landscape level change (Southworth to make informed management decisions. et al. 2004). Continuous data analyses can provide Particularlyinforests,thenumberanddiversityof more revealing spatial analyses, can focus more on organisms can be increased by disturbance, as biophysical indicators,and can examinemoresubtle, pioneer and invasive plant species may support a within-class variability, and not just across-class panoply of herbivorous insects, smaller mammals, conversion (Foody and Curran 1994; Southworth andbirds.However,thegoalsofconservationandthe et al. 2004). The Normalized Difference Vegetation measures of diversity that we currently use are not Index (NDVI) has been widely shown to be a robust always in concordance. In a study comparing urban and reliable proxy of vegetation productivity and andestablishedruralplantcommunities,Knappetal. quality in many environments (Jensen et al. 1991; (2008) discovered greater phylogenetic diversity in Ehrlich et al. 1994; Rey-Benayas and Pope 1995; established areas, despite greater species richness in Serneels et al. 2001) and an indicator of forage disturbed areas. This pattern may be important when quality(Petorellietal.2011).Netprimaryproduction protecting remaining tropical forests. Endemic or is strongly correlated with the NDVI because it endangered animal species that are tropical old- emphasizes the red and near infrared bands to detect growth habitat specialists will not fare well in changes in biomass (Roughgarden et al. 1991). In disturbed habitats and recovered habitat may take dense, closed-canopy mature forests, we would toolong togrow.Speciesthatcan exploitrecovering expect greenness to be of lower value, due to high habitat, or which are habitat generalists, and less woody biomass and relatively low leaf area, and impacted by disturbance, such as crop-raiding pri- remain stable as a sign of health and persistence. mates or elephants, may slow or prevent sapling Therefore, we can use traditional classifiers to target recruitment and halt reversion to old-growth entirely specific land cover classes (e.g., forest) and NDVI (Lawes and Chapman 2006). Thus, it is important to can be used to examine within-class variability. distinguish old-growth and recovering forest types, In this study, we examine Kibale National Park, and understand their recovery trajectories and how Uganda. It comprises a mosaic of intact primary they contribute to the ecosystem. forest and formerly logged areas, and the ‘‘extended Thepatternsoflandcoverchangeinmosttropical park landscape’’, which contains remnant forest developing countries relate significantly to anthropo- fragments in an agricultural mosaic surrounding the genic forcings occurring across multiple spatial and gazetted park. We examine forest productivity of temporal scales (Woods and Skole 1998; Duncan different components of the PA landscape, using et al. 1999). Therefore, landscapes around PAs, NDVI. This paper builds on previous land cover particularly in tropical forested areas, are important change analyses for Kibale using traditional discrete 123 LandscapeEcol(2011)26:877–890 879 classifiers (e.g., Laporte et al. 2008; Hartter and intensive smallholder agriculture, high levels of land Southworth2009;Southworthetal.2010;Naughton- and resource pressures, and high rates of habitat loss Treves et al., in press), but represents the first use of and conversion, making it a high priority area for continuous data in this fashion and the first detailed conservation (Brooks et al. 2001). Kibale National evaluation of NDVI with respect to areas of known Parkisamedium-altitudetropicalmoistforestwithin disturbance histories. theAlbertineRiftthatcoversapproximately795 km2 Within the Kibale landscape, we examined four in western Uganda (Fig. 1). This transitional forest differentforesttypes:in-park,intactforest,whichisold- (between lowland rainforest and montane forest) has growth; logged-a conglomerate of former logging an average elevation of 900–1590 m and the climate concessionsinwhichcuttingceasedin1975,andwhich is warm throughout the year, with an average daily isrevertingtomorecontinuousforest;forestfragments minimum of 15(cid:3)C and maximum of 23(cid:3)C, and a outside the park within a 5 km buffer (Hartter and mean annual rainfall of 1,662 mm (1970–2010, Southworth 2009; Hartter et al. 2009), which are in T. Struhsaker and C. Chapman, unpublished data). varying states of degradation and use; and, a former The bi-modal rainfall pattern produces two major gamecorridorthatextendsfromthewesternboundary rainyseasons:‘‘short’’(lateFebruary-earlyMay)and of the park. Using land cover classifications of forest, ‘‘long’’ (late August-late November). agriculture, wetlands, tea, and crops/bare ground cre- PriortoKibalebecomingaNationalParkin1993,it ated in previous research (Hartter and Southworth, was designated a Forest Reserve with the main 2009; Southworth et al. 2010), we can compare these objective to provide a sustained production of hard- fourforesttypes/landscapelocations(park,loggedpark, wood timber (Struhsaker 1997) and softwoods from forest fragments and corridor) over time, in terms of plantations. Logging was conducted at regular inter- landscapechangesandchangesinforestproductivityto valsbetween1950and1975atvariableintensities,but addressthefollowinghypotheses: mostly between 1967 and 1969, resulting in large forest gaps and variable forest disturbance patterns 1. Intact forest cover will remain stable over time, (Hartter et al. 2009). Harvested areas were typically while fragmented, logged, and corridor areas are allowed toregeneratenaturally, although undesirable expected to change. trees were poisoned in some areas. During Uganda’s 2. Differenttypesofmanagement(i.e.,foresttypes) political upheaval in the 1970s and 1980s, the will lead to different NDVI signatures. plantationstypicallyestablishedonanthropogenically 3. Establishment of a corridor area will lead to createdgrasslandswerenotmanaged(Chapmanetal. reforestationandchangingNDVIvalues,resulting 2002). Management plans changed when Kibale in higher NDVI values initially and becoming became a national park in 1993. Plantations were moresimilartothoseofintactforestovertime. harvested from 1993 to 2006 (using manual pit saws 4. Asthepreviouslyloggedareasoftheparkrevertto and portable sawmills) and the harvested pine areas moreintact, morematureforest,theproductivity werelefttoregeneratetonativeforest(Chapmanetal. signatureofloggedforestwillbecomesimilarto 2002; C. Chapman unpublished data). In 1964 the theforestintherestoftheparklandscape. Kibale Forest Corridor Game Reserve was gazetted 5. Forest fragments outside the park will have and administered as a separate entity by the Uganda earlier successional stages of plant cover, corre- GameDepartmentundertheMinistryofTourismuntil sponding to similar NDVI values as other it was combined with the Kibale Forest Reserve in regenerating forest. 1993tocomprisethepresent-dayKibaleNationalPark (Struhsaker1997).Asearlyas1971,illegaldestruction andencroachmentoccurredinthecorridor,whichled Study area toforestconversiontofarms.Theestimatesofpeople who settled in the corridor varies considerably TheAlbertineRiftregioninEastAfricaisoneofthe (8,800–170,000),butallwereevictedin1993(Chap- world’s hotspots for biodiversity (Plumptre 2002; manetal.2010a). Plumptreetal.2003;2007;Cordeiroetal.2007).Itis Outside Kibale, a mosaic of small farms (most also one of the most threatened, due to dense \5 ha in size), large tea estates ([200 ha), and 123 880 LandscapeEcol(2011)26:877–890 Fig.1 KibaleNational Parkandsurrounding landscapeandareas classifiedasforest interspersed forest fragments and wetlands, effec- and resource availability depending on human use tivelyisolatetheparkfromothertractsofforest.Tea and disturbance (Hartter 2010). isconfinedtothenorthwestandnortheastsidesofthe Nearly 95% of the population in this area (pre- park. Most tea lands are held by large multi-national dominately Batoro and Bakiga tribes) sustains their companies consisting of hundreds of hectares, but livelihoods through farming or other agricultural- other smaller, individual holdings also dot the based activities (National Environment Management landscape. A vast network of bottomland forest Authority2001).Thisareaisoneofthemostdensely fragments and wetlands either extend out from the populated areas in Sub-Saharan Africa (Lepp and park or are isolated within this mosaic landscape. Holland 2006), with the population around Kibale Theseareasvaryinsize(\0.5 hato[200 ha),shape, having grown bymorethan 300% between1959and 123 LandscapeEcol(2011)26:877–890 881 1990 (Naughton-Treves 1998) and estimated at an layer.Thefinalfive-classclassification(forest;crops/ averageofapproximately300individuals/km2within bare land—including pastures, cultivated crops, and 5 km of the park boundary by 2006 (Hartter 2010). kitchen gardens; papyrus (Cyperus papyrus L.) and However,theeasternsidecarrieshigherdensitiesthan elephant grass (Pennisetum purpureum); tea; and the west. water) obtained an overall accuracy of 89.1% and an overall kappa statistic of 0.867 (Hartter and South- worth 2009). The classifications were done indepen- Methods dently for each image and not based on the same spectralsignatures due to a lack ofexact anniversary Image pre-processing dates, although the pattern of land cover signatures was consistent across all image dates. SevenLandsatTMandETM?sceneswereobtained: To address our hypotheses, the 1984, 1995, and 26 May 1984, 4 August 1986, 20 August 1989, 17 2003 land-cover composites were used to create January1995,9January2001,31January2003,and11 change trajectories. Given that we wanted to under- September 2008. All were acquired at the end of the standthedominantlandcovertrajectories,anddueto dryseasonwhenfallowagriculturallandscanbeeasily how sensitive this method is to the number of initial distinguishedfromforests,exceptfor1984,whichwas classes and image dates, we used only three of the acquiredattheendoftherainyseasonsincethiswas availableimagedateswhichwerespreadouttodetect theonlyavailablehaze-andcloud-freeimage.Images the patterns of change. (No data were available to weregeometricallyregisteredto1:50,000scalesurvey include the 2008 image as part of the land cover topographic maps of the region, followed by radio- analysis and we limited the number of possible metric calibration and atmospheric correction. All trajectories). Land-cover/use classes of interest here images were geo-registered to within a root mean are areas forested on all three image dates (forest), square error of\0.5 pixels (below 15 m accuracy). areas ofwetland andgrassesonallthree imagedates SinceLandsatTMandETM?satellitesdonotmake (wetland),andareasofagricultureonallthreeimage geographicallyexactreturnvisits,weoverlaidthegeo- dates (crops or tea) which represents the three stable registeredimages andclipped outareasfor which, in land cover classes in the region. Next we have two any year, there were missing data. This mainly conversion classes of interest: reforestation (when occurredinverysmallareasofthefarnorthandeast, anynon-forestclass(cropsorwetland vegetation)on giving a slightly truncated edge to the buffer date one or date two is forested by date three; and (describedbelow)inthoseregions(seeFig. 2). deforestation (any area of forest on date one or date two, which is non-forest (wetland vegetation, tea or Image classification and land cover change crops) on date three). Water was excluded from this detection analysis since the crater lakes predominant in this region do not vary in area over time. Areas of forest We created a buffer zone outside the park boundary cover vary in terms of geography and past land use to a distance of 5 km to describe the surrounding history, and here we discuss four different forest landscape(769 km2).Thisdistancewashypothesized ‘‘types’’.(1)Intactforest—thisisnaturalforestcover, asbeingfarenoughtocapturesocio-economiceffects found within the park boundaries, not logged or ofthepark,asappliedinotherstudies(DeFriesetal. otherwiseimpactedwithinrecenthistory.(2)Logged 2005; Goldman et al. 2008). Land-cover maps were forest—forest areainthe northernmostsectionofthe derived for each date by independent supervised park which was previously logged. (3) Fragments— classification for three Landsat scenes: 1984, 1995, degraded remnants of the intact forest found in the and2003.Duringfieldseasonsin2004and2005,180 landscape around the park. Most of these forest trainingsampleswerecollectedandusedtoconstruct fragments have seen extensive human use. (4) a supervised classification using the Gaussian max- Corridor—formergamecorridorconnectingtoneigh- imum likelihood classifier. The classified image was boringQueenElizabethPark.Ittransectsthewestern constructed using a layer stack including all bands domesticated landscape, and represents recent regen- 1–5, 7, texture bands of layers 1–5, 7, plus an NDVI erating forest. 123 882 LandscapeEcol(2011)26:877–890 Fig.2 Trajectoriesofland use/coverchangecategory occurringwithinthepark boundariesandsurrounding landscapefrom1984to 1995to2003.Reforestation Areasthattransitionedto forestfromagricultural land,Forestconversion areasthattransitionedfrom foresttoagriculture, Agriculturecrops,baresoil, shortgrasses,tea;elephant grassinallyears,Wetland wetlandvegetationinall years,StableForestforest inallyears NDVI analysis analysistodetermineandaccountfordifferencesdue to past management. If we do not separate this out, We calculated the NDVI value for each image, and the signal from the NDVI change analysis may be a adjusted NDVI values for the 2001 and 2003 ETM? mixed signal and prove to be confusing. By incor- images to TM values (Steven et al. 2003; Table 2). poratingthepastuseandmanagement(intact,logged, To examine the trajectory of productivity over time, regenerating, and fragment) we can then evaluate wecomparedthemeanNDVIwithineachforesttype their respective impacts. (Table 1; Fig. 3) for all image dates. Given the Each of the images was captured following differences in forest type, we then used the NDVI abundant rainfall during the rainy seasons and all 123 LandscapeEcol(2011)26:877–890 883 Table1 Land Cover/Use trajectories for the study region for interest:stableforestcover,stableagriculture,stablewetland/ 2003,shownforintactforest(P)andtheloggedarea(L)within grass,reforestationordeforestation,intermsofthepercentof Kibale,forestfragmentsoutsidepark(within5kmaroundthe thelandscape park)(F),andthegamecorridor(C),forthefivetrajectoriesof Trajectory Intactforest(P) Loggedarea(L) 5kmBuffer(F) Corridor(C) (%area) (%area) (%area) (%area) Stableforest 79.2 82.5 15.7 20.5 Stableagriculture 4.5 2.6 42.3 42.3 StableWetland 1.2 1.1 5.5 3.4 Reforestationa 11.3 9.0 17.1 16.6 Recentreforestation 6.4 4.4 10.1 9.0 Olderreforestation 5.0 4.6 7.0 7.7 Deforestationb 3.9 4.8 24.7 16.4 Recentdeforestation 1.7 3.6 15.6 10.8 Olderdeforestation 2.2 1.2 9.1 5.6 a OlderreforestationAreasthatbecameforestfrom1984to1995andremainedforest,Recentreforestationareasthatbecameforest from1995to2003 b OlderdeforestationAreas thatlostforestfrom 1984to1995,Recentdeforestation areasthatlostforestfrom1995to2003and remained seven images are not anomalous (not in peak or Results trough precipitation months). Mean rainfall totals for the4 monthspriortoimageacquisitionforallimages Spatial patterns of land cover change were not significantly different than the 4 months prior to the 2008 image capture (Dunnett’s test, Forest remains the dominant land cover type within P[0.05). Kibale (not including the corridor—Fig. 2). Over time, park boundaries have been maintained with no Change detection evidence of large-scale encroachment. The park showed a consistent pattern of reforestation since We used a two-factor (type and year) ANOVA 1984, while only 3.9% of intact forest and 4.8% of framework to explore differences between mean the logged area deforested (Tables 1, 2). The logged NDVI values in the four forest types. A post-hoc area had 9% reforestation identified (Table 1). Tukey’s Honest Significant Difference test (HSD), In contrast, outside the park the situation is quite a = 0.05 was used to determine which forest types different. This domesticated landscape is in a con- differed from one another. To assess the increase in stantstateoffluxasaresultofthevariouslivelihoods similarity of disturbed forest types to the intact type, being supported. Conversion of forest occurred at a we compared differences in types over time. Differ- high rate, while there is still some effort towards encesbetweenmeanNDVIvaluesoftheloggedarea, reforestation.Althoughthere isanoveralllossof2% corridor, fragmented forests, and intact forest were total forest, only about 16% of the forest fragments calculated for each image year. We regressed these outsideKibalehaveremainedstablesince1984.This differences upon year (nominal) to determine if the area has experienced higher reforestation rates than differences were decreasing over the study period. the park, but has also had nearly 25% deforestation, Sincetheimageswerecollectedatdifferentintervals, suggesting a high rate of landscape turnover. Simi- wecouldnotperformformaltimeseriesanalyses.As larly,only20%oftheforestinthecorridorremained thesedataarequiteheterogeneouswithinforesttype, since 1984, while the corridor also sustained 16.4% wealsocalculatedthemedianvalueforeachtypeand deforestation, with almost 11% of it recent (since a image, and found that medians differed from the year before being gazetted). At the same time, the mean by an average of only 1.35%, suggesting that corridor has had 16.6% reforestation, which also the mean was not biased. indicates a high rate of landscape turnover. 123 884 LandscapeEcol(2011)26:877–890 Fig.3 GraduatedNDVIcompositesforforestonlyfortheparkandthe5kmbuffer NDVI change The two-factor ANOVA on forest type and year explained 95% of the variance in the data, with the Itisevidentthatthereisaspatialpatternofchangein majority explained by year (R2 = 0.95, SS = 0.22, Y NDVIfromimagetoimage(Fig. 3).Withinthepark, SS = 0.01), but both factors were significant (forest T the highest NDVI values are found in the northern type: P = 0.003, Year: P = 0.0001). The Tukey’s sectionsofthepark,whichwereloggedbetween1950 HSD showed that the logged area had the highest and 1975, and are now regenerating forest (Table 2). NDVIvalueswhencontrollingforyear(althoughnot The rest of the park which was not commercially significantly higher than the fragments), followed by logged shows NDVI values below the mean. The the corridor, and the intact forest had the lowest logged area within the park has higher mean NDVI values (although not significantly lower than the valuescomparedtotheentireparkandthesurrounding corridor; Table 3). The fragments have a higher than landscapeforallimageyears(Fig. 3).Thereisalsoan average NDVI value in all years except 1989, and in increasinggradientofNDVIintheparkfromthelower 2001 and 2003, the highest mean NDVI values elevationsintheeastandsouthtothehighernorthern overall (Fig. 4). In addition, most individual frag- forests, which may also be related to the logged- ments had above-mean NDVI values for each image unloggedareasand/orrainfallpatterns(Fig. 3). year (Fig. 2). 123 LandscapeEcol(2011)26:877–890 885 Table2 Forestcover andNDVIvaluesforintactforest(P)andtheloggedarea(L)withinKibale, forestfragmentsoutsidepark (within5kmaroundthepark)(F),andthegamecorridor(C) Intactforest(P) Loggedarea(L) Year Forest NDVI Year Forest NDVI (%area) Mean StdDev (%area) Mean StdDev 1984 86.1 0.570 0.071 1984 88.5 0.669 0.050 1986 85.8 0.581 0.053 1986 89.6 0.638 0.050 1989 85.8 0.604 0.051 1989 89.6 0.650 0.046 1995 85.8 0.533 0.046 1995 89.6 0.600 0.049 2001a 90.3 0.336 0.067 2001a 91.0 0.386 0.066 2003a 90.3 0.448 0.049 2003a 91.0 0.488 0.050 2008 90.3 0.609 0.049 2008 91.0 0.644 0.046 5kmBuffer(F) Corridor(C) Year Forest NDVI Year Forest NDVI (%area) Mean StdDev (%area) Mean StdDev 1984 31.9 0.645 0.063 1984 36.0 0.630 0.068 1986 34.3 0.603 0.071 1986 39.2 0.584 0.070 1989 34.3 0.600 0.085 1989 39.2 0.575 0.091 1995 34.3 0.575 0.045 1995 39.2 0.548 0.045 2001a 29.2 0.413 0.069 2001a 37.2 0.407 0.078 2003a 29.2 0.503 0.051 2003a 37.2 0.376 0.050 2008 29.2 0.629 0.078 2008 37.2 0.637 0.068 a StandardizedNDVIcalculationsfromETM?toTM(Stevenetal.2003),RemoteSensingofEnvironment88,pp412–422 The corridor NDVI values varied about the 3 negative changes and alternating year patterns landscape average within a similar range to the respectively (Fig. 5). fragments and logged area, with the striking excep- tionof2003.Inthecorridor,therewasalowerNDVI value than any other forest category in 2003, and the Discussion largest difference from the intact forest of any category in any year (Fig. 5). Visual inspection of composite satellite images and Likely due to low sample size (n = 7), the the comparison of NDVI values show that Kibale regressions of differences between disturbed forest National Park has become an island of forest in a types and within-park intact forest across years were landscape surrounded by intensive agriculture. This not significant. However, the formerly logged area, domesticated landscape is characterized by highly was marginally significant (P = 0.06) and negative fragmentedandrapidlychanging landcovers,imply- and captured 52% of the variability (logged = inghighlydynamiclanduses.Ourresultsindicatean 3.44 - 0.002*year, R2 = 0.52), while the fragments increase in forest cover inside the park since 1984. (fragment, P = 0.88, R2 = 0.005) and the corridor Specifically, the majority of Kibale’s in-park forest (corridor, P = 0.81, R2 = 0.013) showed no trends. has remained intact with only a small proportion lost In addition, results from the logged area (in which 5 to deforestation over 20 years. of 6 between-image transitions in difference (83%) Althoughitdoesnotcompletelycapturetheeffec- were negative, including the last 3 transitions) tiveness of park management, remotely sensed data suggest convergence to the intact forest signature. does reveal the impacts of various management In contrast, the fragments and the corridor had 4 and approaches on the vegetation cover and dynamics. 123 886 LandscapeEcol(2011)26:877–890 Table3 Least-squares Mean (LSM) NDVI differences in forest types, after controlling for year, using a post-hoc Tu- key’sHSDona2-factorANOVA Type LSM Group L 0.582 1 F 0.567 1,2 C 0.537 2,3 P 0.526 3 Summarized by forest type (Type: Logged (L), Fragments in the surrounding domestic landscape (F), Corridor (C), and intact forest within the park (P)), least-squares mean NDVI value (LSM), and grouping (group), where types in the same group (1–3) are not significantly different from one another. Fig.5 The differences of mean NDVI value between forest Overallmodelstandarderror=0.01 typesL(black),F(grey)andC(white)andintactforest(P)at successiveimagesdatesinthestudy landscape had significantly higher apparent produc- tivity (as measured by NDVI) than did the intact forest within the park. Interestingly, the corridor appearedtohaveanoverallsimilarityinproductivity, statistically, to both the forest fragments outside the park and to the intact portion of the forest within the park. It is important to note that these images were obtained in the same season within each year, and that the precipitation patterns preceding each image werenotstatisticallydissimilar.Sincetheseasonality of the area has been changing (Hartter unpublished Fig.4 Mean forest NDVI values for intact forest within Kibale,theloggedareawithinthepark,fragmentswithinthe5 data), using the same Julian date in each image year kmbufferoutsidethepark,andtheformergamecorridor would have been inappropriate. It is thus interesting to see the variability in the NDVI values between Forexample,oneofStruhsaker’s(2002)indicatorsof image years, and due to the corrections between park success isadecreaseinillegalactivities. Large- sensors(TMandETM?),thevaluesarecomparable. scale encroachment into Kibale has virtually halted It is important to understand the within-class since formal park establishment in 1993 (Hartter and variability and its potential ecological significance Goldman 2011) and there are well understood and withinourregions.Itislogicalthattheintactforestin maintained boundaries. When compared to the sur- the park has lower NDVI values (i.e., NDVI in rounding landscape, the area within the park has matureforest levels off and then decreases over time experienced limited land cover change in absolute comparedtonewer,secondarysuccessionalgrowthin terms,buttherearenotablechangesormodificationsin regenerating areas) than the logged area and buffer, cover as seen from the NDVI analysis. While effec- butperhapspuzzlingisthatithaslowermeanvalues tively protected from human exploitation, the park thanthecorridorin5outof7 years.Whilethebuffer itselfisnotunchanging,withvariationfromoneland- area outside the park is an active transition zone cover class to another. For example, areas misclassi- where there is ongoing deforestation, there is also fied as crops in the park are naturally occurring transition of former agricultural lands back to forest. grasslands, which undergo succession in savanna Sincemanyoftheseforestsareatearliersuccessional areas,andsomevariationfromwetlandstobareareas. stages, the productivity was expected to be higher Wefoundthattheformerlyloggedareawithinthe than forests at later successional stages, resulting in parkandtheremnantforestfragmentsinthedomestic above-mean NDVI values for the forest fragments. 123
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