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MODELING ECOLOGICAL CONNECTIVITY IN A PROTECTED AREA NETWORK IN SOUTHEAST TANZANIA A THESIS PRESENTED TO THE DEPARTMENT OF HUMANITIES AND SOCIAL SCIENCES IN CANDIDACY FOR THE DEGREE OF MASTER OF SCIENCE By ADAM P. DIXON NORTHWEST MISSOURI STATE UNIVERSITY MARYVILLE, MISSOURI JUNE, 2012

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Page 1: MODELING ECOLOGICAL CONNECTIVITY IN A PROTECTED …World Heritage Site and 107 protected areas of various level of management. Ecological connectivity between protected areas is modeled

MODELING ECOLOGICAL CONNECTIVITY IN A PROTECTED AREA NETWORK IN SOUTHEAST TANZANIA

A THESIS PRESENTED TO THE DEPARTMENT OF HUMANITIES AND SOCIAL SCIENCES

IN CANDIDACY FOR THE DEGREE OF MASTER OF SCIENCE

By ADAM P. DIXON

NORTHWEST MISSOURI STATE UNIVERSITY MARYVILLE, MISSOURI

JUNE, 2012

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MODELING ECOLOGICAL CONNECTIVITY IN SOUTHEAST TANZANIA USING A FLAGSHIP SPECIES

Modeling Elephant Movement

Adam P. Dixon

Northwest Missouri State University

THESIS APPROVED

Thesis Advisor, Dr. Ming-Chih Hung Date Dr. Patricia Drews Date Dr. Yi-Hwa Wu Date Dean of Graduate School, Dr. Gregory Haddock Date

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MODELING ECOLOGICAL CONNECTIVITY IN A PROTECTED AREA

NETWORK IN SOUTHEAST TANZANIA

Abstract

The increased fragmentation of habitat capable of sustaining the life cycle of paradigm

species, such as the African elephant, is occurring due to agricultural and urban expansion. In

addition to its intrinsic value as one of the largest animals on the planet, the elephant can serve

as an umbrella species, whose habitat also sustains a large number of other plant and animal

species. Predicting the migration patterns of elephants may offer clues to how to best develop

an effective use of resources in the development of protected and managed areas for the

conservation of nature. Using habitat preference values developed from a habitat selection

framework which uses a combination of elephant GPS point locations with a land cover dataset,

a least cost corridor can be developed. Conditional Minimum Transit Cost is used to develop a

more realistic passage space between protected areas in southeast Tanzania. The network of

protected areas contain some of the last vestiges of east African coastal forest, which contain

world class ecosystems of high endemism and species richness. Using the elephant as a flagship

species might improve the chances of conservation in this region which contains a UNESCO

World Heritage Site and 107 protected areas of various level of management.

Ecological connectivity between protected areas is modeled amongst a network of

protected areas. A new strategy - Conditional Minimum Transit Cost - is successfully employed

for modeling least cost movement across a vast protected area network across the southeast

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Tanzania landscape. Several bottlenecks, areas of concern and areas where increased

protection could increase the effectiveness of corridor space are identified. Using the protected

area as a surrogate for habitat patches may be problematic, since it does not accurately reflect

the landscape as an elephant experiences it. This is most reflected in the results as most

corridors with high resistance to movement are those with a greater distance between origin

and destination. An analysis that identifies habitat patches of adequate size and using those

results as origin and destination nodes is suggested in subsequent iterations of similar analyses.

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Table of Contents

List of Figures ............................................................................................. vii

List of Tables................................................................................................ ix

Acknowledgements ...................................................................................... x

List of Abbreviations ................................................................................... xi

Chapter 1: Introduction ............................................................................... 1

1.1 Research objective ...............................................................................................6

Chapter 2: Literature review ....................................................................... 7

2.1 Least cost modeling .............................................................................................9

2.2 Modeling habitat................................................................................................16

Chapter 3: GIS modeling of connectivity ................................................... 22

3.1 Study area ..........................................................................................................22

3.2 Methodology overview ......................................................................................25

3.3 Water .................................................................................................................28

3.4 Land cover .........................................................................................................31

3.5 Slope ..................................................................................................................34

3.6 Roads .................................................................................................................35

3.7 Human density ...................................................................................................36

3.8 Cost surface .......................................................................................................38

3.9 Corridor modeling with CMTC ............................................................................41

3.9.1 Modeling corridor width and alternate dispersal corridors ..............................45

3.10 Calculating CMTC .............................................................................................47

3.10.1 Finding five percent.......................................................................................48

3.10.2 Extracting five percent ..................................................................................49

Chapter 4: Results and discussion ............................................................. 51

4.1 Cost surface results and discussion ....................................................................52

4.2 Zonal characteristics discussion ..........................................................................60

4.3 CMTC results and discussion ..............................................................................62

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4.3.1 Regional zone results and discussion ...............................................................63

4.3.2 Results from a single corridor ..........................................................................71

4.3.3 Entire study area results and discussion ..........................................................74

Chapter 5: Conclusion ............................................................................... 78

5.1 Least cost modeling ...........................................................................................79

5.2 The use of data ..................................................................................................80

5.3 Model processing issues.....................................................................................83

5.4 The exclusion of scale sensitivity ........................................................................85

5.5 Final thoughts ....................................................................................................85

Appendix A: Africover land cover classes reclassified .............................. 87

Appendix B: Modeled PA connections ..................................................... 91

Appendix C: List of corridors between PAs modeled ................................ 95

References ............................................................................................... 101

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List of Figures Figure 1. Theoretical least cost movement across a cost surface ...................................10

Figure 2. Study area .......................................................................................................23

Figure 3. Flowchart of cost surface development ...........................................................27

Figure 4. Data used for water source .............................................................................29

Figure 5. Flowchart for modeling corridors ....................................................................42

Figure 6. The extent of each zone of analysis within study area. ....................................44

Figure 7. Results of the distance to water analysis .........................................................53

Figure 8. Land cover reclassification ..............................................................................54

Figure 9. Result of the slope analysis .............................................................................55

Figure 10. Result of the distance to roads analysis .........................................................56

Figure 11. Human population density reclassification ....................................................58

Figure 12. The final cost of movement surface...............................................................59

Figure 13. Total area in each zone .................................................................................61

Figure 14. Percent of land cover type for each zone of analysis ....................................62

Figure 15. Model result for Zone A ................................................................................65

Figure 16. Model Result for Zone B ...............................................................................66

Figure 17. Model result for Zone C .................................................................................67

Figure 18. Model result for Interzone A .........................................................................69

Figure 19. Results from Interzone B ...............................................................................70

Figure 20. Model result between Uluguru South and Kimboza in Zone A ......................72

Figure 21. Model result between Ntama and Lionja in Zone C ......................................73

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Figure 22. Model results for entire PA network .............................................................76

Figure 23. Model result classified for PA network .........................................................77

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List of Tables

Table 1. Values given to distance to water criterion .......................................................30

Table 2. Integrated Galanti Land Cover Classes into Preferential Scale...........................33

Table 3. Values used for slope criterion in model ...........................................................35

Table 4. Results of distance to roads analysis .................................................................36

Table 5. Values applied to weight human population density criterion in model............38

Table 6. Final amalgamation of movement cost surface values ......................................41

Table 7. Number of CMTC analyses completed ..............................................................47

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Acknowledgements

The author would like to thank Dr. Neil Burgess of WWF-US for providing the research

concept as well as my thesis advisor Dr. Ming Hung for providing feedback during the

nearly two year process which was required for this thesis. The other two members of

the thesis committee, Dr. Patty Drews and Dr. Yi Hwa Wu, improved the quality of the

thesis considerably through the review process. The author’s parents, grandparents,

siblings and friends were of immense value for their constant support and love.

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List of Abbreviations

AHP Analytical Hierarchy Process

CMTC Conditional Minimum Transit Cost

EROS Earth Resources Observation Satellite

ESA European Space Agency

ESRI Environmental Systems Research Institute, Inc.

FAO Food and Agriculture Organization

GIS Geographic Information Systems

IUCN International Union for the Conservation of Nature

NDVI Normalized Difference Vegetation Index

ORNL Oak Ridge National Laboratory

PAs Protected Areas

SRTM Shuttle Radar Topography Mission

TM Thematic Mapper

UNESCO United Nations Educational, Scientific and Cultural Organization

WDPA World Database of Protected Areas

WWF World Wildlife Fund for Nature

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Chapter 1: Introduction

Geographic Information Systems (GIS) have been employed in recent decades in

the natural sciences arena to model natural phenomena. The discipline of ecology has

been particularly enhanced due to its primary focus on the interactions of the biotic and

abiotic while occurring in some measure of space. The utility of GIS in this type of

research is vast. Ecosystems and individuals in those ecosystems can be modeled from

the global scale to the local. The two forms of visual geographic data, vector and raster,

are well suited for ecological analysis and visualization. Points, lines and polygons as

vector data bound to coordinate systems are powerful allies in the construction and

deconstruction of natural space. Raster surfaces as cell values help to effectively define

ecological characteristics in a simple or complex matrix framework. It is within this

nexus of ecology and GIS that this paper will demonstrate the utility of modeling

ecological connectivity across geographic space using a megafauna species in east Africa

as a surrogate for the connectivity.

Land use conversion is an increasing reality over much of the world’s terrestrial

surfaces. An expected 8.9 billion people are expected to occupy the earth by the year

2050 (United Nations 2005). Agricultural intensification and expansion will need to

increase substantially in order to provide nourishment for many of these people (Gibbs

et al. 2010). Urban areas are expected to triple in size by the year 2030 (Angel et al.

2005). African road networks are projected to increase substantially to support new

economies across the continent (Buys et al. 2006). In northern Tanzania, a road is

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planned to cross the ancient migration routes of the large mammals of the Serengeti

grasslands (Dobson et al. 2010).

The effective result of land use conversion is habitat fragmentation, which is

correlated with declining species abundance (Donovan and Flather 2002), and is an

increasing trend on a global scale (Fahrig 2003). Habitat fragmentation occurs when an

ecosystem goes through a disturbance, which can be natural or anthropogenic in origin.

Natural disasters may include fire, meteorological storms or other natural processes.

Anthropogenic sources are the origin of land use conversion. Agriculture, urban areas

or infrastructure all may be cited as reasons for land use conversion. Species dependent

on the ecological characteristics of fragmented areas are forced to move to a different

location and often experience early mortality. The reduction of habitat area also

contributes to the insularization of species populations (Newmark 1996), since the total

area of potential habitat space reduces in size resulting in an ecosystem unable to

sustain the same population as before.

The first proposed method to deal with land use conversion is to find the most

ecologically significant regions and propose areas for reservation from further human

activity. These reserves establish protected areas (PAs), which have various levels of

management and administration. Protected areas are managed at the international,

national and local levels. The enforcement of various restrictions on the use of lands

within PAs for human activities can be poorly managed in some countries. While an

imperfect solution to the problem of habitat loss, the creation of PAs have provided a

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high degree of successful species conservation which otherwise would have been lost

(Naugton-Treves et al. 2005).

As stated, the creation of PAs has not decreased the overall acceleration of

worldwide species extinction at rates which are estimated to be occurring 1000 times

faster than normal background evolutionary processes (Primm et al. 1995).

Explanations for this rapid loss of unique life forms include anthropogenic-caused land

use conversion, as well as associated vulnerabilities correlated to habitat degradation.

Therefore, the strategies for conservation of biodiversity employed by non-

governmental organizations (NGOs) and government institutions have not sufficiently

decreased the rate of extinctions. The traditional paradigm of discrete populations

within discrete geographic regions has proven inadequate to deal with species loss.

Isolation of wildlife populations within protected areas has been shown to reduce

genetic diversity and increase genetic drift (Frankam 2005), as well as increase the risk

of competitors, predators, parasites, diseases, and natural catastrophes (Shaffer 1981).

Thus, perceiving healthy wildlife populations as continuously dispersed across a

functional geographic space may more adequately address the concern of the species

loss.

The concept of modeling habitat connectivity has been proposed as a landscape

planning tool to avoid the negative effects of habitat insularization. Habitat connectivity

refers to the realization of natural pathways a species will take while moving across

geographic space. Habitat selection and the path of least resistance might help predict

the likelihood for an individual to take a certain route. The concept of random walking

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serves as a baseline for the concept of connectivity, which posits that an individual will

take the path of least resistance. The path is often characterized by slope, elevation,

foraging and mating potential amongst other parameters which may be used to predict

the most likely route traveled by an individual. The rationale of connectivity is that if

individuals can have some protected route to continue their life cycles and spread

genetic diversity amongst macropopulations, then perhaps the conservation objective

of stabilizing species loss can be achieved. By extension, a perfect place to find a route

is between protected areas which suffer from insularization and might have increased

capacity to conserve species if some degree of connectivity with the exterior landscape

is realized.

Large mammals work well as a surrogates for the overall success of conservation

efforts. It has frequently been demonstrated that the first observable species loss from

an area due to habitat disturbance are large mammals (Newmark 1993). This has been

observed for the American Bison on the North American prairie (Newmark 1993), as

well as the black panther, whose historic range covered Florida through to south central

South America (Rabinowitz and Zeller 2010). The African elephant Loxodonta africana

is a large mammal species whose ability to range across the landscape is limited by the

abundance of foraging material and other traditional habitat characteristics, as well as

the more recent human-elephant conflict events that have resulted in large numbers of

elephant mortality in recent times (Blanc et al. 2007). The population of African

elephants plummeted fifty years ago down to only 100,000 individuals from 2 million,

and then in recent decades has finally risen to somewhere around 500,000 (Blanc et al.

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2007). Perhaps the lower numbers of elephants from historical times indicates the

pressures facing many ecosystems across eastern Africa, and indeed across the globe.

The concept of flagship species is helpful when developing conservation

scenarios for an area of conservation concern. The term flagship species in this paper is

intended to have similar connotations to the terms surrogate species, umbrella species,

indicator species or focal species. The elephant is certainly a prime example of a species

with enough malleable marketing prowess and ecological significance to fit nicely into

any definition produced for any of the terms. It is a species that needs large tracts of

habitat that by conserving will automatically operate as effective conservation for a

number of other species (Simberloff 1998). To fully assess the effectiveness of a flagship

species, an ecological characterization of similar habitat can evaluate the true value of a

flagship species. Epps et al. (2011) found that elephant presence was positively

correlated with a number of other large mammals. Therefore, predicting the pathway

an elephant may take from one protected area to another might help to ensure the

continued existence of the species, as well as serve as a flagship species for a network of

interconnections between the biotic, abiotic and physiographic that serves to stabilize

ecosystems.

Developing predictive parameters for modeling ecological connectivity within a

GIS is a complex exercise. There are many decision points in designing a methodology

to execute a model. There is, however, a vast collection of literature that has been

developed over the last several decades that can serve as a guide to methodology

development.

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1.1 Research objective

The protected areas of southeast Tanzania currently lack a conservation plan

with an incorporated connectivity analysis. This has implications for coastal forest and

elephant populations in the region. The research objective is to offer a methodology to

model elephant corridors for ecological pattern and process within protected areas in

southeast Tanzania between Selous Game Reserve and the eastern coast.

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Chapter 2: Literature review

The definition of connectivity has been described as the degree to which the

landscape facilitates or impedes movement among resource patches (Taylor et al.

1993). Reviewing the topic of connectivity modeling yields a great deal of research

dealing with the GIS and mathematical basis for connectivity modeling and perhaps in

higher quantity the ecological application that serves as the major driver for inquiry into

the concept. Connectivity from a GIS perspective is perhaps as important as any other

subsection of research. Ensuring that nodes of connectivity are designed properly plays

a major role in the overall effectiveness of the effort. The concept of connectivity from

the ecological perspective can be developed once the GIS theory is developed.

Establishing of a focal species for the landscape being modeled is a critical component

to the effort since the behavioral characteristics of the species are what the model is

calculating. Evaluating the role of geographic information within the project is also of

great concern. The quality of the data governs to a large degree the outputs.

Addressing issues in corridor effectiveness ensures outputs are actually correlated with

the natural occurrence of phenomena.

Connectivity modeling may have originated with the concept of graph theory.

Two major concepts in graph theory are that of the node and the edge, which are both

not necessarily bound to geographic coordinates and can be employed outside of a GIS.

Perhaps most similar is the ESRI geometric network, which is used for routing network

problems like shortest path analysis and the traveling salesman problem. The

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Geometric Network borrows heavily from graph theory. An additional first concept that

emerged to make connectivity modeling possible is Dijkstra’s algorthim, which finds the

shortest path between two nodes in a network (Urban et al. 2009).

In ecological applications, a node represents a habitat patch, which may have a

greater or lesser value in habitat quality, and an edge may represent some structural

component to the landscape. Urban and Keitt (2001) described the graph theoretic

perspective in landscape applications as a way to evaluate functional connections

between habitat patches and dispersal corridors, and that the gain or loss of edges and

nodes is a way to evaluate different management strategies. They demonstrated how

Mexican Spotted Owl habitat in the American Southwest can be evaluated using patch

structure and edge length to determine impact of different habitat alteration scenarios.

Urban et al. (2009) argued that graph theory is perhaps the best suited device to model

landscape connectivity since it provides insights into conservation and ecology that no

other geostatistcal device can. This might be the evaluation of habitat patch resistance

to disturbance or how to optimize a habitat mosaic.

McRae et al. (2008) took a permutation of graph theory as it evolved in electrical

engineering theory and proposed that electrical circuit theory may best represent an

ecological network. Nodes were defined as resistors and edges as currents with a

positive or negative charge. They suggested that statistical software is able to compute

landscapes of much higher complexity with thousands of habitat patches in a

reasonable amount of time rather than some GIS applications and is therefore a more

apt tool in the understanding of landscape complexity. Shah and McRae (2008)

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developed a software package called CircuitScape that automatically converts

geographic vector and raster data into a circuit network and creates measures of

voltage and resistance across a landscape. They argued that CircuitScape will help to

analyze larger landscapes due to its ability to take large datasets and provide a

sophisticated analysis without the aid of large research-grade computers.

2.1 Least cost modeling

Also progressing along a parallel track throughout the last couple decades is the

use of GIS software to compute connectivity, such as ESRI’s ArcGIS. Graph theory and

GIS methods to measure landscape connectivity adapt Dijkstra’s 1959 algorithm which

computes least cost paths between edges and nodes. ESRI’s Spatial Analyst extension

contains a set of tools to find the least cost path between two points on a landscape

using a raster surface that represents cost of movement through each raster cell. This is

essentially modeled as a line the width of the raster lattice cell between the two points.

Most research is an applied use of connectivity measurement in the form of

least-cost analysis with the development of cost surfaces (Adriaensen et al. 2003;

Driezen et al. 2007; LaRue and Nielsen 2008; Pinto and Kiett 2008; Urban et al. 2009).

Cost surfaces measure the cost to traverse from one point on the landscape to another.

Least cost modeling begins from the notion that terrestrial habitat patches are often

surrounded by a complex mosaic of land cover types ranging from perfectly hospitable

to hostile to movement by wildlife (Ricketts 2001). It is then complemented by the

ideas of graph theory, which postulates that connectivity can be quantified though the

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measurement of edges and nodes along possible paths between two points (Pinto and

Keitt 2008). The general idea of connectivity is then applied to ecological space by

defining the ecological linkages across a landscape.

Landscape connectivity can be broken down into structural components, as well

as functional components. Structural connectivity refers to characteristics of the

landscape such as habitat suitability, while functional connectivity refers to the potential

mobility in a given landscape and takes into consideration elevation change across a

landscape, a waterway or human interference to movement (Adriaensen et al. 2003).

Least-cost modeling can be used to model landscape connectivity based on the patterns

and processes of a landscape based on the potential mobility of a species.

Calculating a least-cost path is based on a simple algorithm that calculates any

given movement from one cell to another and chooses the neighbor cell with the lesser

value. Each cell has eight neighbors and thus eight potential directions that may offer

the least cost of travel (Figure 1).

Figure 1. Theoretical least cost movement across a cost surface

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Adriaensen et al. (2003) described the algorithm as movement from cell Ni to

Ni+1, as shown in Equation 1 for least-cost movement. Ni is the accumulated cost of

movement along the path and Ni+1 is the final value of the neighbor cell. The cost to

reach cell Ni is calculated using resistance values (r) of each cell plus the average cost to

move through cell Ni and Ni+1.

푵풊 ퟏ = 푵풊 + 풓풊 풓풊 ퟏퟐ

Equation 1

Using an average cost make the relationship symmetrical since movement can occur in

either direction, as shown in Equation 2 for diagonal least-cost movement. When

moving between diagonal cells, the cost is multiplied by the square root of two to

compensate for the increased distance between the cells.

푵풊 ퟏ = 푵풊 + (풓풊 풓풊 ퟏퟐ

) × √ퟐ Equation 2

Cost is described as a friction or resistance value and is the first of two surface

layers used in the calculation in the GIS. The second surface layer is the source layer,

which represents the habitat patch or the source from where the connectivity will be

calculated. Several studies utilize the least-cost tool available within ESRI ArcGIS

Toolbox to derive a calculation (Adriaensen et al. 2003; Cushman et al. 2009; La Rue and

Nielsen 2007; Driezen et al. 2007; Rabinowitz and Zeller 2010).

Determining friction/resistance is an essential component to modeling

connectivity. Several studies have developed resistance surfaces derived from habitat

suitability models of one to several focal species (La Rue and Nielson 2007; Beier et al.

2007; Rouget et al. 2006). Chetkiewicz et al. (2006) had argued that resource selection

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functions, which are based on species occurrence data, provide a better framework for

modeling wildlife movement. Rouget et al. (2006) produced a least-cost model based

on an amalgam of environmental factors, including vegetation type, habitat suitability,

protected area location, and future land-use changes. Beier et al. (2007) suggested that

a resistance surface be developed by essentially creating a grid of values that are the

inverse of the habitat suitability model. Raster cells that have high suitability or

likelihood of species occurrence will score low on resistance to travel across the cell.

The least-cost resistance layer is formed by combining multiple GIS datasets that

represent the landscape matrix, and then reclassifying them to reflect either arbitrary,

field-based, or empirical values representing resistance. An example of field-based

values comes from Ricketts (2001) where butterflies were measured for resistance of

movement between two points between landscape matrices defined by the tree

species, which were either conifer forest or willow thicket. Individual resistance values

were obtained for several butterfly species by mark recapture and a maximum

likelihood equation, which produced index values ranging from 0.9 to 12.6. Another

example is Cushman et al. (2009) who attempted to measure resistance values for

movement of grizzly bears using molecular genetics to determine bear movement, and a

set of GIS datasets (land cover, roads, slope and elevation) within a statistical modeling

framework to determine the correlation between movement and matrix. The bear

movements that correlated with landscape matrix heterogeneity provided evidence for

the determination of resistance values. Another variation in determining resistance

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from empirical studies comes from Driezen et al. (2007) who assigned resistance values

based on hedgehog Erinaceus europaeus occurrence data points within land cover type.

Beier et al. (2007) reviewed two common algorithms to combine the relative

influence of multiple criteria. The weighted arithmetic mean, which is differentiated

from a normal mean by its exclusion of dividing the sum by the number of inputs,

assigns a percentage weight to each criterion, as shown in Equation 3 for the weighted

arithmetic mean. It is multiplied by the likelihood score

푺풖풊풕풂풃풊풍풊풕풚 = ∑(푺풏 × 푾풏) Equation 3

where S is the score for criterion n and W is the weight or percentage for criterion n.

The weighted geometric mean is an appealing algorithm to use because it is very

sensitive to each criterion contribution to overall likelihood. It is found by again

assigning a percentage weight to each criterion, but then exponentiating the percentage

with the suitability score, as shown in Equation 4 for weighted geometric mean, where

푺풖풊풕풂풃풊풍풊풕풚=∏(푺풏푾풏) Equation 4

S is the score for criterion n and W is the weight or percentage for criterion n; ∏ means

to multiply the n terms. For instance, if the species is highly sensitive to roads or urban

areas, but other criteria score high, the geometric mean will ensure that the final

suitability score is low. As is noted in Beier et al. (2007), this reflects Liebig’s law of the

minimum, where the potential of an individual is determined by the most limiting

factor.

When species occurrence data is not available, researchers have asked experts

of the species or ecosystem being modeled to score criteria relating to likelihood of

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occurrence. The criteria then incorporate expert-based opinion into a resistance layer

through the use of an analytical hierarchy process (AHP). La Rue and Nielsen (2007)

derived weights for five criteria (land cover, human density, distance to paved roads,

slope, and distance to water) into a habitat suitability model using an AHP derived from

the inputs of eleven experts. AHP is a two-step process that utilizes pair-wise

comparison matrices that clarify the relative weight of each attribute (e.g. grassland)

within a criterion (land cover), and then summing them to determine a weight for each

attribute. The attribute weights and then criterion weights from each expert are

averaged to derive the final weights to use within the suitability raster surface. They

found that through the use of their AHP, they were able to predict species occurrence

68% of the time.

Criticism of expert-based species prediction and the subsequent resistance

layers produced has been leveled on the basis of the “subjective translation” that such a

process incorporates into the scientific process (Beier et al. 2008). The overwhelming

tendency for researchers to incorporate subjective measurements into the design

process may originate from the availability of enough GIS datasets to form a prediction

hypothesis and a lack of field derived species occurrence and movement GIS datasets.

Moreover, with several studies of field-based occurrence data in which expert-based

models are validated, such as La Rue and Neilsen (2007), there is no empirical evidence

for eliminating the incorporation of expert-based opinion. However, if species

occurrence data does exist, a new suite of analysis tools may be employed, such as

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resource selection framework, or habitat selection framework that can help provide

more meaningful evaluations of efficacy.

The criteria used to develop resistance surfaces within a GIS are often dependent

on available data resources. A model is therefore frequently constructed with GIS data

that is easily accessible and available for download via the World Wide Web. Land cover

is an often used data layer and is frequently the largest determinant of reclassified

resistance values. Other frequently used criteria include factors related to human

disturbance (roads, population density) and topographic factors such as hydrography,

elevation, slope, and aspect (Beier et al. 2008).

Determining the width of the wildlife corridor is the next major step in the

design process. Through a simple least-cost path analysis, a corridor is initially designed

to be one cell raster in width, which is not representative of the entire area needed to

assure representation of wildlife movement. Several methods have been explored in

the quest to obtain the most representative width needed to assure the patterns and

processes of ecological functions are attained. Rouget et al. (2006) used the concept of

irreplacability of biodiversity to determine the final corridor width. They used a

separate modeling process to quantify irreplacablity values of what they deemed

planning units. Planning units were used to measure the overall contribution of raster

cells to biodiversity, as well as to incorporate conservation management practices into

the overall design of the corridor. Beier et al. (2007) advised to widen corridors to

buffer against edge effects (e.g. invasive species, human disturbance) to conserve

ecosystem processes through the accommodation of multiple focal species’ minimum

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area for viability, and to ensure the ability of wildlife to adapt to climate change. The

minimum area needed for viability of a species ensures that a focal species can carry out

many of its life functions, such as foraging, breeding and nesting within the space

provided by the corridor. Applying the least-cost algorithm in several iterations that

models the top tier of least-cost paths available may be a useful technique to determine

the biological usefulness of the modeled corridor (Adriaensen et al. 2003). Pinto and

Keitt (2008) employed this idea and used an iterative technique to model the top ten

percent of least-cost pathways that ultimately modeled a wide variety of possible

movement routes through the landscape. They used a modified least-cost algorithm

called Conditional Minimum Transit Cost (CMTC) to determine multiple least-cost

pathways between habitat patches, as shown in Equation 5. The raster cell that is used

as the origination point contains a vertex (V) at the center of the cell that contains a line,

or technically called an edge, extended to each of its eight neighbor cells. The nearest

habitat patch cells are considered source vertices (S). The target vertex (T), is the

destination habitat patch cell. CMTC is calculated as:

푪푴푻푪(푽, 푺,푻) = 푪풖풎풎풖풍풂풕풊풗풆 푪풐풔풕(푽,푺) + 푪풖풎풎풖풍풂풕풊풗풆 푪풐풔풕(푽,푻)

Equation 5

2.2 Modeling habitat

As previously mentioned, the concept of a focal species has emerged as an

effective conservation target when planning due to the simplicity in designing for only

one species. The idea is that a paradigm species, such as the African elephant, can serve

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as an umbrella for a great number of species whose habitat will also be conserved as the

elephant’s habitat is conserved. The elephant serves as a habitat generalist that should

serve well as a surrogate for a great many other species. Caro and O’Doherty (1999)

suggested that effective protection of a viable population is assumed to protect

populations of other sympatric members of the same guild, biota at lower levels or

appreciable parts of the ecosystem. Landscapes are dynamic and species requirements

vary. While this project is only using one focal species, it should not be taken for

granted that one species be considered adequate for comprehensive conservation

planning (Beier et al. 2008).

The African elephant is a widely studied species. A great deal of literature has

been published that describes elephant habitat use, movement patterns and behavioral

choices around human populations. These papers form the basis for a cost surface since

a new set of field measurements is not a viable option for this Master’s thesis.

Furthermore, the models were developed within the limitations of a desktop analysis.

Galanti et al. (2006) investigated the space and habitat use of the elephant in an

area several hundred kilometers to the northwest of this research paper’s study area in

central Tanzania. They used GPS satellite radio-tracking to develop a spatial

representation of home range of several individuals in multiples herds of both male and

female elephants during the wet and dry season. Home range is a polygon of the

combined GPS point locations taken. The Africover Tanzania Multipurpose Landcover

Database (Di Gregario 2002) was used to determine land cover characteristics which are

essentially an explanation of habitat type. Habitat utilization was calculated by dividing

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the area of habitat type by the total home range of the individual. This produced a total

percentage of habitat use by averaging each individual’s habitat utilization.

Harris et al. (2008) found that elephants in southern Africa have preferences for

proximity to water and available vegetation for forage, and a low tolerance for humans.

They put GPS collars on nine elephants in two locations, one in an elephant reserve in

Mozambique and another in a national park in Namibia. For vegetation data, they

created a land cover dataset developed from Landsat 7 ETM+ imagery classified into

nine classes. Water was also interpreted from satellite imagery for the wet and dry

seasons. They also mapped homesteads and village locations from regional maps. Then

they took their tracking data and calculated the probability of movement from one

location fix from the GPS to the next location fix. From this they concluded that

elephants traveled up to seven to nine kilometers away from water sources during the

dry season and ten to twenty-three kilometers away from the water during the wet

season. They also concluded that elephants prefer areas of high vegetation rather than

low. Finally, they found that elephants prefer distances from humans of at least five

kilometers. The final conclusion of the paper was that, “elephants prefer to move little,

eat well, drink easily and avoid people (Harris et al. 2008).”

Ngene et al. (2009) determined the percentage of positive correlation to habitat

factors that affect elephant distribution around a national park and reserve in north

central Kenya. They investigated proximity to water, shrubland, forest, distance to

settlements and distance to roads. Using elephant location data from GPS collars, they

developed point data following nine elephants over an eleven month period. They then

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used the point data to extract distance values from each factor. They found that factors

explaining the presence of elephants in their study area are thirteen percent distance to

drinking water points, eleven percent distance to seasonal rivers, fifteen percent due to

elevation, ten percent due to presence of shrubland, nine percent presence of forest,

eight percent distance to human settlements, and seven percent distance to minor

roads (Ngene et al. 2009). They found that by using these factors they could predict

elephant distribution seventy-three percent of the time.

Of additional note in the evaluation of factors involved in the prediction of

elephant distribution is the total slope that an elephant is willing to tolerate as preferred

habitat. Wall et al. (2006) observed that elephants occur in a downward trend as the

elevation gradient increases and that a slope of thirty degrees is the maximum angle

they can tolerate. This corresponds with evidence suggested in Ngene et al. (2009)

where they found that elephants most prefer flatter areas and that they risk potential

injury when traversing along a high gradient.

Evaluating the final corridor design may be the final step in the ultimate design

process. All layers used are subject to inaccuracy (land cover, etc.), and a corridor

should be ultimately evaluated for contribution to overall conservation goals. However,

a universal theory on how to do this remains somewhat elusive since there are so many

variables within the design of a corridor to evaluate. Rouget et al. (2006) focused on

evaluating designed corridors through comparison with simple river buffers. They then

contrasted the extent of natural area, thicket representation (a major ecotype in their

study area), elephant suitability, conservation of vegetation targets, achievement of

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process targets (along hydrologic gradients), avoidance of land-use pressures and

amount of protected areas found within the designed corridors and within the river

buffers. Beier et al. (2007) suggested evaluating a corridor on the basis of three tools:

frequency distribution of habitat quality for focal species, quantification of intensity and

length of bottlenecks within the corridor design, and a list of the longest distances focal

species would have to travel between habitat patches. A frequency distribution of

habitat suitability for focal species would provide a metric for comparing alternative

corridors. Bottlenecks occur within corridors when optimal cell rasters are limited, and

might create an impassable zone within the corridor. These should therefore be

identified and evaluated as potential disrupters to corridor functionality. Rabinowitz

and Zeller (2010) decided to label bottlenecks within their modeled corridor “corridors

of concern,” to be evaluated on a case by case basis. They feared the bottlenecks could

sever corridor functionality and prohibit the genetic exchange between species

populations. Similar to bottlenecks within modeled corridors is the concern of longer

distances needed to travel between habitat patches. Statistical analyses can be

performed based on the total length of least-cost pathways, which can provide a

measure of corridor effectiveness.

Lastly, it may be of use to discuss the importance of stakeholders for the design

and implementation of wildlife corridors. Beier et al. (2007) found that ignoring the

stakeholders who would ultimately implement the plan and simply focusing on the

design of a wildlife corridor was the first error made in their study. Involving

stakeholders at the beginning of the process engenders ownership of a wildlife corridor

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implementation project, helps to eliminate “turf battles” between disparate agencies

working in the area and has shown to increase overall chances a project will be

successful (Beier et al. 2007). Since this project is located in Tanzania, some lessons may

be learned from previous conservation projects there. Harris and Hazen (2006)

described how the exclusion of Maasai knowledge in Northern Tanzania marginalized

local populations and discouraged flexible management strategies, to which local

knowledge could have been helpful. Understanding these flaws of corridor design from

a remote location may prove useful later, as concern for a rigid methodology may be

tempered with the notion that the design is perhaps flawed from the commencement of

any seriously designed conservation project without stakeholder involvement.

Conservation around the globe is at a critical moment due to explosive

population growth, agricultural intensification and land use conversion. The traditional

paradigm of islands of protected areas amongst an inhospitable matrix will lead to the

decline of ecosystem health. Douglas-Hamilton et al. (2005) described their study area

in southern Kenya and northern Tanzania as an archipelago of isolated protected

“islands” within a “sea” of pastoral areas. The protected areas of southeastern Tanzania

–which protect some of the last vestiges of east African coastal forests – are similarly

becoming “islands” in a sea of inhospitable matrix of human development. These

protected areas are of global biodiversity significance, and it may serve conservation

goals well to define their ecological connectivity using the elephant as a flagship species.

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Chapter 3: GIS modeling of connectivity

The concept of a connectivity model is fairly simple. There is an origin, a

destination and the path of least resistance found in between them. However, applying

this method to a network of 107 protected areas over a 500,000 km2 area is rather

complicated. There are a multitude of decisions, tradeoffs and determinations of

whether the methodology is consistent with the ecological theory made throughout the

process. Additionally, the entirety of this research was completed through a desktop

analysis. There was no possibility of field work, nor is it within the scope of a master’s

thesis to complete such a large project. The overall goal is to create a foundational

prototype of ecological connectivity amongst a protected area network in the study

area. To actualize the research, field validation through the monitoring of actual

elephants, and perhaps a few other focal species, as well as an accuracy assessment of

geographic information would have to be completed.

3.1 Study area

The study area (Figure 2) contains 107 protected areas that extend to the

eastern coast of Tanzania. Selous Game Reserve, which is positioned just to the west of

the study area, has been designated as a United Nations Educational, Scientific and

Cultural Organization (UNESCO) World Heritage Site due to its globally significant

biodiversity (United Nations Environment Programme 2008).

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Tanzanian national parks and reserves contain one of the world’s most

impressive assemblages of large mammals worldwide (Newmark 1993). These include

the critically endangered Black Rhinoceros Dicero bicornis, African Wild Dog Lycaon

pictus, and the African Elephant Loxodonta africana. Also within the study area, and

represented in many of the protected areas east of Selous Game Reserve, are coastal

forests and directly along the coast, mangrove forests. Coastal forests of eastern Africa

Figure 2. Study area

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support a high number of endemic genera and species of plants and animals, including

six bird species, two mammals, six reptiles, five amphibians and at least 50 invertebrates

and 100 vascular plant species (Burgess et al. 1993). There are 92 endemic forest tree

species (White 1983). Conservation International - a US-based international

conservation organization - has rated the coastal forests of eastern Africa as a global

hotspot for biodiversity (Myers et al. 2000).

The network of protected areas was chosen due to their contribution to the

protection of the last vestiges of coastal forest in east Africa (Burgess and Clarke 2008),

and because population growth, infrastructure development, and agricultural expansion

are all threats to conservation in the region. The elephant was chosen as a focal species

due to the species’ tendency to be a habitat generalist and a roamer across wide areas

of land. The elephant is known to occur with high frequency across the south Tanzanian

landscape and has widespread recognition. The study area extent was drawn to include

the protected area network surrounding the final vestiges of coastal forest in the region.

A total of 107 protected areas are established in the study area. A buffer of ten

kilometers was created to prevent the constraining of the analytical window, so that

areas of high habitat quality that might facilitate movement were given enough area to

be identified within a reasonable distance (Beier et al. 2008). Providing an initial design

for the conservation of ecological connectivity could be helpful to the preservation of

ecological integrity of the protected areas within the study area. During the last 35 - 85

years, there have been six species of large mammals that have become locally extinct

within six Tanzanian parks (although not located in this study area). Habitat

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insularization from habitat conversion outside of protected areas within the Tanzanian

protected area reserve system (Newmark 1996) highlight the urgency to adapt

management strategies.

3.2 Methodology overview

The first step is to determine where to place the origin and destination points.

Modeling connectivity is widely defined as occurring between source habitat patches.

Therefore choosing which habitat patches would represent a node within the

connectivity network represents a major design point since the inclusion of each habitat

patch in the study area would present a major computational challenge since there

would likely be thousands of points. If it was decided that each habitat patch needed to

be found, a minimum area would need to be established as a threshold for what is a

suitable and not a suitable patch size. This was not the chosen procedure due to the

theoretical and computational challenge of establishing criteria for suitable patches.

Considering the scale of the study area and the large protected area network, it was

decided that each protected area could be considered a kind of meta-habitat patch, and

that the centroid of each protected area polygon could be used as origin/destination

points. The centroid of each polygon was not always found to be well within the

boundary of the polygon. As can be seen in Figure 2, the shapes of the polygons were

often polymorphic and the centroid could be found even outside of the polygon

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boundary. This is due to the GIS operation that determines the centroid by determining

the average geometric center of the polygon.

Using the protected area polygon centroid, rather than habitat patches,

represents a slight departure from traditional corridor design, but it presents a viable

option for the scale at which the model was run and is in line with the concept that

protected areas are the main nodes of analysis. A major assumption, with some

reasonable justification, is that protected areas in the study region always contain high

quality habitat for elephants. Douglas-Hamilton et al. (2005) found that elephants spent

an average fifty-five percent of their time within protected areas, with the remaining

time spent outside of protection. This indicates that elephants use protected areas at

least a majority of the time, but depend on the landscape outside of protected area

boundaries to complete the movement necessary to forage, breed and reproduce.

All geographic data was clipped to a drawn rectangle surrounding the study area.

A buffer of ten kilometers from the outlying PAs was ensured so that ecological

connectivity would have a reasonable space to be modeled. All geographic data was

then reprojected out of its native format into UTM Zone 37S, which is the UTM zone in

which the PA network in southern Tanzania is located.

The next major step is the development of a cost surface (Figure 3). A cost

surface is the raster grid that is used to calculate least-cost paths. Based on the

literature review and on available geographic datasets, it was determined that the

factors used to model connectivity would be proximity to water, land cover, slope,

proximity to roads, and human density. These factors are assumed to be fairly good

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predictors of elephant habitat suitability according to Ngene et al. (2009). The cost

surface is essentially a habitat suitability grid representing the ease of movement across

each raster cell. If habitat suitability is high, the cost of movement will be low; and the

inverse for a cell with low suitability.

Figure 3. Flowchart of cost surface development

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Elephants exhibit different behaviors in the wet and dry season primarily due to

forage and drinking water opportunities that are less frequently encountered during the

dry season. For the purpose of finding connectivity and thus finding areas of high

conservation value, it was decided that the model would be designed for the dry season.

During the dry season, intermittent streams are often not longer viable drinking water

sources, and an elephant’s range is reduced significantly due to their frequent need for

hydration (Ngene et al. 2009). The dry season is the most critical for the elephant’s

resource availability and a loss in critical dry season habitat would be detrimental. The

model was therefore developed to reflect this seasonal characteristic since potential

movement is most restricted during this time of the year.

3.3 Water

The water dataset (Figure 4) was obtained from two sources. Water is an

extremely important predictor of wildlife in Tanzania, and is the single biggest predictive

factor for elephant presence (Ngene et al. 2009). There were a few available water

datasets to choose from. World Wildlife Fund (WWF) Hydrosheds data provides global

drainage network data derived from Shuttle Radar Topography Mission (SRTM)

elevation data (Lehner et al. 2008). Hydrosheds was modeled by finding the elevation

gradients suitable for water drainage and does not verify the actual presence of water,

however when visually compared with satellite imagery on Google earth (date

unspecified) the drainage gradient is unmistakable across the landscape. A shortcoming

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of Hydrosheds data is that it does not represent wetlands, lakes or ponds that might

also serve as potential drinking spots for the elephant. Therefore, polygon data

obtained from a website data warehouse of Tanzania GIS data called the Tanzania GIS

User’s Group was used in addition to the Hydrosheds data. The Tanzania GIS User’s

Group lakes dataset included mostly small lakes along drainage gradients. It is assumed

Figure 4. Data used for water source

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that this combination of Hydrosheds and the Tanzaniza GIS User’s Group geographic

data would create the best option for using water within the model.

To process the water data, the polygon data was converted to polyline and

unioned with one another. The distance to the nearest polyline was then calculated

across the study area. Harris et al. (2008) found that during the dry season elephants in

a reserve in a fairly proximate region south of the study area typically stay within two to

three kilometers of water and are found with much less frequency seven kilometers

away. Ngene et al. (2009) observed that elephants require drinking water every one to

two days and that the average for an elephant point location was 4.3 km, with up to

seventy-five percent of locations being between 1.7 and 6.1 km from drinking water

points. Elephants were sometime found up to 25.6 km away from water sources. These

two papers were synthesized to a set of categorical distances to water (0-1 km, 1-3 km,

3-5 km, 5-26km, and >26 km). The categories were then ranked on a scale of 1 to 10 of

likelihood that the distance to water in the raster cell would facilitate elephant

movement (Table 1). The values were chosen based on imprecise selection of values

offered by Ngene et al. (2009) and Harris et al. (2008). Each paper contained graphs

with values derived from analyses of elephant location in relation to resource location.

These graphs were used to heuristically derive the selection of values of 0 to 10.

Table 1. Values given to distance to water criterion

Km Scale (0 -10) 0 - 1 10 1 - 3 6 3 - 5 4

5 - 26 2

>26 0

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3.4 Land cover

Land cover is another critical component to model design. Perhaps the best

descriptor of landscape composition, land cover describes the types of habitat of an

elephant. Choosing a source for the land cover data was a rather difficult process, since

it was not clear which dataset would be most suitable. It was determined that there

were two appropriate choices in land cover datasets. Globcover is a global land cover

dataset produced by the European Space Agency (ESA). It contains twenty three land

cover classes, has a resolution of 300 meters, and was derived from the EROS satellite

imagery from the year 2008. An accuracy assessment provided by ESA found that

Globcover correctly identifies landscape characteristics fifty-eight percent of the time.

Africover is officially called the Tanzania Multipurpose Land Cover Database. The

Africover project has a set of land cover datasets available for several African countries.

The Tanzania land cover dataset will be referred to as Africover for simplicity. Africover

was developed through visual interpretation of Landsat 5 Thematic Mapper (TM)

imagery obtained between 1997 and 1998 (Di Gregario 2002). Since it is a visually

interpreted land cover dataset and is polygon data, there is no resolution given. They

do, however, measure the smallest land area as a minimum mapping unit of 25 hectares

(Di Gregario 2002). They do not provide an accuracy assessment, but provide some

guidelines to assess accuracy if needed.

Obtaining cost surface values for land cover was perhaps the most difficult part

of this research. There are a variety of methods used to determine elephant habitat

preferences in regards to vegetation and potential foraging opportunities. Each of the

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papers reviewed a different measure of land cover to derive preference values. Many

used a dataset that was privately created and is publicly unavailable. It was considered

a possibility to create an original land cover dataset for this research paper, or perhaps

something as simple as NDVI or a greenness index, but it would have been difficult to

assess accuracy and to use literature to demonstrate corresponding elephant

preferences.

The only study that did use a publicly available dataset and happened to be in

close proximity to the study area of this research was Galanti et al. (2006), whom used

Africover. Their study area was Tarangire National Park, located about 250 km

northwest of the study area of this resarch. Due to the proximity and the likelihood that

the elephants in the study area exhibit similar habitat foraging characteristics, results

from their research were incorporated into the land cover evaluation. Additionally,

since they used the Africover dataset, there would perhaps be less propagation of error.

This is another unverified, but acknowledged assumption made in the design of the

model.

The Africover dataset was produced by the Food and Agriculture Administration

(FAO) of the United Nations and was obtained through direct contact with them. It was

designed to be a multipurpose dataset with a customizable classification system for

agricultural and environmental applications. There are 119 available land cover classes.

These are listed in Appendix A. The land cover classes were generalized and reclassified

to reflect the eleven classes that were used in the Galanti et al. (2006).

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Galanti et al. (2006) determined percentage of habitat usage by elephants within

each land cover class during the dry season. The percent use was converted to the

likelihood from one to ten by creating a ratio to determine preferential scale. The land

cover class with the highest usage was open shrubland with 32.6 percent use. This was

classified as a 10. Each land cover class thereafter was divided by 32.6 and multiplied by

10 to derive habitat preference by the placement on the scale of 1 to 10 (Table 2). The

scaled values were accurate to the millionths place.

Table 2. Integrated Galanti Land Cover Classes into Preferential Scale

Percent Use

(Galanti et al. 2008) Scale (0 – 10) Agriculture

2.2 0.674847

Closed forest

1.7 0.521472 Open forest

17.8 5.460123

Savannah

7.4 2.269939 Savannah with trees and shrubs 17.9 5.490798 Closed shrubland

6 1.840491

Open shrubland

32.6 10.000000 Non-permanent water 13.3 4.079755 Permanent water

0.9 0.276074

Urban 0 0.000000

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3.5 Slope

Elevation across the study area is not a significant factor to elephant movement.

There are no regions with significant hills or mountains except in the northwest region

and some patches of hills scattered throughout. For this reason, slope was used as a

derivative of elevation.

The elevation dataset was derived from SRTM provided on the ESRI Data and

Maps 9.3 Media Kit. The SRTM data is minimally processed by ESRI to remove sinks and

other anomalies. The data was extracted and clipped to the study area and then

processed through the Spatial Analyst slope tool to provide a degree slope on a per

raster basis.

Wall et al. (2006) found that elephants can navigate a slope of up to 30 degrees,

and that any additional inclination prevents their movement because of their physical

frame and walking mechanics, even when foraging potential exists beyond that

threshold. They also found a fairly consistent trend of elephants preferring flat ground,

with an overwhelming majority of elephants found between 1 and 5 degrees slope. To

incorporate this into the model, the slope raster was reclassified into five classes; 0 -5

degrees was ranked 10, 5 – 10 was ranked 7, 10 – 20 was ranked 5, 20 – 30 was ranked

2, and above 30 was ranked as 0 to reflect the impossible nature of elephant movement

beyond that point (Table 3). Wall et al. (2006) provide a graph with elephant fix density

(fixes per km2) in relation to slope gradient in degrees. A fix is a record noting the

geographic coordinate of the elephant occurrence. The graph in Wall et al. (2006)

indicated there were nearly 100 elephant fixes near 0 degree slope and very near 0 fixes

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Table 3. Values used for slope criterion in model

Degrees slope Scale (0-10) 0 - 5

10

5 - 10

7

10 - 20

5

20 - 30

2 >30 0

at 30 degrees slope. This range of values was used to heuristically evaluate the ranking

of the slope classes.

3.6 Roads

The roads dataset was provided as ancillary information in the Africover dataset

produced by FAO. There was no distinction placed on the characteristics of the roads,

such as major or minor. Furthermore, the dataset may be slightly out of date, but

nonetheless provides a perspective on human presence within an elephant’s

interpretation of the landscape. The literature provides divergent viewpoints on the

influence of roads in an elephant’s habitat preferences. Ngene et al. (2009) found that

elephants in their study area are actually at ease around roads because they are

regularly patrolled by security forces who prevent poaching. They acknowledged,

however, that their case may be an exception, since many other studies found roads to

be avoided by elephants (Blom et al. 2005; Barnes et al. 1991). Elephants generally

avoid things that are associated with people (Harris et al. 2008). With this research in

mind, roads were buffered and all areas were classified as being a short, medium or high

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Table 4. Results of distance to roads analysis

Distance (Km) Scale (0-10) 0 - 1 5 1 - 3 8 >3 10

distance from a road. On the ease of movement scales of one to ten, a high distance to

road was ranked as a 10, medium distance was an 8 and short distance was ranked as 5

(Table 4). These values were assigned somewhat arbitrarily and not based on any

numbers offered within the literature. They were chosen to give roads at least a partial

influence in the design of elephant movement corridors.

3.7 Human density

Similar to roads on elephant movement, human population is another factor

related to the presence of humans and the likelihood that elephants will make a

decision to travel one way or the other. Many of the same rationales were used to

quantify resistance, such as interpreting observations about human presence and

elephant preferences from literature and assigning a somewhat arbitrary number to

reflect the spirit of the statement. Harris et al. (2008) observed that female elephants

tend to stay at least 5 km from human settlements, but that male elephants were less

cautious around humans.

The human density data was derived from LandScan global population dataset

created by the Oak Ridge National Laboratory (ORNL) for the year 2008. Landscan is a

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gridded, 1 kilometer resolution product that predicts population density based on a

series of inputs, including census counts, land cover, slope, urban areas, village locations

and high resolution imagery (ORNL 2009). The data is combined using a dasymetric

modeling approach. LandScan 2008 was obtained through special agreement with

ORNL to only use the data for educational purposes.

Since elephant movement through a matrix of resistance values was the goal of

this study, the LandScan product appeared to compliment the objectives very well. The

alternative would have been to use incomplete point data on village and settlement

locations and derive distances from population point, and then assign resistance values.

This may have made it easier to reflect statements made in the literature referring to

the exact distances elephants generally prefer from human populations. Unfortunately,

there is no authoritative point population dataset for the study area.

There was no literature found that directly referenced human density in

observing elephant behavior; however, it seemed logical to apply the scale of 0 to 10 to

the population density grid. Areas with highest population are most likely urban areas

while the sparsely populated regions have very few people per square kilometer.

To prepare the data for inclusion into the analysis, the LandScan product was

extracted by mask for the study area and reclassified to five categories of population

density. These were referred to as low, low-moderate, moderate, high-moderate and

high population density and assigned a preference factor of 10, 8, 3, 2, and 0

respectively (Table 5).

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People per Km² Scale (0-10) 0 - 5

10

6 - 50

8

50 - 100

3

100 - 300

2

>300 0

The values were chosen based on reports that elephants tend to avoid human

population (Harris et al. 2008; Wall et al. 2006; Ngene et al. 2009). The ranking of the

values were not in relation to any graph or parameter set by the literature, rather a best

guess based on the notion in the literature that elephants do their best to avoid contact

with human populations.

3.8 Cost surface

The final composition of model data was then amalgamated into a weighted

conductance surface raster. Ngene et al. (2009) provided a percent factor contribution

to elephant distribution. Through a correlation matrix of explanatory variables, they

calculated that distance to water points, distance to seasonal rivers, elevation,

shrubland, forest, distance from settlements and distance from minor roads help to

explain 73 percent of elephant distribution (Ngene et al. 2009). The remaining factors

explaining 27 percent of elephant distribution were listed as agriculture, closed

grassland, mollic andisol and chromic combisol soil types. These types had on average a

less than significant influence on elephant distribution and therefore were excluded

Table 5. Values applied to weight human population density criterion in model

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from use in the current analysis. To create an easily understood weighting system, the

significant explanatory variables were curved to represent a scale of 0 to 100 percent.

The land cover variables of forest and shrubland were combined to represent

the influence of land cover. It should be acknowledge that these two land cover types

do not compose the entirety of the land cover types used in the model. Furthermore,

Ngene et al. (2009) determined that other land cover types such as agricultural do not

factor as a significant predictive variable for elephant habitat preference. This is a

significant shortcoming of the model; however, there did not appear to be another

choice from the desktop analysis perspective. A combination of concepts derived from

literature will certainly create at least some theoretical dissonance within the

methodology. It did not seem prudent to dismiss execution the model for this reason,

and the land cover types suggested by Ngene et al. (2009) were used as surrogates for

all land cover types in Galanti et al. (2006).

Distance to water points and seasonal rivers from Ngene et al. (2009) were

combined to represent distance to water. Distance to minor roads from Ngene et al.

(2009) was considered distance to all roads, since the roads dataset did not have any

descriptive attributes. Therefore, explanatory variables to describe the ease of

movement, or conductivity of a cost surface cell was influenced 33 percent by distance

to water, 26 percent by land cover, 20 percent by elevation, 11 percent by human

density and 10 percent by distance to roads (Table 6) (Ngene et al. 2009).

The final amalgamation borrowed from concepts provided by Beier et al. (2007).

They suggest either using a geometric mean or an arithmetic mean as described in

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Chapter 2. An arithmetic mean simply multiplies each factor’s interim value, in the case

of this analysis, the conductivity score of 0 to 10, by the weight being applied to the

distribution factor. The final values are then added together to form a final

conductance value. This means that if a value of 0 is present within the factors, it will

not necessarily result in a pixel value with maximum resistance. This is particularly

unhelpful when demonstrating the effect of potential slope values that exceed an

elephant’s capability. The geometric mean takes each factor and exponentiates the

factor’s interim value by the percentage of the distribution factor. The remaining values

are then all multiplied together. If a 0 exists in the conductance factors, the ultimate

score will result in a pixel with maximum resistance, or 0 (Beier et al. 2007). The

geometric mean was used in this analysis by using the Spatial Analyst Power tool with

each raster of conductance factors and then with a simple map algebra statement

multiplying the factors together. This resulted in a raster surface of landscape

conductance values ranging between 0 and 10.

It should be noted at this point that there was an issue with the Spatial Analyst

Math tools. The land cover dataset contained values with several decimal places.

Values with decimals occur in an ESRI GRID raster as a floating point grid. The Power

tool used to exponentiate the factors was unable to process the floating point values.

To overcome this obstacle, each factor was multiplied by 1,000,000. This allowed for at

least six additional significant digits in each factor, which minimized the loss of accurate

data and maintained the integrity of the conductance value. Each value’s magnitude

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Table 6. Final amalgamation of movement cost surface values

was simply adjusted to occur in front of the decimal point. This resulted in a raster

surface of landscape conductance values ranging between 0 and 1,000,000.

Before advancing to the calculation of CMTC could commence, the preferential

value structure of the conductance surface needed to be inverted to represent

resistance rather than conductance. Using the Spatial Analyst Map Algebra tool, the

raster was multiplied by -1 and then 1,000,000 was added to it. This reversed the

conductance raster to place the most facilitative movement values at the low end of the

spectrum. This concluded the first half of the analysis, and the cost distance calculation

could commence.

3.9 Corridor modeling with CMTC

The second half of the analysis (Figure 5) consisted of finding the least cost

corridor between each PA centroid in the PA network. There were 107 total PAs. A

method was devised to run an analysis between PAs that were in direct path of each

other. Developing the list of PAs to run the least cost analysis was completed by

manually generating straight line paths between each PA centroid. So that PA names

were easily entered into path file names in the variety of GIS tools used, unique ID

Criterion

Weight given (%) Distance to water

33

Land cover

26 Slope

20

Human density

11 Distance to roads 10

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numbers were assigned as a surrogate to the text names of the PAs. This significantly

facilitated ease of model execution. The original PA dataset, obtained from World

Database of Protected Areas (International Union for the Conservation of Nature and

United Nations Environment Programme 2010) was a collection of PAs in one shapefile.

The centroid of each PA polygon was used as the origin and/or destination of the least

cost corridor. Therefore, using the Data Management toolbox Feature to Point tool,

each polygon was converted to a point within the centroid of the polygon.

A simple model in Model Builder was created to extract each PA individually.

The Iterate Feature Selection tool was used to run through each row of the centroid

shapefile of PAs. Then the Copy Features tool was used to extract each PA individually.

In this way, each protected area was made into its own point feature class.

Figure 5. Flowchart for modeling corridors

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To manage the list of least cost paths, the study area was compartmentalized

into five zones (Figure 6). Each zone was identified based on an evaluation of major PA

point clustering (Appendix B). There appeared to be three major zones of clusters.

These were identified as Zone A, Zone B and Zone C. In order to model between each of

the zones, two interzones were created; these were named Interzone A and Interzone

B. Zone A consisted of the twenty-nine PAs found in the northwest section of the study

area. Zone B consisted of fifty PAs found in the central portion of the study area. Zone

C consisted of thirty PAs found in the southern section of the study area. Interzone A

consisted of the eleven PAs that link Zone A and Zone B, and Interzone B contains

eleven PAs that link Zone B and Zone C.

A zone of analysis was drawn as a bounding box around the protected area

centroids using the Feature Envelope to Polygon tool in Data Management Tools in

ArcGIS Toolbox. Then, using the Buffer tool, ten kilometers were added to the bounding

boxes to provide enough distance from the exterior PAs to allow for ecological

processes to be modeled . To find the least cost path between PAs that connected

across the zone boundaries two sets of interzone areas were delineated in the same

fashion as each zone area. Interzone A was inadvertently cut short due to the

previously set analysis mask, however, this was only along the north sections and did

not impact corridor modeling.

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Figure 6. The extent of each zone of analysis within study area.

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3.9.1 Modeling corridor width and alternate dispersal corridors

At this point in the analysis modeling the correct corridor width becomes an

issue. Beier et al. (2007) suggested corridor width should be set according to the

species being modeled. There are two types of species, corridor dwellers that might

carry out an entire life cycle within the corridor, or passage species which simply may

use the corridor for movement between habitat patches. For this research, the African

elephant is considered a passage species since movement between protected areas are

being modeled. For corridor dwellers it might be best to set the width as the minimum

area of viability for the species. Additionally, if more than one species is being modeled,

it is possible that the width of the corridors will be set by overlapping requirements of

each species and that this will be an effective way to model species. Minimizing

bottlenecks is perhaps the most crucial component so that species always have free

access through the most inhospitable matrix of the corridor. It might also be

advantageous to decide corridor width through a stakeholder engagement process; so

that conservation goals can be reasonably assessed with the reality of local

circumstances (Bier et al. 2007). Ultimately, they do not provide a clear rationale why to

choose widening a corridor based on species habitat requirements or through the

stakeholder engagement process.

Pinto and Kiett (2008) criticized focusing on corridor width by expanding a least

cost path because it ignores potential alternate dispersal routes that exist amongst

comparable routes with similar costs of movement. They proposed focusing on

methods that find redundant least cost paths to evaluate corridors. Drawing upon

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graph theory, they describe Conditional Minimum Transit Cost (CMTC) as a method to

find redundant least cost paths between origin and destination points. CMTC is an

analysis that can be carried out through ArcGIS processing of cost surface grids. The

research by Pinto and Keitt (2008) proposed to integrate species habitat preferences

into regional-scale depictions of habitat connectivity through the identification of

redundant least-cost corridors. This appeared to be most in line with the goals of this

analysis considering large protected area network study area. The advantage of this

approach is that redundancies are modeled through a user defined threshold of

conductive pixels across the landscape, and the top percentage of conductive pixels is

identified. The alternative is that a single least-cost path (one pixel in width) is

identified, which provides a less descriptive view of the landscape as the focal species

would experience it.

CMTC is relatively simple to conceptualize in a GIS. As described in Chapter 3, it

simply adds the two cost distance grids in both directions of movement. The cells with

the lowest values will emerge as the corridor. A cost distance raster is calculated using

the Spatial Analyst Distance tool Cost Distance. Once the cost distance in both

directions is calculated, a threshold must be set to determine cells that represent the

most conductive possible movement between PAs. Pinto and Keitt (2008) suggested

that this is somewhat of an arbitrary decision that can be made based on goals of the

analysis. In their study they chose to select the top 10 percent of cells with the lowest

resistance values. In this analysis 10 percent appeared to include too many cells, so the

percent value was reduced from 10 to 5.

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3.10 Calculating CMTC

The first step to calculate CMTC was to use the Cost Distance tool to calculate

the total value of cost of movement from each PA centroid. It was not necessary at this

point to focus on the destination of the corridor. As with the previous analyses the

processing of the cost distance results were completed through the

compartmentalization of the zones of analysis. The batch command of Cost Distance

was utilized to process through each zone. Each value was named according to its

respective PA unique ID it represented.

Obtaining the CMTC was completed by using the batch command and the Plus

tool in the Spatial Analyst toolbox to add the cost distance rasters from the source and

destination of the corridor. The manually generated connections can be viewed in

Appendix B; and the list of protected areas modeled for CMTC in Appendix C. In total,

194 PA connections were produced from the manually generated list (Table 7). Each of

the CMTC analyses was completed using the Plus tool in batch mode.

Table 7. Number of CMTC analyses completed

Zone of Analysis Number of PAs Number of CMTC analyses completed

Zone A 29 51 Zone B 49 83 Zone C 29 42 Interzone A 11* 11 Interzone B 11* 7 Total 107 194

*Interzone PAs are composed of PAs within Zones A though C and therefore are not counted in total

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3.10.1 Finding five percent

Obtaining the top five percent of cells was a relatively complicated process.

Model Builder in ArcGIS was used to create a series of tools that processed the

information obtained for each PA corridor. The process began using the Iterate Rasters

tool to automatically run through the folder of CMTC rasters from each zone that had

been processed. Each cell value in the cost surface was in the magnitude of tens of

millions from the upscaling to rid each value of a decimal point so that Spatial Analyst

would perform more smoothly. When the cost distance analysis was performed, it

created some raster cell values in the tens of billions range since the operation

accumulates cost surface cell values as distance increases from the source cell. To

create cell values in a more manageable magnitude, each raster was divided by 100,000

to return it back to the millions range. Even without decimal values the divide tool

creates rasters that are floating point values; therefore each raster had to be converted

back to an integer with the Int tool. Next an attribute table was built for each raster

since one did not yet exist and would be needed to create a process that finds the top

five percent values in each CMTC raster. The next steps, described next, involved

creating a series of new fields for each newly created attribute table and then

computations were executed to get to the top five percent value of each raster grid.

The Summary Statistics tool was used to find the minimum and maximum value

of each CMTC raster which is now an integer raster with an attribute table and

generates an ESRI Info table. The Summary Statistics tool automatically created two

new fields for minimum and maximum of the corridor raster. Next, using the Add Field

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tool, a field was added named “range” as a long integer, and the Calculate Field tool was

used to subtract the minimum value from the maximum value. This new value

represented the total range found in each CMTC raster so that a clean five percent could

be found. Next, using the Add Field, a new field was added named “fiveperc” as a long

integer and the Calculate Field tool was used to multiply “range” by 0.05, or 5 percent.

Finally the top value of the five percent needed to be found to establish the value that

represented up to five percent of all cell values. To do this a new field was added using

the Add Field tool, naming the new field “topvalue” as a long integer, and again the

Calculate Field tool was used to add the minimum value of the original raster found in

the “min” field first created by Summary Statistics, to the value found in the “fiveperc”

field, which was the value of five percent of the total range of raster values.

This model was executed five times through each of the zones of analysis. Once

this step was complete, a series of ESRI info tables existed with the value that

represented the top of the five percent range. This number was used as the main input

into the next step of the analysis, which was to extract the top five percent of values

from each least cost corridor raster.

3.10.2 Extracting five percent

This step extracted only the most conductive cells to movement so that the

network of PAs could be combined to display a landscape of high potential movement.

It was also possible that this step would provide individual least cost corridor

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visualizations. With these two purposes in mind, it seemed appropriate that extracting

the most conductive cells while ignoring all other values was the best way to provide a

representation of the outputs of the analysis.

To accomplish a visualization of the most conductive cells, map algebra in the

Spatial Analyst Raster Calculator was used in batch mode to calculate multiple CMTC

corridors simultaneously. The value at the top of the five percent range of most

conductive cells in each corridor was manually inserted into a conditional statement

using map algebra. The value at the top of the five percent range was found in the

“fiveperc” field of the attribute table created in the previous step. All cells not meeting

the conditional requirement were set to NoData. This was repeated throughout the

CMTC corridor rasters in each zone of analysis.

To visualize the landscape of conductive cells, the Mosaic to New Raster tool in

the Data Management Tools toolbox was used to patch together each least cost

corridor. The minimum value of raster cells that fell on top of one another was used as

the mosaic output value so that the cell most conducive to movement would be

represented.

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Chapter 4: Results and discussion

Demonstrating ecological connectivity between protected areas amongst a

backdrop of coastal forest areas was the goal of this analysis. The product

demonstrated the potential for geospatial information to model movement of a

paradigm species through a habitat matrix framework that predicted movement paths.

The following results and discussion review the initial criteria used to model elephant

habitat preference used as a predictor of movement under the assumption an elephant

will likely move along a path in structural landscape of preferred habitat with low

functional resistance (i.e. low elevation, avoidance of human population, etc.).

To interpret the findings of the corridor modeling, there are several

considerations to take into account. First is the scale at which one is trying to interpret

the findings. Each of the zones in the study area has slightly different habitat

characteristics that should be considered. Furthermore, conservation activities that

take place in one locality of the network may not be comparable to the results of

another area. Therefore increasing and decreasing the scale in order to provide a more

useful interpretation may be important. Finally, analyzing the corridor found between

each PA may be necessary when considering what types of conservation action to take.

Placing the modeled corridor in the context of a local situation may provide a

cartographic scenario - such as its placement within the matrix of agriculture, natural

land and human population - that would provide a more meaningful graphic for

visualizing elephant movement across geographic space.

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As the results are presented, the modeled corridors are presented in the local,

regional and network wide scales of analysis. This provides the best framework for

interpreting the results as a structural and functional part of the landscape, and acts as a

suggestion for how landscape-scale corridor models should be interpreted.

4.1 Cost surface results and discussion

The distance to water results (Figure 7) demonstrate the availability of drinking water

sources. The WWF Hydrosheds network of drainage channels forms a network pattern without

significant interruption across the landscape of the study area. The values incorporated into the

model were chosen to represent water availability during the dry season, since this is the time

of year that most limits the movement potential of elephants.

The results may represent an overestimation of available water sources since each

drainage network is not verified for the presence of water. The WWF Hydrosheds dataset was

processed by applying an algorithm to find the pathway of the lowest areas of elevation across

the landscape. This analysis usually results in the presence of water, since water typically

follows drainage pathways. The ubiquity of water availability across the landscape is further

perhaps overrepresented by the lack of any location farthest than the longest distance (more

than 26 km) the literature mentioned elephants were willing to travel without water (Harris et

al. 2008). It seems unlikely that there are no places in the study area more than 26 km from

water in the dry season, but it is difficult to verify without validated water presence data. Other

factors that are not included, such as subsurface hydrology, precipitation, temperature and

human use, may also govern the availability of a drinking water source for elephants, but are not

a component to this research.

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Figure 7. Results of the distance to water analysis

The land cover dataset (Figure 8) displays a landscape that is composed of roughly 30

percent agricultural, 25 percent open forest, 20 percent open shrubland, with the other land

cover classes making up the final 25 percent. An elephant would likely perceive this landscape

to be somewhat hostile due to the agricultural areas, but largely accommodating to movement

in the remaining areas. The central region of the study area is largely devoid of agricultural

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Figure 8. Land cover reclassification

activity and may present an elephant with the least amount of resistance to movement in any

direction.

The results of the slope analysis (Figure 9) show a landscape that has a low degree of

slope throughout much of the study area. Only in the northwest portions of the landscape are

there any significant degrees of slope that would impede movement. The areas coincide with a

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matrix of agricultural land use and higher human density. The northwest portion of the

landscape may therefore see a higher cost of movement. The area in the southwest portion of

the study area does not pose any real impedance to movement since this region is deep within

Selous Game Reserve and will not have any potential corridors modeled in between.

Figure 9. Result of the slope analysis

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The distance to roads analysis (Figure 10) demonstrates a landscape that does not

contain a very robust network of roads. This will likely aid elephant movement across much of

the landscape. Roads are likely associated with agriculture and human density since they

represent not only human movement, but also economic corridors. At all edges of the study

area there are higher densities of roads. This will likely make the central portion of the study

area most accommodating to movement.

Figure 10. Result of the distance to roads analysis

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The final criterion for the development of the cost surface raster is human density

(Figure 11). Following many of the patterns found in the previous datasets, the edges of the

study area have higher levels of human density while the central portion remains largely devoid

of any significant human presence. The northwest, northeast and southeast areas have the

highest density and will likely prevent elephants from moving freely across the landscape. Also

revealed are several patches of human absence. The central patch is the largest, with two

medium patches to the northwest and a series of much smaller patches along the eastern

portion of the study area. These areas will likely improve the chances that an elephant will

choose to move in these regions rather than in areas of higher human density.

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Figure 11. Human population density reclassification

The final cost surface (Figure 12) is the amalgamation of the input parameters of

distance to water, land cover, slope, distance to roads and human density. As many of the

criteria that discourage elephant movement were found roughly along the edges of the study

area, the final cost surface shows the highest costs of movement in the northwest, northeast

and southeast corners. There are several smaller pockets in the interior of the study are that

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will likely produce resistance to movement and perhaps encourage an elephant to move around

these pockets of hostile matrix.

Figure 12. The final cost of movement surface

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4.2 Zonal characteristics discussion

As previously described, the PA network was subdivided into a set of five zones

where the cost surface raster was extracted and run amongst the set of PAs located

within the zone. This helped to compartmentalize the task of running 194 CMTC

analyses across the PA network. An unintended consequence of this is that the total

area of analysis changed within each zone (Figure 13). This is significant since the land

cover prediction values were developed using likelihood values based on potential

habitat availability which are represented by the land cover classes (Figure 14). The final

corridor was delineated using the top five percent of all values within the zone. Since

each zone has slightly different ranges of total values, the top five percent probably

changes a little bit between each zone. This is most likely the most significant when

considering the effects of land cover on the model output, since habitat availability is

distinctly different across the PA network.

In each zone of analysis there was a range of approximately 17,000 to 45,000

square kilometers. The study area in Galanti et al. (2006) was approximately 17,000

square kilometers. In addition to the differences in total size of each zone of analysis,

there are different habitat compositions within each zone. Each zone has slightly

different proportions of land cover types (Figure 14).

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Figure 13. Total area in each zone

There was a certain amount of congruency, however, within each land cover

type. The percent composition was within a reasonable range with a few exceptions.

Interzone A and Zone C had much higher amounts of the open forest land cover type

than the rest of the zones. And the amount of open shrubland was somewhat more

variable than the rest of the land cover types throughout each of the zones. Open

shrubland was the land cover type most utilized by elephants, as pointed out by Galanti

et al. (2006).

0

5,000

10,000

15,000

20,000

25,000

30,000

35,000

40,000

45,000

50,000

Km²

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Figure 14. Percent of land cover type for each zone of analysis

4.3 CMTC results and discussion

As can be seen in each of the following representations of model results (Figures

15-23), the most important predictor of modeling connectivity using a least cost path

algorithm may be proximity. PAs that are near are most likely to have lower travel cost

and thus movement within the clusters of PAs is predictably conductive. When high

cost of movement does appear in areas with high clustering, or near proximate PAs,

there must be substantial restrictions to movement that originated with one of the

original parameters of the model, such as high human density or steep slopes. These

areas should be evaluated from a management perspective since they present risk to

0

5

10

15

20

25

30

35

40

45

Perc

ent o

f Tot

al A

rea

wit

hin

Zone

Galanti et al. 2006 Zone A Zone B Zone C

Interzone A Interzone B Entire Study Area

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the degradation of habitat for elephants as well as potential sites of human elephant

conflict.

Another possible lesson taken from the fact that PAs that are in closer proximity

are more effective at providing high levels of conductivity is that the PAs should be

expanded to provide more habitat for elephants and the rest of the biodiversity that

depends on similar habitat. The PAs with the highest conductivity values could be

identified as targets for expansion. It might be expected that external threats degrade

the quality of the habitat in these areas, and therefore an expansion of the total area

currently under protection could be increased.

4.3.1 Regional zone results and discussion

In the cluster of PAs in Zone A (Figure 15), there are several sets of PAs that are

close to one another, but they are surrounded by a fairly high level of inhospitable

matrix. This is especially apparent in the southwest portion of the zone where the cost

of movement between Mikumi and Shikurifumi appears to be rather high. Similar

distances are present throughout the network without such high cost of movement

values. There must be considerable barrier to movement in this area, which is most

likely due to agricultural or human population density.

Several of the PAs are in a cluster pattern and could be used as a starting point

to strengthen the reserve network in contrast to the difficult terrain of the southwest

portion of the zone. The northern portion of the zone, however, appears to be of higher

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habitat value since even in the case of longer distances between PA centroids, there is a

relatively low cost of movement.

There also appears to be a degree of clustering of PAs in Zone B where several

PAs surround Namakutwa Nyamulete. It is also likely in this area that human population

density is high. Therefore conservation effort could be established in these areas as the

matrix of hospitable habitat is less than would be expected.

There is considerable resistance to movement in the northern and the southern

sections of Zone B (Figure 16). This is a coastal region with higher human population

density than more inland. Given the current situation modeled, elephant movement

between these PAs appears to be very hostile.

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Perhaps most importantly, the areas where movement is found to be the most

facilitated by ideal conditions for movement should be used as a core elephant zone.

Several of the PAs in this region are especially important because they contain coastal

forest ecosystem remnants. It would be of high conservation value to ensure that

functional corridors are established between these PAs.

Figure 15. Model result for Zone A

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Zone C contains up to four sets of clustered PAs (Figure 17). These clusters seem

to facilitate the movement of elephants relatively well with low cost of movement

values. There are however some clear corridors that have emerged from the analysis.

Figure 16. Model Result for Zone B

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High movement costs can be found along several places in the southeast portion

of the zone. Additionally, a large gap exists in the central southern region where it does

not appear that elephant movement is likely. This leaves the most southwestern PAs

isolated from movement in and out of PAs.

Figure 17. Model result for Zone C

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The results from the interzones had notably high resistance to movement. The

distances between PAs in these areas were greater than most other connections;

however, this fact might be of value since habitat patches in these areas could be

overlooked. The area between Zone A and Zone B has a paucity of PAs and could be

evaluated as a potential site for the establishment of more protection. Additionally,

along the western edge of Zone B and Zone C is among the highest resistance in the

study area. The southern edge of the study area has the highest values of resistance.

Additionally the corridors show up as very narrow pathways that would make

movement through this region extremely difficult for an elephant.

The exception to difficult movement across Interzone A (Figure 18) is the area

between Ruvu North Fuel and Kitulanghalo, and to a lesser degree directly south at

Mkungwe. This corridor is certainly a pathway most likely taken across this space. The

high amounts of open shrubland, the most preferred land cover type by elephants, is

perhaps in highest density in this section of Interzone A.

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Figure 18. Model result for Interzone A

The space in Interzone B is not as far a distance as that in Interzone A, but there

are a variety of outputs in this area (Figure 19). The eastern section contains a cluster of

smaller PAs where the distance between centroids is minimal. It also appears that the

habitat matrix is relatively hospitable for the movement of elephants. The western

section however is perhaps the exact opposite with among the highest cost of

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movement values in the PA network. This may be in part due to the distance traveled

between Lungonya and Nyera Kipere; however, it seems undeniable that the matrix

restrains movement in this region. It may be important to analyze the threats to

conservation in this region and focus on containing additional pressures from

agriculture or human population growth.

Figure 19. Results from Interzone B

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4.3.2 Results from a single corridor

Many of the local results resembled a least cost pathway that did not

demonstrate very much differentiation between a straight line path, as was the case

with the Uluguru South to Kimboza path in Zone A (Figure 20). The habitat matrix

between the two PAs must be relatively hospitable to movement since there are no

major deviations or abrupt changes in course of the least cost pathway. It is in these

types of situations that the advantage of using CMTC and deriving a top percentage of

movement values becomes clear. The pathway reveals a corridor of sufficient width to

allow for movement across an area and thus could be used as a blueprint for

conservation planning. This is in contrast with methods that find a least cost path,

which does not give the context of the surrounding habitat matrix. A variety of

potential paths are visualized. Using CMTC, the total width of the corridor is

determined using a select group of movement values. In the case of our study, the top

five percent of values were used to represent the best corridor.

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Figure 20. Model result between Uluguru South and Kimboza in Zone A

Perhaps the most interesting corridor modeled throughout the network was the

Ntama to Lionja pathway found in Zone C (Figure 21). Rather than taking a shorter path

between the two PA centroids, the path took a circuitous and meandering form. The

most conductive pathway is to go the most circuitous way around what must be a very

inhospitable matrix. However, an alternate dispersal corridor also exists in a straight

across trajectory. Agriculture is a major land cover type in this area, in addition to

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substantial human population density. These factors must present an elephant with

little choice but to move via less resistant pathways.

Figure 21. Model result between Ntama and Lionja in Zone C

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4.3.3 Entire study area results and discussion

Each of the zones were mosaicked together to produce a full visualization of the

PA network (Figure 22). The results here should be regarded with caution due to the

difference in habitat characteristics and total size in each zone, and its effect on the top

five percent slice of movement values. The results, however, are undeniably similar to

the outputs found in the compartmentalized view of each zone. There are areas of

clustered PAs where movement is relatively easy mostly due to the short distances

traveled between PA centroids, and several bottlenecks and hostile matrix areas

become apparent. When developing a conservation strategy for this area, a

visualization such as the one produced here would provide an excellent overview of

barriers to movement in the region.

Three major gaps in movement across the region emerge from the entire PA

Network. First is the Interzone A area in the north with high resistance to movement

values in the northern and southern portions of the interzone. The next two areas of

hostile matrix are in the south, leaving the southwestern section of the PA network

isolated from the rest of the region. Movement across this space is likely rather difficult

for elephants, and the region may be experiencing declines in elephant populations due

to the pressures exerted from habitat degradation due to human activities.

On the positive side a clear set of pathways exists between many of the PAs in

the network, especially along the coastal region of the network. The most abrupt

changes in movement values exist between Zone A and Interzone A.

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Visualizing the entire study area by classifying the regions into ten movement

classes further illuminates several zones where barriers to movement are higher and

perhaps helps to place corridors in a hierarchy of resistance and conductivity to

movement (Figure 23). As mentioned previously there are several specific areas where

resistance to movement is highest. These are both located in the interzones where the

distance traveled between PAs is high.

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Figure 22. Model results for entire PA network

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Figure 23. Model result classified for PA network

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Chapter 5: Conclusion

This model of connectivity between protected areas was derived from estimates

found in literature to predict the habitat preferences of elephants in the southeastern

section of Tanzania. It is meant to provide a prototype of corridor modeling in the

region. A comprehensive model would include several other species with habitat

requirements different than the elephant as well as a habitat preference study with

point location data of elephant movement throughout the study area. This would avoid

several of the issues related to subjective translation, which Beier et al. (2008) described

as the incorporation of subjective measurements into the model. The criteria for

habitat selection were derived from Ngene et al. (2009) for elephants in eastern Kenya,

which has a very different ecology than southeastern Tanzania. This undoubtedly will

introduce uncertainty in the model. The values chosen for the habitat criteria were

derived from Galanti et al. (2006) in a region close to this study area, which may provide

some reliable estimation of elephant behavior. To test for certainty, there are several

different methods that could be used. One possible method could be to conduct a

sensitivity analysis to see how different weights given to each criterion might produce

different outcomes. Another could be to use field-based measurements from radio or

GPS collars to monitor the movement of elephants in the region. If the elephant’s

movement mimics the predictions given by the model, the results could be validated as

such. The predictions could be statistically evaluated using these field-derived points of

occurrence over time and using logistic regression or the Maximum Entropy modeling

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program to complete accuracy metrics to reflecting the appropriate prediction values

(Pittiglio et al. 2012). These analyses are outside of the scope of this study, which was

composed entirely through desktop analysis without access to field-based occurrence

points or recent land cover data. This study does provide a starting point for completing

such an exercise, however, and can be used to develop a new study that takes these

considerations into account.

A better analysis of PA management strategies also needs to occur so that the PA

network is robustly managed enough to ensure that corridors identified between each

PA can be expected to have long term persistence. The PA data was obtained from the

WDPA and contains the initial attribute data describing the management authority and

the classification by IUCN, which describes a basic evaluation of PA management.

5.1 Least cost modeling

Least cost modeling might also be more effective using habitat patches as the

node of connectivity. PA centroids were used in lieu of habitat patches due to the

difficulty in defining a habitat patch. There would likely be some threshold for

determining patch suitability for elephants as well as some requirement for a large

enough area to qualify as a patch. This again was outside the scope of this report, but

should be evaluated if connectivity is going to be modeled more accurately in the area.

The likely benefits of such an analysis would include a derivation of elephant migration

pathways with the highest degree of certainty. This is especially clear when considering

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the high resistance seen in Interzone A even though the region has the highest

percentage of open shrubland, which is by far an elephant’s preferred habitat.

The modeling process was based off of literature, however, elements of

improvisation did occur throughout due to the lack of a clear example to follow as well

as difficulty processing in the ArcGIS modeling environment. Employing a least-cost

model using CMTC across such a large landscape has never been published. Given these

circumstances, there were numerous instances when the best possible data and

methods were compromised. A new iteration of this model with better data and the

lessons learned during this modeling process would surely provide a more accurate

prediction of elephant movement across the southeast Tanzanian landscape.

5.2 The use of data

Much of the data acquired for use within the model had unique circumstances

that should be discussed as potential agents that may limit the accuracy of the model,

these include land cover, water, slope, human density and roads. Additionally the

process of constructing the model was one of improvisation based on examples cobbled

together from several research examples and trial and error within the GIS processing

environment. It is hoped that this discussion will provide some insight to the thought

process involved during the construction of this research process.

The land cover data was derived from the Africover Tanzania project by FAO.

There were no documents provided that verify the accuracy of the data, nor was it

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within the scope of this project to validate this land cover dataset. An evaluation of

accuracy for this data would have provided an element of credibility to the model.

Additionally the data was quite old. Imagery used to derive the land cover classification

was derived from the 1997 to 1998 timeframe. Imagery obtained from a more up-to-

date period would benefit the model by providing a more recent view of the landscape.

It was certainly considered an advantage that Galanti et al. (2006) used Africover data in

their research project. This may have provided an element of congruency, but

unfortunately it also may have given another opportunity for error to creep into the

analysis. Galanti et al. (2008) did not provide a document of how the land cover data

was reclassified to the eleven land cover classes identified as being surrogates for the

entirety of the land cover classes available in Africover. The process of reclassifying the

data used within this thesis project was completed using assumptions based on an

expectation of reasonable reclassifications. For example, the wide variety of agricultural

crops detailed in the Africover land cover dataset were generalized simply as

agriculture. A complete list of the land cover reclassification is provided in Appendix A.

The water data is of particular concern since it has the most influence out of all

of the model criteria. Additionally, Galanti et al. (2006) suggest that elephants have

different movement behaviors in the wet season, ranging from December to May, and

the dry season, ranging from June to November (Voeten and Prins 1999). Galanti et al.

(2006) provide separate values for elephant habitat used in the wet and dry season as

elephants may choose to forage in areas of higher primary productivity that provide

nourishment even in the dry season. The other study used to provide values in the

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model, Ngene et al. (2009), incorporated water into their model using permanent

drinking water points and seasonal rivers as separate measurements of elephant

preference. This thesis combined the two measurements made in Ngene et al. (2009)

and classified them as one measurement of water source. Additionally complicating the

distance to drinking water issue was the use of WWF Hydrosheds data, combined with

lakes data from Tanzania GIS User’s Group. These two data sources do not distinguish

between wet and dry season availability of water. The WWF Hydrosheds data only

provides a representation of the drainage network, rather an actual mapping of

perennial river networks.

These combined issues decrease the credibility of the distance to water criterion

used in the model, especially since it was declared that the model was being sensitized

to respond to the habitat requirements of elephants in the dry season. This uncertainty

should be considered while evaluating the use of the model in conservation planning,

however, the construction of a flawless model was not the intent of this thesis. More

importantly, the issues with water data, as well as its commanding influence on

elephant behavior should provide future GIS researchers with the evidence needed to

focus on the water issue in as great of detail as possible.

The slope data could have been potentially misused during the modeling

process. Values for elephant’s ability to navigate slope came from Wall et al. (2006)

which were combined with the value Ngene et al. (2009) developed for explaining

elephant behavior in regards to elevation. The difference between slope and elevation

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was not distinguished in the thesis model and therefore would need to be evaluated

more robustly in a new modeling scenario.

The roads data could be improved in several different ways. The improvement

status of roads was never distinguished in the model. All roads, whether used

frequently or not, paved or dirt, were counted as having equal influence in the model.

Additionally the roads data was derived as ancillary data delivered in the same database

that the Africover land cover data was delivered in. Although it is not verified, it would

be reasonable to assume that roads may have been constructed since the 2003 release

of this dataset.

5.3 Model processing issues

In the process of constructing the model methodology several issues arose that

deserve mention in this discussion so that new research on this topic may benefit from

the lessons learned. First, was the use of the Summary Statistics tool in ArcGIS Toolbox.

The Summary Statistics tool gives a variety of statistics in raster data, such as minimum

and maximum value, average and range. It was not known at the time of construction

that range was an output of the tool and its use was inadvertently disregarded during

the modeling process which made the modeling process slightly more complicated than

necessary. In the model the total range of values were arranged to subtract the

maximum value from the minimum value to produce a total range value.

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Also of note during the modeling process was the use of magnitudes to derive

cost of movement values that could be processed in the ArcGIS modeling environment.

Section 3.9 describes the process of developing the cost surface used to derive the

CMTC corridors. It was found that the raster file types output from process contained

decimal values and therefore would not process successfully in ArcGIS ModelBuilder. To

remedy the situation the values were multiplied by 1,000,000 to ensure at least six

significant digits. Maintaining this level of detail of each value was not derived from

literature, rather was assumed to be a conservative enough to capture enough available

data values to provide a proper valuation of the cost surface as an elephant may

experience the landscape. This decision created a cost surface with resistance values

ranging from 0 to 1,000,000. When the CMTC analysis was completed corridor values

ranged from zero to the tens of billions.

At this point, it was discovered that values in the tens of billions would also not

process successfully in ModelBuilder. Therefore the values were reduced by 100,000 to

return all values to at least the tens of millions magnitude. A brief review of the data

values indicated that all values could be reduced by five digits without any loss of data.

This review, however, turned out to be flawed. It was discovered later that the lowest

values in the CMTC corridors was actually 613,260,000. Dividing this value by 100,000

results in a value of 6132.6; further, when the Int tool is used to remove the decimal

value and create only integer values this data value is lost. This may have occurred

more than once in the model. A full investigation of data values that were in the

hundreds of millions range and therefore would have lost this unit of value was not

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completed. The results were compromised to a slight degree, but it was deemed

reasonable to accept this small degree of loss. These values were likely especially

prevalent when the distance between origin and destination was minimal. In the vast

majority of cases, there was minimum value lost.

5.4 The exclusion of scale sensitivity

The issue of scale is also of concern for the analysis of the model. Improvements

could be made to a new iteration of this model by taking into account the sensitivities of

the differences in scale used while modeling. It was assumed that scale is a non-factor

throughout the model, as can be evidenced by the use of zones of analysis with

different total areas and the evaluation of connectivity using the same modeling process

and data at the single corridor level all the way up to the region-wide model

5.5 Final thoughts

Given the quantity of problems associated with the data and the flaws in the

modeling process, the question arises whether this thesis research has any value at all.

It is clear that improvements made to the data would certainly increase the credibility of

the modeling exercise. It is also clear that the amount of inputs needed to model

connectivity are abundant and not easily obtained. Therefore, it is hoped that this

discussion may serve as evidence to the need to increase the quality of data in the

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southeast Tanzanian region. Developing data for regional water availability during the

dry season, for instance, might serve as a critical input to a variety of research projects

trying to model wildlife occurrence and movement in the region. As discussed in

Chapter 1, the area is home to a multitude of endemic species and some of the last

remaining vestiges of east African coastal forest. Improving the data availability so that

more research may contribute to conservation in the region would be of immense

value. The added value of a thesis project such as this, is that may serve as a prototype

for conservation planning in one of the biologically outstanding places on planet earth.

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Appendix A: Africover land cover classes reclassified

Africover Land Cover Class Galanti et al. (2008)

Reclassification

1 Forest Plantation - Acacia mearsi 1 Agriculture

2 Forest Plantation - Eucalyptus spp. 1 Agriculture

3 Forest Plantation - Pinus spp. 1 Agriculture

4 Forest Plantation - Teak 1 Agriculture

5 Forest Plantation - Needle Leaved Evergreen 1 Agriculture

6 Forest Plantation - Large Fields 1 Agriculture

7 Rainfed Tree Crop, Large Fields 1 Agriculture

8 Rainfed Tree Crop, Medium Fields 1 Agriculture

9 Rainfed Tree Crop, Plus 1 Herbaceous Crop, Medium Fields 1 Agriculture

10 Rainfed Orchard (1 add. Herbaceous Crop), Medium Fields - Cashew 1 Agriculture

11 Rainfed Tree Crop, Clustered Medium Fields 1 Agriculture

12 Rainfed Tree Crop (1 add. Herbaceous Crop) - Clustered Medium Fields 1 Agriculture

13 Rainfed Tree Crop, Small Fields 1 Agriculture

14 Rainfed Tree Crop (1 add. Herbaceous Crop), Small Fields 1 Agriculture

15 Rainfed Tree Crop (1 add. Shrub Crop), Small Fields 1 Agriculture

16 Rainfed Tree Crop, Clustered Small Fields 1 Agriculture

17 Rainfed Tree Crop (1 add. Herbaceous Crop), Clustered Small Fields 1 Agriculture

18 Rainfed Tree Crop (1 add. Shrubs Crop), Clustered Small Fields 1 Agriculture

19 Rainfed Tree Crop, Isolated Small Fields 1 Agriculture

20 Rainfed Tree Crop (1 add. Herbaceous Crop), Isolated Small Fields 1 Agriculture

21 Rainfed Shrub Crop, Large Fields 1 Agriculture

22 Rainfed Shrub Crop, Large Fields - Tea 1 Agriculture

23 Rainfed Shrub Crop, Medium Fields 1 Agriculture

24 Rainfed Shrub Crop, Medium Fields - Tea 1 Agriculture

25 Rainfed Shrub Crop, Small Fields 1 Agriculture

26 Rainfed Shrub Crop, Clustered Small Fields 1 Agriculture

27 Rainfed Shrub Crop, Isolated Small Fields 1 Agriculture

28 Rainfed Shrub Crop (1 add. Herbaceous Crop), Small Fields 1 Agriculture

29 Rainfed Shrub Crop (1 add. Herbaceous Crop), Clustered Small Fields 1 Agriculture

30 Rainfed Shrub Crop (1 add. Shrub Crop), Small Fields - Coffee, Banana 1 Agriculture

31 Rainfed Shrub Crop (1 add. Herbaceous Crop), Isolated Small Fields 1 Agriculture

32 Irregularly Flooded Cereals, Small Fields - Rice 1 Agriculture

33 Irrigated Herbaceous Crop, Large Fields 1 Agriculture

34 Irrigated Herbaceous Crop, Large Fields - Sugarcane 1 Agriculture

35 Irrigated Herbaceous Crop, Small Fields 1 Agriculture

36 Post Flooding Herbaceous Crop, Medium Fields 1 Agriculture

37 Post Flooding Herbaceous Crop, Medium Fields, Clustered 1 Agriculture

38 Post Flooding Herbaceous Crop, Small Fields 1 Agriculture

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39 Post Flooding Herbaceous Crop, Clustered Small Fields 1 Agriculture

40 Post Flooding Herbaceous Crop, Isoloated Small Fields 1 Agriculture

41 Rainfed Herbaceous Crop, Large to Medium Fields 1 Agriculture

42 Rainfed Herbaceous Crop, Large to Medium Fields - Sisal 1 Agriculture

43 Rainfed Herbaceous Crop, Large Fields 1 Agriculture

44 Rainfed Herbaceous Crop, Large Fields - Sugarcane 1 Agriculture

45 Rainfed Herbaceous Crop, Large Fields - Sisal 1 Agriculture

46 Rainfed Herbaceous Crop, Large Fields - Wheat 1 Agriculture

47 Rainfed Herbaceous Crop, Medium Fields 1 Agriculture

48 Rainfed Herbaceous Crop, Medium Fields - Wheat 1 Agriculture

49 Rainfed Herbaceous Crop, Clustered Medium Fields 1 Agriculture

50 Rainfed Herbaceous Crop, Isolated Medium Fields 1 Agriculture

51 Rainfed Herbaceous crop, Small Fields 1 Agriculture

52 Rainfed Herbaceous Crop, Clustered Small Fields 1 Agriculture

53 Rainfed Herbaceous Crop, Isolated Small Fields 1 Agriculture

54 Rainfed Herbaceous Crop (2 add. Herbaceous Crops), Small Fields 1 Agriculture

55 Rainfed Herbaceous Crop (2 add. Herbaceous Crops), Clustered Small Fields 1 Agriculture

56 Rainfed Herbaceous Crop (2 add. Herbaceous Crops), Isolated Small Fields 1 Agriculture

57 Rainfed Herbaceous Crop (2 add. Herbaceous Crops), Isolated Small Fields 11 Urban

58 Vegetated Urban Areas 2 Closed forest

59 Closed woody (broadleaved deciduous) with sparse 2 Closed forest

60 Open woody with closed to open herbaceous 3 Open forest

61 Closed trees (needlelaved evergreen) 2 Closed forest

62 Closed trees with closed to open shrubs 2 Closed forest

63 Closed low trees with closed to open shrubs 2 Closed forest

64 Open general trees with closed to open shrubs 3 Open forest

65 Open general trees with open shrubs 3 Open forest

66 Open general trees (broadleaved evergreen) with open shrubs 3 Open forest

67 Open trees (broadleaved deciduous) with closed to 3 Open forest

68 Open trees (broadleaved deciduous) with open herbaceous and sparse shrubs 7 Open shrubland

69 Open low trees (broadleaved deciduous) with open herbaceous and sparse shrubs 7 Open shrubland

70 Very open trees (broadleaved deciduous) with 3 Open forest

71 Very open low trees (broadleaved deciduous) with open herbaceous and sparse shrubs 7 Open shrubland

72 Very open low trees (broadleaved deciduous) with open tall herbaceous and sparse shrubs 7 Open shrubland

73 Closed multilayered trees (broadleaved evergreen) 2 Closed forest

74 Closed Shrubs 6 Closed shrubland

75 Closed medium shrubs (broadleaved deciduous) - Fern 6 Closed shrubland

76 Open general shrubs with closed to open herbaceous 7 Open shrubland

77 Open general shrubs with closed to open herbaceous and sparse trees 7 Open shrubland

78 Open shrubs with closed to open herbaceous and 7 Open shrubland

79 Very open shrubs with closed to open herbaceous 7 Open shrubland

80 Very open shrubs with closed to open herbaceous and sparse trees 7 Open shrubland

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81 Sparse shrubs with sparse herbaceous 7 Open shrubland

82 Closed to very open herbaceous 4 Savannah

83 Closed to very open herbaceous with sparse shrubs 5 Savannah with trees and shrubs

84 Closed to very open herbaceous with sparse trees 5 Savannah with trees and shrubs

85 Sparse herbaceous 7 Open shrubland

86 Cereals, Rice - Large Fields 1 Agriculture

87 Cereals, Rice - Medium Fields 1 Agriculture

88 Cereals, Rice - Small Fields 1 Agriculture

89 Closed herbaceous on temporarily flooded and - fresh water 8 Non-permanent water

90 Closed herbaceous with sparse trees on temporarily flooded land - fresh water 8 Non-permanent water

91 Closed Herbaceous (on permanently flooded land - Fresh Water) 8 Non-permanent water

92 Closed to very open herbaceous with sparse shrubs on temporarily flooded land - fresh water 8

Non-permanent water

93 Closed shrubs on temporarily flooded land - fresh water 8 Non-permanent water

94 Closed shrubs (broadleaved evergreen) on permanently flooded land - brackish water 9 Permanent water

95 Open shrubs with closed to open herbaceous on 8 Non-permanent water

96 Very open shrubs with closed to open herbaceous on temporarily flooded land - fresh water 8

Non-permanent water

97 Closed trees on temporarily flooded land - fresh water 8 Non-permanent water

98 Open general woody with closed to open herbaceous on temporarily flooded land 8

Non-permanent water

99 Closed trees (broadleaved evergreen) on permanently flooded land - brackish water 9 Permanent water

100 Open general trees with closed to open herbaceous on temporarily flooded land - fresh water 8

Non-permanent water

101 Urban areas (general) 11 Urban

102 Rural settlements 11 Urban

103 Refugee camp 11 Urban

104 Port 11 Urban

105 Airport 11 Urban

106 Industrial area - general 11 Urban

107 Quarry 11 Urban

108 Bare rock 7 Open shrubland

109 Bare soil 7 Open shrubland

110 Bare soil very stony 7 Open shrubland

111 Salt crusts 7 Open shrubland

112 Sand 10 Waterbodies

113 Artificial Lakes or Reservoirs 10 Waterbodies

114 River 10 Waterbodies

115 Natural Lakes 10 Waterbodies

116 River banks 10 Waterbodies

117 Lake shore 10 Waterbodies

118 Sand beaches 10 Waterbodies

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119 Snow 10 Waterbodies

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Appendix B: Modeled PA connections

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Interzones

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Appendix C: List of corridors between PAs modeled Zone A PA number Origin Protected Area Name

PA number Destination Protected Area Name

1 Wami-Mbiki to 2 Morogoro Fuel

1 Wami-Mbiki to 5 Kitulanghalo

2 Morogoro Fuel to 3 Nguru ya Ndege

2 Morogoro Fuel to 4 Dindili

3 Nguru ya Ndege to 4 Dindili

3 Nguru ya Ndege to 7 Nyandira

3 Nguru ya Ndege to 6 Mindu

4 Dindili to 5 Kitulanghalo

4 Dindili to 12 Mkungwe

4 Dindili to 9 Pangawe East

5 Kitulanghalo to 12 Mkungwe

6 Mindu to 10 Konga

6 Mindu to 28 Bunduki 3

7 Nyandira to 10 Konga

7 Nyandira to 8 Pangawe West

7 Nyandira to 11 Uluguru North

8 Pangawe West to 9 Pangawe East

9 Pangawe East to 11 Uluguru North

9 Pangawe East to 13 Mangala

9 Pangawe East to 15 Ruvu

9 Pangawe East to 12 Mkungwe

11 Uluguru North to 16 Bunduki

11 Uluguru North to 27 Milawilila

12 Mkungwe to 15 Ruvu

12 Mkungwe to 23 Mkulazi

13 Mangala to 14 Kimboza

13 Mangala to 27 Milawilila

14 Kimboza to 15 Ruvu

14 Kimboza to 19 Uluguru South

14 Kimboza to 20 Kasanga

15 Ruvu to 22 Chamanyani

15 Ruvu to 23 Mkulazi

16 Bunduki to 27 Milawilila

16 Bunduki to 28 Bunduki 3

17 Vigoza to 28 Bunduki 3

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17 Vigoza to 19 Uluguru South

17 Vigoza to 18 Nyandiduma and Nyandira

18 Nyandiduma and Nyandira to 19 Uluguru South

18 Nyandiduma and Nyandira to 29 Shikurifumi

19 Uluguru South to 29 Shikurifumi

19 Uluguru South to 20 Kasanga

20 Kasanga to 21 Mvuha

20 Kasanga to 29 Shikurifumi

20 Kasanga to 25 Kilengwe

21 Mvuha to 22 Chamanyani

21 Mvuha to 26 Vigoregoro

23 Mkulazi to 26 Vigoregoro

24 Mikumi to 29 Shikurifumi

24 Mikumi to 25 Kilengwe

25 Kilengwe to 29 Shikurifumi

25 Kilengwe to 26 Vigoregoro

Zone B

PA number Origin Protected Area Name

PA number Destination Protected Area Name

30 Ruvu North Fuel to 31 Pande

30 Ruvu North Fuel to 34 Ruvu South

31 Pande to 35 Pugu

33 Mbinga to 35 Pugu

33 Mbinga to 37 Forest Reserve Name Unknown (TZA) (Mangrove) No.20

34 Ruvu South to 35 Pugu

33 Mbinga to 37 Forest Reserve Name Unknown (TZA) (Mangrove) No.20

34 Ruvu South to 35 Pugu

34 Ruvu South to 38 Masanganya

35 Pugu to 36 Kazimzumbwi

36 Kazimzumbwi to 33 Mbinga

37 Forest Reserve Name Unknown (TZA) (Mangrove) No.20 to 41

Forest Reserve Name Unknown (TZA) (Mangrove) No.22

40 Forest Reserve Name Unknown (TZA) (Mangrove) No.23 to 41

Forest Reserve Name Unknown (TZA) (Mangrove) No.22

40 Forest Reserve Name Unknown (TZA) (Mangrove) No.23 to 43 Marenda

40 Forest Reserve Name Unknown (TZA) (Mangrove) No.23 to 51 Mchungu

39 Masanganya to 43 Marenda

36 Kazimzumbwi to 38 Masanganya

38 Masanganya to 45 Mtanza Msona

39 Masanganya to 42 Mtita

42 Mtita to 43 Marenda

40 Forest Reserve Name Unknown (TZA) to 51 Mchungu

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(Mangrove) No.23

39 Masanganya to 42 Mtita

42 Mtita to 43 Marenda

42 Mtita to 46 Ruhai River

42 Mtita to 47 Ngulakula

47 Ngulakula to 44 Kingoma

52 Forest Reserve Name Unknown (TZA) (Mangrove) No.25 to 33 Mbinga

48 Nyumburuni to 33 Mbinga

44 Kingoma to 48 Nyumburuni

43 Marenda to 44 Kingoma

44 Kingoma to 51 Mchungu

51 Mchungu to 50 Kikale

51 Mchungu to 52 Forest Reserve Name Unknown (TZA) (Mangrove) No.25

58 Nyamwage Village to 48 Nyumburuni

50 Kikale to 58 Nyamwage Village

52 Forest Reserve Name Unknown (TZA) (Mangrove) No.25 to 59 Mohoro

48 Nyumburuni to 58 Nyamwage Village

48 Nyumburuni to 78 Yelya Village

48 Nyumburuni to 57 Katundu

47 Ngulakula to 48 Nyumburuni

46 Ruhai River to 47 Ngulakula

45 Mtanza Msona to 46 Ruhai River

55 Kipo to 47 Ngulakula

45 Mtanza Msona to 55 Kipo

55 Kipo to 57 Katundu

55 Kipo to 56 Rupiange

56 Rupiange to 57 Katundu

56 Rupiange to 61 Ngarambe-Tapika

56 Rupiange to 76 Kichi Hill

57 Katundu to 76 Kichi Hill

57 Katundu to 78 Yelya Village

47 Ngulakula to 57 Katundu

76 Kichi Hill to 72 Mbinga

76 Kichi Hill to 75 Lungonya

61 Ngarambe-Tapika to 75 Lungonya

75 Lungonya to 72 Mbinga

72 Mbinga to 74 Ulabo

73 Mchonga to 74 Ulabo

71 Tong-omba to 73 Mchonga

70 Kibambo to 71 Tong-omba

67 Kilungulungu to 70 Kibambo

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66 Kiwengoma to 67 Kilungulungu

63 Nambunju Village to 66 Kiwengoma

62 Tawi Village to 66 Kiwengoma

62 Tawi Village to 63 Nambunju Village

76 Kichi Hill to 62 Tawi Village

62 Tawi Village to 78 Yelya Village

78 Yelya Village to 77 Mbwara Village

77 Mbwara Village to 64 Namakutwa Nyamulete

64 Namakutwa Nyamulete to 68 Mchekela

68 Mchekela to 69 Kianika

64 Namakutwa Nyamulete to 68 Mchekela

69 Kianika to 79 Rufiji-Mafia-Kilwa Marine

64 Namakutwa Nyamulete to 79 Rufiji-Mafia-Kilwa Marine

64 Namakutwa Nyamulete to 65 Tamburu

58 Nyamwage Village to 59 Mohoro

58 Nyamwage Village to 65 Tamburu

65 Tamburu to 79 Rufiji-Mafia-Kilwa Marine

59 Mohoro to 60 Mohoro River

59 Mohoro to 65 Tamburu

79 Rufiji-Mafia-Kilwa Marine to 81 Forest Reserve Name Unknown (TZA) (Mangrove) No.27

60 Mohoro River to 81 Forest Reserve Name Unknown (TZA) (Mangrove) No.27

81 Forest Reserve Name Unknown (TZA) (Mangrove) No.27 to 33 Mbinga

Zone C

PA number Origin Protected Area Name

PA number Destination Protected Area Name

80 Kisangi Village to 82 Forest Reserve Name Unknown (TZA) (Mangrove) No.28

80 Kisangi Village to 85 Migeregere Village

80 Kisangi Village to 83 Kikole Village

82 Forest Reserve Name Unknown (TZA) (Mangrove) No.28 to 85 Migeregere Village

82 Forest Reserve Name Unknown (TZA) (Mangrove) No.28 to 88

Forest Reserve Name Unknown (TZA) (Mangrove) No.30

85 Migeregere Village to 88 Forest Reserve Name Unknown (TZA) (Mangrove) No.30

85 Migeregere Village to 86 Ruhatwe Village

84 Kikole Village to 86 Ruhatwe Village

86 Ruhatwe Village to 89 Mitundumbea

86 Ruhatwe Village to 87 Mitaurure

88 Forest Reserve Name Unknown (TZA) (Mangrove) No.30 to 89 Mitundumbea

88 Forest Reserve Name Unknown (TZA) (Mangrove) No.30 to 90 Kiwawa Village

87 Mitaurure to 89 Mitundumbea

87 Mitaurure to 91 Rungo

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91 Rungo to 92 Ngarama

90 Kiwawa Village to 92 Ngarama

89 Mitundumbea to 90 Kiwawa Village

92 Ngarama to 94 Pindiro

92 Ngarama to 95 Ndimba

94 Pindiro to 95 Ndimba

93 Kihundu Village to 94 Pindiro

91 Rungo to 93 Kihundu Village

93 Kihundu Village to 109 Malehi

109 Malehi to 108 Matapwa

108 Matapwa to 96 Ruawa Munimburo

95 Ndimba to 96 Ruawa Munimburo

100 Chitoa to 96 Ruawa Munimburo

100 Chitoa to 101 Litipo

107 Ntama to 101 Litipo

106 Nndawa Village to 107 Ntama

105 Rondo to 106 Nndawa Village

105 Rondo to 99 Mkangala

98 Nandimba to 108 Matapwa

98 Nandimba to 99 Mkangala

98 Nandimba to 105 Rondo

98 Nandimba to 105 Rondo

98 Nandimba to 104 Lionja

109 Malehi to 97 Nyera Kiperere

109 Malehi to 98 Nandimba

97 Nyera Kiperere to 103 Angai

103 Angai to 104 Lionja

104 Lionja to 107 Ntama

Interzone A

PA number Origin Protected Area Name

PA number Destination Protected Area Name

1 Wami-Mbiki to 30 Ruvu North Fuel

5 Kitulanghalo to 30 Ruvu North Fuel

12 Mkungwe to 30 Ruvu North Fuel

12 Mkungwe to 34 Ruvu South

23 Mkulazi to 30 Ruvu North Fuel

23 Mkulazi to 34 Ruvu South

23 Mkulazi to 38 Masanganya

23 Mkulazi to 42 Mtita

23 Mkulazi to 45 Mtanza Msona

26 Vigoregoro to 45 Mtanza Msona

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25 Kilengwe to 45 Mtanza Msona

Interzone B

PA number Origin Protected Area Name

PA number Destination Protected Area Name

75 Lungonya to 97 Nyera Kiperere

75 Lungonya to 87 Mitaurure

72 Mbinga to 83 Kikole Village

72 Mbinga to 80 Kisangi Village

74 Ulabo to 80 Kisangi Village

69 Kianika to 85 Migeregere Village

79 Rufiji-Mafia-Kilwa Marine to 82 Forest Reserve Name Unknown (TZA) (Mangrove) No.28

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