Biodiversity and Conservation (2006) 15:855878

DOI 10.1007/s10531-004-2937-4

Springer 2006

-1

A new approach to diversity indices modeling and

mapping plant biodiversity of Nallihan
(A3-Ankara/Turkey) forest ecosystem in frame of
geographic information systems
HAKAN METE DOGAN1,* and MUSA DOGAN2
1
GIS and RS Department, Central Research Institute for Field Crops (CRIFC), Gulderen Sok.
Ziraat Loj. No: 3, Yenimahalle, 06070 Ankara, Turkey; 2Biology Department, Middle East Technical
University, Ankara, Turkey; *Author for correspondence (e-mail: hmdogan@hotmail.com; phone:
+90-312-327-01-50; fax; +90-312-315-14-66)

Received 10 February 2004; accepted in revised form 4 August 2004

Key words: Flora, Geographic information systems, Mapping, Modeling, Plant biodiversity, Plant
ecology, Remote sensing, Spatial analysis
Abstract. Modeling and mapping possibilities of ShannonWiener, Simpson, and number of
species (NS) indices were researched using geographic information systems (GIS) and remote
sensing (RS) tools in Nallihan forest ecosystem of Turkey. The relationships between the indices
and a number of independent variables such as topography, geology, soil, climate, normalized
dierence vegetation index (NDVI), and land cover were investigated to understand relationships
between plant diversity and ecosystem. Georeferenced eld data from the established 56 quadrats
(50 20 m) were used to calculate the indices. Principle component analysis (PCA) and multiple
regression were employed for data reduction and model development, respectively. Three diversity
maps were produced using the developed models. Residual maps and logical interpretations in
ecological point of view were used to test the validity of the models. Elevation and climatic factors
formed the most important components that are eective determinants of plant species diversity,
but geological formations, soil, land cover and land-use characteristics also inuenced plant
diversity. Considering the dierent responses of the models, ShannonWiener (SWI) and NS
models were found suitable for rare cover types, while Simpson (SIMP) model might be appropriate for single dominant land covers in the study area.

Introduction
Conservation Biology is an emerging discipline dedicated to the preservation of
endangered species and habitats. To develop eective protection strategies,
experts need to understand the relationship between species and ecosystem.
Most importantly, they need to decide which areas are the most important to
protect. Consequently, mapping the areas with high plant biodiversity has a
priority for decision-makers. Eective management plans and actions can only
be achieved with this valuable spatial information. Ecologists dene species
diversity on the basis of two factors: species richness and species evenness. The
number of species (NS) in the community is called species richness, while the
relative abundance of species is described as species evenness (Molles 1999).

856
How environmental structure aect species diversity is one of the most fundamental subject of investigation about communities (Barbour et al. 1987;
Molles 1999). Although having been much criticized for the imperfect denition of the concept of diversity and sampling diculties, diversity indices are
still widely used to evaluate, survey, and conserve ecosystems (Pielou 1966;
Barbour et al. 1987; Riitters et al. 1995; Mouillot and Lepretre 1999). The most
popular indices that have been used to quantify landscape composition are
Shannons index, believed to emphasize the richness component of diversity,
and Simpsons index, emphasizing the evenness component (Magurran 1988;
Nagendra 2002). Choosing appropriate methods and tools, it is believed that
these indices have a potential to map diversity. At this point, geographic
information systems (GIS) that has been recently recognized by conservation
biologists can be an appropriate and powerful tool in the spatial analyzes
performed in conservation biology (Kadmon 1997; Dogan 1998; Kress et al.
1998; Lenton et al. 2000; Dogan 2001). The spatial nature of the biological data
lets GIS to develop spatial models of which they might also be used as a
solution for predictive mapping (Franklin 1998; Gottfried et al. 1998). Where
as the monitoring results and mapping of earlier periods are considered as vital
information for such kind of GIS databases. This need is fullled generally by
aero-space remotely sensed data (Fjeldsa et al. 1997) in which at some regions
of the globe the data set can go back to early 1950s via aerial photographs to
recent via high resolution multispectral global coverages for the diversity
studies. Within this frame, the aim of this study is to create a new approach to
the conventional diversity (Shannon-Wiener, Simpson, NS) indices using GIS
and remote sensing (RS) tools. Consequently, in order to reach this goal plant
biodiversity of Nallihan forest ecosystem was modeled and mapped within the
frame of this new approach between the years 2001 and 2002.

Materials and methods

Study area
This study was conducted in Nallihan administrative district of Ankara
province in Turkey. According to the grid system based on two degrees of
latitude and longitude (Davis 19651988); the study area is located in the A3
grid square of Central Anatolia (Figure 1a). This location is within the IranoTuranian phytogeographical region with some Mediterranean penetrations
(Davis 1971), and the records of Turkeys Plant Database (TUBIVES 2003)
pointed out 119 family, 553 genera, and 1350 plant species in this square.
Recently, a new Acantholimon (Plumbaginaceae) species was published from
the area (Dogan and Akaydin 2002). The study area is specically called
Erenler forest region, and covers 327.31 km2 (32731.29 ha) area. The general
topography of the study area is mountainous (Figure 1b). Generally, agricultural lands are concentrated along the river basins, while forests dominate the

857
higher elevations. There are 28 settlements in the study area, and majority of
them are small villages. Major human eects on the forest can be seen in the
area like agriculture and urbanization. Moreover, a considerable part of this
area faces erosion. About 5.6% forest was degraded by natural or anthropogenic causes in the area.
According to Emberger classication system, the climate of the study area
showed Semi-arid Upper Mediterranean Bioclimate characteristics with cold
winters (Akman and Daget 1971; Akman 1999). Basically, four climatic seasons are recognized in the study area. Precipitation is mostly in the form of rain

Figure 1. Physiographic setting (a) and physical geography (b) of the study area (projection
systems of physiographic setting and physical geography maps were dened as geographic with
European datum (spheroid international 1909) and UTM with European datum (zone: 36, spheroid
international 1909), respectively).

858
throughout the year except winters, and total number of snowy days does not
exceed 20 days. The main tree species of the study area are black pine (Pinus
nigra), juniper (Juniperus spp.), red pine (Pinus brutia), and oak (Quercus spp.).
According to the digital forest stand map, the study area can be generalized in
six categories as non forest (28.39%), oak forest (0.47%), erosion/stony
(5.60%), degraded forest (33.85%), Black Pine forest (31.47), and Red Pine
forest (0.21%).

Methods
A owchart of the methodology is given in Figure 2. Digital geological and soil
maps of the study area were obtained from the General Directorate of Mineral
Research and Exploration (MTA) and the General Directorate of Rural Affairs (KHGM), respectively. Topographical and forest stand maps were digitized in UNIX Arc/Info 7.0.4 and PC Arc/Info 3.5 software (ESRI 1994; ESRI
1997). The LANDSAT-TM image, acquired on August 21st 2000, was utilized
to develop land cover and normalized dierence vegetative index (NDVI) maps
in Erdas Imagine 8.5 software (ERDAS 1997). Supervised classication
method (maximum likelihood parametric rule), 4-5-3 band combination, and
statistical ltering (7 7) were used to develop a land cover map. The unsigned
8-bit NDVI model was utilized to establish NDVI classes. Arc/View 3.2 software (ESRI 1996) and Inverse Distance Weighted (IDW) method were employed to produce the interpolated surfaces (grid maps) of climatic
(temperature, precipitation, and potential evapotranspiration (PET)), and
additional soil (K2O, P2O5, organic matter, pH, salt, CaCO3, saturation and
texture) variables. To conduct spatial analysis, all developed maps were converted to grid themes by using 30 30 m grid size in Arc/View 3.2. Universal
Transverse Mercator (UTM) projection system (spheroid international-1909,
datum: European-1950, zone: 36) was applied to all map data.
Georeferenced point data (791 points) were collected to classify LANDSATTM image and to conduct accuracy assessment (Figure 3a). Land-use (FAO
1990) and formation classes (UNESCO 1973) were utilized to identify the main
land-use and vegetation types in the eld. Detailed plant data for diversity
indices were collected from the established quadrats (Figure 3b). The number
of quadrats was determined as 56 considering the quadrat surveys of Magurran
(1981), and the quadrat sites were established according to stratied random
sampling design (McGrew and Monroe 1993). The size of each quadrat was
20 50 m following Grossman et al. (2003). Plant parameters collected from
each quadrat were (1) species component, (2) NS, (3) species cover (%), and (4)
species density (number of plant/m2). From the quadrats, soil samples were
also taken according to the certain soil sampling methods (Atesalp 1976; Ulgen
and Yurtsever 1995). On the map of study area, total 570 points were determined to aggregate climatic data (Figure 3c). The LOCCLIM software
(Grieser 2002) was employed with the digital elevation model (DEM) to

859

Figure 2. The owchart of the methodology (the rounded rectangles indicate the analyses and
processes, rectangles show output products).

calculate the best estimates of focused climatic variables in each determined

point. Climatic variables were investigated in both annual and seasonal
basis. The time period between May and September was taken as seasonal
because of low precipitation and high PET values within this period.
Total 752 plant specimens, pressed and dried following the rules and denitions explained by Davis and Heywood (1965), were identied in the
ANKARA Herbarium of Ankara University. The Davis Flora of Turkey and
the East Aegean Island Vol. 110 (Davis 19651988) were used as the main
reference throughout the herbarium studies. Species diversity indices were
calculatedPfor each quadrat at
P the2 end of this work. The formulas
H =
(pi loge pi) and D =
(pi ) were employed to calculate Shannon
Wiener and Simpson indices, respectively (Barbour et al. 1987; Molles 1999). In
both formulas, pi values indicate the proportional abundance of the ith species
in a quadrat. On the other hand, NS index has no formula, and it was determined by using the total species number in a quadrat.

860

Figure 3. Georeferenced point data for supervised classication (a), established quadrats (b), and
established point data to derive climatic variables (c).

Spatial analyses and model development were conducted in four steps

(Figure 2). KaiserMeyerOlkin (KMO)Bartlett tests were conducted to test
the suitability of the data for factor analysis. Then, principle component

861
analysis (PCA) with varimax rotation was applied for data reduction (SPSS
2001). Multiple regression, regressing a variable on a series of independent
variables (Sokal and Rohlf 1995), was chosen to formulate the relationships.
This was achieved by applying the linear regression with enter method in
SPSS-11 software (SPSS 2001). Applying the models, species richness maps
were produced in Arc/View 3.2. The reliability of the maps was tested by
residual maps and ecological interpretations. Residuals were calculated by
using the observed and computed values of indices in each quadrat, and IDW
method was employed to map them. To evaluate dierent indices in the same
base, interpolating surfaces of the residuals were developed by using standard
deviation values of each index.
Results
Plant species
Total 239 species belonging to 45 families were determined in the study area.
According to Davis (19651988) and the records of Turkeys Plant Database
(TUBIVES 2003); 14 species were detected as endemic for the study area. NS
recognized in each family is stated in Table 1. Leguminosae, Compositae,
Labiatae, Rosaceae, Cruciferae, and Gramineae families have more species
comparing to the others. The full list of identied species was given in
Appendix 1

Table 1. NS recognized in each family.

Family

No. of species Family

No. of species Family

No. of species

Leguminosae
Compositae
Labiatae
Rosaceae
Cruciferae
Graminae
Liliaceae
Boraginaceae
Scrophulariaceae
Caryophyllaceae
Umbelliferae
Campanulaceae
Rubiaceae
Cupressaceae
Plumbaginaceae

37
34
28
15
10
10
9
8
8
7
7
5
5
4
4

3
3
3
3
2
2
2
2
2
2
2
2
2
2
1

1
1
1
1
1
1
1
1
1
1
1
1
1
1
1

Ranunculaceae
Cistaceae
Papaveraceae
Fagaceae
Santalaceae
Illecebraceae
Rhamnaceae
Geraniaceae
Linaceae
Berberidaceae
Cyperaceae
Paeoniaceae
Pinaceae
Valerianaceae
Dipsacaceae

Iridaceae
Acanthaceae
Anacardiaceae
Chenopodiaceae
Convolvulaceae
Coryllaceae
Crassulaceae
Equisetaceae
Euphorbiaceae
Globulariaceae
Guttiferae
Malvaceae
Orchidaceae
Polygalaceae
Urticaceae

Number of determined plant species in this study were given with their families in this table. In this way, overall
results about the recognized species were summarized eciently. Details about the determined species were also
given in Appendix 1.

862
Remote sensing data
Supervised classication obtained 92.16% overall accuracy with a Kappa
coecient of 0.8828, and produced a reliable result. Moreover, NDVI map
delineated the areas where the plants dominated. Therefore, these two map
layers supplied valuable spatial information that can be eective on plant
species diversity. The grid maps of land cover and NDVI were given in Figure 4 with the original LANDSAT-TM (band 3) image.
Data reduction
Initial results of KMO measure of sampling adequacy indicated that factor
analysis would be an appropriate statistics for data reduction. The best solution was found after the second pass with the removal of (1) aspect, (2) slope,
(3) P2O5, (4) K2O, (5) salt, (6) erosion, and (7) seasonal maximum temperature.
After the removal of these seven variables the KMO measure of sampling
adequacy was increased for all indices (Table 2).
Total ve factors were determined according to their Eigenvalues in the
second pass (Table 3). Consequently, stable models were produced. The stability of the models can be seen in the generic dierentiation of the factors and
their responsible variables. For instance, the rst component consists of elevation and climatic variables for all indices that is reasonable because of the
clear relationship between elevation and climatic factors. Similarly, classes
derived from satellite images (NDVI and land cover classes) take part in the
fth component of all indices.
Modeling
The results of linear (multiple) regression were summarized in Table 4. The
Analysis of Variance (ANOVA) showed the acceptability of the models from a
statistical perspective, and the model summary reported the strength of the
relationship between the models and the dependent variables (Table 4). Large
values of the multiple correlation coecient (R) indicated a strong relationship.
Table 2. KaiserMeyerOlkin and Bartletts Test results for the 22 variables in second pass.
Second pass

SWI index

SIMP index

Number of species

KMO measure of sampling adequacy

Bartletts test of sphericity
Approx. v2
df
Signicance

0.808
2502.219
210
0.000

0.811
2454.079
210
0.000

0.808
2426.494
210
0.000

This table summarized the KMO measure of sampling adequacy results after the second pass with
the removal of (1) aspect, (2) slope, (3) P2O5, (4) K2O, (5) salt, (6) erosion, and (7) seasonal
maximum temperature variables. Increasing KMO and Bartletts Test results (0.808 for Shannon
Wiener, 0.811 for Simpson and 0.808 for NS) for the last 22 variables and the low signicance levels
(0.00 for all indices) indicated the suitability of the for data reduction.

863

Figure 4. Original LANDSAT-TM (band 3) image (a), land cover map (b), and NDVI (8 bit)
classes (c) of the study area.

Moreover, the signicance values of the F statistic are less than 0.05 in all models,
which means that the variation explained by the models is not due to chance. The
unstandardized coecients were dened as the coecients of the estimated
regression model, and they were used in the developed models (Table 5).

864
Mapping
The developed grid themes (complementary data set) and map calculator
functions of Arc/View 3.2 were employed throughout the application process
of the three models. With the power of GIS, mathematical operations
were easily conducted on grid themes. Consequently, species diversity maps of
focused indices were developed (Figure 5).

Discussion
The reliability of species diversity maps was questioned in two ways. These
were (1) mapping residuals to predict the locations where the models work
perfectly and (2) logical interpretations in ecological point of view.
Residual from regression is simply the dierence between observed and
computed value (Berry and Marble 1968; McGrew and Monroe 1993), and is a
good indicator to show where the models work perfectly or imperfectly. In
general, low residual values indicate the robust models. Produced residual maps
were given in Figure 5. The percent area covered by each distinct residual class
indicated the credibility of three models. The less predictive areas for three
models covered small percentages (SWI: 7.82%, SIMP: 6.60% and NS: 7.84%),
while the strongly predictive areas contained signicant parts (SWI: 64.85%,
SIMP: 68.12%, and NS: 67.57%). Moderately predictive areas were also
determined as approximately one fourth of the total area (SWI: 27.33%, SIMP:
25.28% and NS: 24.59%) for each index. Considering strongly and moderately
predictive areas together, it seems that each model runs very well in itself.
Overall results of the study indicated that Simpson model worked inversely
comparing the ShannonWiener and NS models (Figure 5). Low Simpson (0
0.42) high ShannonWiener (2.703.59 and 3.60<) and high NS (1726 and
2758) values were detected in higher altitudes where settlements and agricultural lands are rare. On the contrary, high Simpson (0.630.84 and 0.84<),
low ShannonWiener (01.19 and 1.201.89) and low NS (2> and 28) values
were found along the rivers where irrigated agriculture, orchards, and settlements are common. The dierence between the models comes from their inherent characteristics. Basically, ShannonWiener index gives more
importance to the richness component and rare cover types, while Simpson
index weights the evenness component and dominant cover types (Barbour
et al. 1987; McGarigal and Marks 1994; Haines-Young and Chopping 1996;
Molles 1999; Riitters et al. 2000). Moreover, assemblages with identical richness, diering only in evenness, might be ranked oppositely by the Shannon
Wiener and Simpson diversity indices (Nagendra 2002). Therefore, opposite
responses of ShannonWiener and Simpson indices are highly possible
according to assemblages diering in richness as well as evenness (Hurlbert
1971; Peet 1974; Nagendra 2002).

0.97
0.15
0.17
0.28
0.16
0.12
0.20
0.23
0.10
0.26
0.21
0.57
0.97
0.95
0.97
0.97
0.97
0.92
0.82
0.89
0.93

0.12
0.53
0.19
0.06
0.79
0.85
0.06
0.09
0.81
0.06
0.00
0.63
0.06
0.06
0.13
0.07
0.05
0.29
0.29
0.25
0.21

0.13
0.48
0.82
0.85
0.38
0.18
0.02
0.04
0.07
0.01
0.12
0.02
0.16
0.20
0.15
0.14
0.15
0.07
0.01
0.14
0.14

4
0.08
0.14
0.03
0.03
0.10
0.07
0.89
0.82
0.15
0.37
0.26
0.34
0.06
0.04
0.09
0.07
0.07
0.17
0.27
0.17
0.14

5
0.10
0.04
0.23
0.06
0.10
0.04
0.01
0.02
0.02
0.75
0.85
0.10
0.11
0.12
0.10
0.10
0.11
0.08
0.05
0.16
0.14
0.97
0.15
0.17
0.28
0.17
0.13
0.19
0.22
0.10
0.26
0.21
0.48
0.97
0.95
0.97
0.97
0.97
0.92
0.81
0.89
0.93

2
0.11
0.53
0.18
0.06
0.80
0.85
0.08
0.07
0.80
0.06
0.01
0.50
0.05
0.06
0.13
0.06
0.04
0.28
0.29
0.25
0.20

3
0.09
0.15
0.03
0.04
0.10
0.07
0.88
0.81
0.17
0.36
0.26
0.46
0.07
0.05
0.10
0.08
0.08
0.18
0.29
0.18
0.15

4
0.14
0.58
0.82
0.83
0.39
0.18
0.03
0.03
0.06
0.01
0.13
0.07
0.16
0.20
0.15
0.14
0.15
0.07
0.01
0.14
0.15

Components for Simpson index

Components for ShannonWiener index

5
0.10
0.05
0.23
0.05
0.10
0.04
0.01
0.03
0.03
0.75
0.85
0.10
0.10
0.12
0.10
0.10
0.11
0.08
0.05
0.16
0.14

1
0.97
0.15
0.19
0.29
0.17
0.12
0.20
0.22
0.10
0.26
0.22
0.22
0.97
0.96
0.97
0.98
0.98
0.92
0.82
0.89
0.94

2
0.12
0.56
0.17
0.05
0.77
0.83
0.05
0.10
0.78
0.08
0.04
0.54
0.06
0.05
0.13
0.07
0.05
0.28
0.27
0.23
0.20

3
0.12
0.45
0.82
0.83
0.42
0.21
0.03
0.04
0.03
0.03
0.15
0.24
0.14
0.18
0.14
0.12
0.13
0.07
0.03
0.16
0.15

4
0.08
0.13
0.02
0.02
0.11
0.08
0.89
0.82
0.13
0.35
0.25
0.34
0.07
0.05
0.09
0.07
0.08
0.17
0.26
0.16
0.13

Components for number of species

5
0.10
0.03
0.22
0.07
0.12
0.01
0.01
0.01
0.02
0.75
0.85
0.04
0.10
0.12
0.10
0.10
0.10
0.08
0.05
0.16
0.13

Rotated component matrices of the three indices in the second pass were summarized in this table. Bigger Eigenvalues (over 0.70) of the components were stated in bold indicating
that they were selected for multiple regression. The stability of the model can be seen in the generic dierentiation of the factors and their responsible variables. For instance, the rst
component consists of elevation and climatic variables, and the second component is composed of some soil variables (organic matter, saturation, and texture) and geology for all
indices.
Abbreviations: ELEV, elevation; ORGM, organic matter; STR, saturation; TEXTR, texture; SOILG, big soil group; SLDPT, soil depth; GEO, geological formations; SPVSD,
supervised classes; INDEXES, Shannon Wiener, Simpson, NS indices; META, annual mean temperature; METS, seasonal mean temperature; MAXTA, annual maximum
temperature; MAXTS, seasonal maximum temperature; MINTA, annual minimum temperature; MINTS, seasonal minimum temperature; PRCPA, annual precipitation; PRCPS,
seasonal precipitation; PETAN, annual potential evapotranspiration; PETSE, seasonal potential evapotranspiration.

ELEV
ORGM
CaCO3
PH
STR
TEXTR
SOILG
SLDPT
GEO
NDVI
SPVSD
INDEXES
META
METS
MAXTA
MINTA
MINTS
PRCPA
PRCPS
PETAN
PETSE

Variable

Table 3. Rotated component matrix of ShannonWiener, Simpson, and number of species indices (Eigenvalues of selected variables were written in bold).

865

Simpson

ShannonWiener

R2
0.907

R
0.952a

R2
0.742

R
0.861a

Model 1

0.133
0.027

Mean square

ANOVA

0.854

Adjusted R2

0.594

Adjusted R2

Model summaryb

2.669
0.929
3.598

Regression
Residual
Total

20
35
55

Sum of squares

Model 1

df

1.459
0.086

Mean square

ANOVAb

Model summaryb

Model 1

20
35
55

29.173
2.993
32.167

Regression
Residual
Total

df

Sum of squares

Model 1

Table 4. The statistics of linear (multiple) regression.

0.162938

Standard error

5.027

0.292452

Standard error

17.055

0.000a

Signicance

0.000a

Signicance

866

Model 1

59.935
29.584

Mean square

0.537

R2

Model summaryb

0.732

20
35
55

1198.695
1035.430
2234.125

Regression
Residual
Total

df

Sum of squares

Model 1

ANOVAb

0.272

Adjusted R2

2.026

5.439

Standard error

0.033

Signicance

The results of linear (multiple) regression produced for each index were unied in this table. The analysis of variance (ANOVA) showed the accepttability of
the models from a statistical perspective, and the model summary reported the strength of the relationship between the models and the dependent variables.
Large values of the multiple correlation coecient (R) indicated a strong relationship. Moreover, the signicance values of the F statistic are less than 0.05 in
all models, which means that the variation explained by the models is not due to chance.
Note: For abbreviations see Table 3.
a
Predictors: (Constant), SPVSD, GEO, PH, SLDPT, ORGM, NDVI, MINTA, TEXTR, SOILG, CACO3, PRCPS, STR, PETAN, METS,
PRCPA, MAXTA, MINTS, ELEV, PETSE, META.
b
Dependent variable: SWI.
c
Dependent variable: Simpson.
d
Predictors: (Constant), PETSE, ORGM, NDVI, GEO, SLDPT, PH, SPVSD, TEXTR, SOILG, CACO3, PRCPS, STR, METS,PRCPA,
MAXTA, MINTA, PETAN, MINTS, ELEV, META.
e
Dependent variable: Number of species.

Number of SP.

867

868

Figure 5. Plant species diversity and residual maps of ShannonWiener (a), Simpson (b), and NS
(c) models.

The relationships between the indices and elevation can be recognized when
the elevation (Figure 1) and diversity maps (Figure 5) were examined together.
A direct relationship between the elevation and indices was detected for
ShannonWiener and NS models. On the other hand, this relationship turned
an inverse character in Simpson model. Depending on these results, an
important question arises: which model has the capacity to delineate real situation in the eld? In basic, there are two general concepts: (1) a monotonic
decrease in species richness with increasing elevation (Stevens 1992; Huston
1994; Rahbek 1995; Brown and Lomolino 1998) and (2) a peak in richness at
intermediate elevations (8001400 m) exemplied by a hum-shaped distribution (McCoy 1990; Rahbek 1997; Fleishman et al. 1998). Considering the
elevation range (1441740 m) of the study area, Simpson model might be
found reasonable within the rst concept. On the other hand, ShannonWiener

This table states the models (regression equations) according to the results of multiple regression. In the equations, the understandardized coecients are the
coecients of the estimated regression model. Each model also has a constant value such as; 22.296 for ShannonWiener, 10.424 for Simpson, and 244.804 for
number of species.
Note: For abbreviations see Table 3.

Number of species index = 244.804 + (0.136 * ELEV) + (0.340 * ORGM) + (0.175 * CACO3) + (4.935 * TEXTR) + (11.565 * SOILG) + (1.099 * GEO) +
(0.026 * NDVI) + (0.365 * SPVSD) + (23.089 * META) + (4.766 * METS) + (3.804 * MAXTA) + (6.697 * MINTA) + (1.152 * PRCPS) + (3.932 * PETSE)
(5.642 * PH) (0.395 * STR) (0.402 * SLDPT) (2.079 * MINTS) (0.204 * PRCPA) (10.770 * PETAN)

Simpson index = 10.424 + (0.437 * MINTA) + (0.087 * PRCPS) + (0.028 * PETSE) + (0.079 * TEXTR) + (0.004 * ORGM) + (0.030 * SLDPT) + (0.269 *
PH) + (0.001 * NDVI) (0.002 * ELEV) (0.009 * META) (0.220 * METS) (0.293 * MAXTA) (0.428 * MINTS) (0.064 * PRCPA) (0.027 *
PETAN) (0.003 * STR) (0.064 * GEO) (0.001 * SOILG) (0.011 * CACO3) (0.053 * SPVSD)

ShannonWiener index = 22.296 + (0.008 * ELEV) + (1.688 * META) + (0.944 * MINTS) + (0.097 * PRCPA) + (0.226 * GEO) + (0.071 * ORGM) +
(0.020 * CACO3) + (0.005 * SOILG) + (0.139 * SPVSD) (0.981* METS) (1.068 * MAXTA) (0.289 * MINTA) (0.143 * PRCPS) (0.168 * PETAN)
(0.032 * PETSE) (0.004 * STR) (0.008 * TEXTR) (0.378 * PH) (0.081 * SLDPT) (0.002 * NDVI)

Models

Table 5. Developed models of each index (ShannonWiener, Simpson, and number of species).

869

870
and NS models could be found acceptable according to the second concept. So,
the question remains as to which diversity index reected the reality. According
to The Ecological Society of America Committee on Land Use (Dale et al.
2000); Shannons index of diversity has greater sensitivity to rare cover types
and it needs to be given greater importance during interpretation. However,
Simpsons index of diversity might be preferred in landscapes where a single
dominant land cover type is of interest. Therefore, the appropriateness of the
models depends on the aims what the decision-makers seek. ShannonWiener
and NS Models might be useful to detect the areas where rare and endangered
species in focus. On the other hand, Simpson model could be best t to
determine the areas where dominant species in point of concentration.

Conclusion
In this study, we tested the modeling and mapping capabilities of some diversity
indices by using a new approach. The forest ecosystem was handled as a whole,
and the relationship between the plant biodiversity and the factors eective on
ecosystem were investigated. The complementary data about topography,
geology, soil, forest, climate, land cover, and NDVI supplied very important
information, and played the backbone role at the spatial analysis and modeling
stages. The importance of quantitative eld data was also emphasized. The
results showed that plant diversity can be modeled by using index values and
complementary data set. Both GIS and RS are important tools at the analysis
and visualizing (mapping) stages. According to the results; both Shannon
Wiener and NS models could be successful to reveal the richness aspect of
species diversity, while Simpson model might be acceptable to delineate the
evenness aspect indicating single dominant land cover types. Although this
study suggested an applicable method, it is implied that researchers should be
cautious to select appropriate index according to their aims.

Acknowledgements
The authors wish to thank the following individuals for their contributions in
various parts of this research study: Vedat Toprak, Lut Suzen, Unal Sorman,
and Zuhal Akyurek from Middle East Technical University (METU); Osman
Ketenoglu from Ankara University (AU); Ali Mermer, Ediz Unal, Tuncay
Porsuk, Oztekin Urla, and Hakan Yildiz from the GIS and RS Department of
Central Research Institute for Field Crops (CRIFC-GIS and RS); Murat
Cetiner and Irfan Artuc from the Nallihan Forest Management District
(NFMD). Thanks are also due to METU Research Fund for making nancial
assistance, Soil and Fertilizer Research Institute for analyzing soil samples, and
the Keeper of the Ankara (ANK) Herbarium for making the herbarium
facilities available.

871
Appendix 1 List of identied species in the study area (endemics were stated in bold and marked
with are asterisk (*))
No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46

Species name

Family

Alhagi pseudalhagi (Bieb.) Desv.

Anthyllis vulneraria L. subsp. boissieri (Sag.) Bornm.
Astragalus angustifolius Lam. subsp. angustifolius
Astragalus densifolius Lam.
Astragalus glycyphyllos L. subsp. glycyphylloides (DC.)
Matthews
Astragalus lycius Boiss.
Astragalus macrocephalus Willd. subsp. Macrocephalus
Astragalus microcephalus Willd.
Astragalus micropterus Fischer
Astragalus squalidus Boiss. & Noe *
Astragalus trichostigma Bunge *
Chamaecytisus pygmaeus (Willd.) Rothm.
Cicer pinnatidum Jaub. & Spach
Conorilla varia L. subsp. Varia
Dorycnium pentaphyllum Scop. subsp. anatolicum
(Boiss.) Gams
Hedysarum varium Willd.
Lathyrus aureus (Stev.) Brandza
Lotus aegaeus (Gris.) Boiss.
Lotus corniculatus L. var. corniculatus
Lotus corniculatus L. var. tenuifolius L.
Medicago polymorpha L. var. vulgaris (Benth.) Shinners
Medicago sativa L. subsp. Sativa
Medicago varia Martyn
Melilotus alba Desr.
Melilotus ocinalis (L.) Desr.
Onobrychis argyrea Boiss. Subsp. argyrea
Onobrychis armena Boiss. & Huet.
Onobrychis hypargyrea Boiss.
Ononis adenotricha Boiss. var. adenotricha
Ononis spinosa L. subsp. Leiosperma (Boiss.) Sirj.
Pisum sativum L. subsp. Elatius var. elatius
Trifolium arvense L. var. arvense
Trifolium barbulatum (Freyn & Sint.) Zoh.*
Trifolium repens L. var. repens
Vicia cracca L. subsp. Stenophylla Vel.
Vicia grandiora Scop. var. grandiora
Vicia narborensis L. var. narborensis
Achillea biebersteinii Afan.
Achillea setacea Waldst. & Kit.
Acroptilon repens (L.) DC.
Anthemis tinctoria L. var. discoidea (All.) DC.
Cardopodium corymbosum (L.) Pers.
Carlina corymbosa L.
Centaurea deprassa Bieb.
Centaurea solstitialis L. subsp. solstitialis
Centaurea triumfettii All.

Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Leguminosae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae

872
Appendix 1. (Continued)
No.

Species name

Family

47
48
49
50
51
52
53
54
55
56
57
58
59
60
61

Centaurea urvillei DC. subsp. Urvillei*

Centaurea virgata Lam.
Chardinia orientalis (L.) O. Kuntze
Chondrilla juncea L. var. juncea
Cichorium intybus L.
Cirsium arvense (L.) Scop. subsp. vestitum
Cirsium hypoleucum DC.
Crepis sancta (L.) Babcock
Doronicum orientale Hom.
Echinops ritro L.
Inula oculus-christi L.
Lactuca serriola L.
Leontodon asperrimus (Willd.) J. Ball.
Petasites hybridus (L.) Gaertner
Pilosella echioides (Lumn.) C.H. & F.W.Schultz subsp.
procera (Fries) Sell & West
Pilosella hoppeana (Schultes) C. H. & F.W. Schultz
subsp. testimonialis (NP.) Sell &West
Scorzonera cana (C.A.Meyer) Hom.
Scorzonera laciniata L.
Senecio vernalis Waldst. & Kit.
Sonchus asper L. Hill subsp. glaucescens (Jordan) Ball.
Tanacetum poteriifolium (Ledeb.)
Tanacetum vulgare L.
Taraxacum seronitum (Waldst. & Kit.) Poiret in Lam.
Tragopogon latifolius Boiss. var. angustifolius Boiss.
Xeranthemum annuum L.
Acinos rotundifolius Pers.
Ajuga chamaepitys (L.) Schreber, subsp. chia (Schreber)
Arcangeli, var. chia
Lamium macradon Boiss. & Huet
Marrubium parviorum Fisch. & Mey. subsp. oligodon
(Boiss.) Seybold *
Mentha spicata L. subsp. tomentosa (Briq.) Harley
Nepeta nuda L. subsp. albiora (Boiss.) Gams
Phlomis armeniaca Willd. *
Phlomis nissolii L.
Prunella vulgaris L.
Salvia aethiopis L.
Salvia hypargeia Fisch. & Mey.
Salvia sclarea L.
Salvia tomentosa Miller (Syn: S. grandiora Etl.)
Salvia verticillata L. subsp. amasiaca (Freyn & Bornm.)
Bornm.
Salvia viridis L.
Scutellaria orientalis L. subsp. macrostegia (Hausskn. ex
Bornm.) Edmondson
Sideriris montana L. subsp. montana
Sideritis galatica Bornm.

Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae

62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89

Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Compositae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae

873
Appendix 1. (Continued)
No.
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137

Species name

Family

Stachys annua (L.) L. subsp. ammophila (Boiss. & Bl.)

Samuelss
Stachys annua (L.) L. subsp. a nnua var. annua *
Stachys cretica L. subsp. anatolica Rech. l. *
Teucrium chamaedrys L. subsp. chamaedrys
Teucrium parviorum Schreber
Teucrium polium L.
Thymus leucostomus Hausskn. &Velen. var. leucostomus
Thymus longicaulis C. Presl subsp. longicaulis
Thymus sipyleus Boiss. subsp. sipyleus
Ziziphora capitata L.
Cotoneaster nummularia Fisch. & Mey.
Crataegus monogyna Jacq. subsp. monogyna
Crataegus orientalis Pallas ex Bieb. var. orientalis
Crataegus tanacetifolia (Lam.) Pers. *
Potentilla recta L.
Prunus avium (L.) L.
Prunus divaricata Ledeb. subsp. divaricata
Prunus spinosa L. subsp. dasyphylla (Schur) Domin
Pyracantha coccinea Roemer
Pyrus elaeagnifolia Pallas subsp. elaeagnifolia
Rosa canina L.
Rubus ideaus L.
Rubus sanctus Schreber
Sanguisorba minor Scop. subsp. muricata (Spach) Briq.
Sorbus umbellata (Desf.) Fritsch var. umbellata
Alyssum desertorum Stapf. var. desertorum
Alyssum murale Waldst. & Kit. var. murale
Alyssum sibiricum Willd.
Arabis nova Vill.
Barbera plantaginea DC.
Cardaria draba (L.) Desv. subsp. draba
Erysimum crassipes Fisch. & Mey.
Iberis taurica DC.
Thlaspi perfoliatum L.
Turritis glabra L.
Agropyron cristatum (L.) Geartner, subsp: pectinatum
(Bieb.) Tzvelev, var: pectinatum
Aegilops umbellulata Zhuk.
Brachypodium sylvaticum (Hudson) P. Beauv
Dactylis glomerata L. subsp. glomerata
Festuca airoides Lam.
Festuca anatolica Markgr.-Dannenb. subsp. anatolica
Festuca ilgazensis Markgr.-Dannenb.
Poa bulbosa L.
Stipa bromoides (L.) Dorer
Stipa lessingiana Trin. & Rupr.
Allium scorodoprasum L. subsp. rotundum (L.) Stearn
Gagea granatellii (Parl.) Parl.
Muscari armeniacum Leichtlin ex Baker

Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Labiatae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Cruciferae
Cruciferae
Cruciferae
Cruciferae
Cruciferae
Cruciferae
Cruciferae
Cruciferae
Cruciferae
Cruciferae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Liliaceae
Liliaceae
Liliaceae

874
Appendix 1. (Continued)
No.

Species name

Family

138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162

Muscari longipes Boiss.

Muscari neglectum Guss.
Muscari tenuiorum Tausch
Ornithogalum oligophyllum E.D.Clarke.
Ornithogalum mbriatum Willd.
Ornithogalum umbellatum L.
Adonis ammea Jacq.
Ranunculus argyreus Boiss.
Ranunculus caria L. subs. cariiformis Rouy & Fouc.
Dianthus anatolicus Boiss.
Dianthus ancyrensis Hausskn. & Bornm. *
Dianthus zonatus Fenzl var. zonatus
Herniaria glabra L.
Minuartia hirsuta (Bieb.) Hand. & Mazz.
Saponaria glutinosa Bieb.
Silene supina Bieb. subsp. pruinosa (Boiss) Chowdh
Astrodaucus orientalis (L.) Drude
Coriandrum sativum L.
Falcaria vulgaris Bernh.
Laser trilobum (L.) Borkh.
Malabaila secacul Banks & Sol.
Turgenia latifolia L. Hom.
Zosima absinthifolia (Vent.) Link
Alkanna orientalis (L.) Boiss. var. orientalis
Anchusa leptophylla Roemer & Schultes subsp. leptophylla
Cerinthe minor L. subsp. minor
Lithospermum ocinale L.
Onosma aucheranum DC.
Onosma bornmuelleri Hausskn.
Onosma isauricum Boiss. & Heldr. *
Onosma tauricum Pallas ex Willd. var. tauricum
Digitalis ferruginea L. subsp. ferruginea
Digitalis orientalis Lam.
Scrophularia scopolii [Hoppe ex] Pers. var. scopolii
Verbascum cherianthifolium Boiss var. cheiranthifolium.*
Verbascum glomeratum Boiss
Veronica chamaedrys L.
Veronica multida L.
Veronica pectinata L. var. pectinata
Asyneuma limonifolium (L.) Janchen subsp. pestalozzae
(Boiss.) Damboldt.
Asyneuma rigidum (Willd.) Grossh. subsp. rigidum
Campanula glomerata L.
Campanula persicifolia L.
Legousia speculum-veneris (L.) Chaix
Asperula stricta Boiss. subsp. latibracteata (Boiss.) Ehrend.
Cruciata taurica (Pallas ex Willd.) Ehrend.
Galium incanum Sm. subsp. elatius (Boiss.) Ehrend.

Liliaceae
Liliaceae
Liliaceae
Liliaceae
Liliaceae
Liliaceae
Ranunculaceae
Ranunculaceae
Ranunculaceae
Caryophyllaceae
Caryophyllaceae
Caryophyllaceae
Caryophyllaceae
Caryophyllaceae
Caryophyllaceae
Caryophyllaceae
Umbelliferae
Umbelliferae
Umbelliferae
Umbelliferae
Umbelliferae
Umbelliferae
Umbelliferae
Boraginaceae
Boraginaceae

163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184

Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Scrophulariaceae
Campanulaceae
Campanulaceae
Campanulaceae
Campanulaceae
Campanulaceae
Rubiaceae
Rubiaceae
Rubiaceae

875
Appendix 1. (Continued)
No.

Species name

Family

185
186
187
188
189

Galium palustre L.
Galium verum subsp. verum
Cistus laurifolius L.
Fumana aciphylla Boiss.
Helianthemum nummularium (L.) Miller. subsp. ovatum
(Viv.) Schinz & Thell
Juniperus communis L. subsp. nana
Juniperus excelsa Bieb.
Juniperus foetidissima Willd.
Juniperus oxycedrus L. subsp. oxycedrus
Scabiosa argentea L.
Quercus cerris L. var. cerris
Quercus pubescens Willd.
Quercus robur L. subsp. robur
Osyris alba L.
Thesium billardieri Boiss
Paronychia dudleyi Chaudhri
Paronychia kurdica Boiss. subsp. kurdica var. kurdica
Iris orientalis Miller.
Acanthus hirsutus Boiss.
Paliurus spina-christi Miller
Rhamnus thymifolius Bornm. *
Geranium robertianum L.
Geranium tuberosum L. subsp. tuberosum
Linum hirsitum L. subsp. anatolicum (Boiss) Hayek *
Linum tenuifolium L.
Fumaria cilicica Hausskn.
Hypecoum procumbens L.
Papaver commutatum Fisch & Mey
Rhus coriaria L.
Berberis crataegina DC.
Berberis vulgaris L.
Salsola ruthenica Iljin
Convolvulus arvensis L.
Corylus avellana L. var. avellana
Sempervivum armenum Boiss. & Huet. var. armenum
Carex acca Schreber subsp. serrulata (Biv.) Greuter
Carex ovalis Good.
Equisetum palustre L.
Euphorbia macroclada Boiss.
Globularia trichosanta Fisch. & Mey.
Hypericum perforatum L.
Malva neglecta Wallr.
Cephalanthera rubra (L.) L.C.M. Richard
Paeonia mascula subsp. mascula
Paeonia peregrina
Pinus brutia
Pinus nigra subsp. pallasiana
Acantholimon acerosum (Willd.) Boiss
Acantholimon glumaceum (Jaub. & Spach) Boiss.

Rubiaceae
Rubiaceae
Cistaceae
Cistaceae
Cistaceae

190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233

Cupressaceae
Cupressaceae
Cupressaceae
Cupressaceae
Dipsacaceae
Fagaceae
Fagaceae
Fagaceae
Santalaceae
Santalaceae
Illecebraceae
Illecebraceae
Iridaceae
Acanthaceae
Rhamnaceae
Rhamnaceae
Geraniaceae
Geraniaceae
Linaceae
Linaceae
Papaveraceae
Papaveraceae
Papaveraceae
Anacardiaceae
Berberidaceae
Berberidaceae
Chenopodiaceae
Convolvulaceae
Coryllaceae
Crassulaceae
Cyperaceae
Cyperaceae
Equisetaceae
Euphorbiaceae
Globulariaceae
Guttiferae
Malvaceae
Orchidaceae
Paeoniaceae
Paeoniaceae
Pinaceae
Pinaceae
Plumbaginaceae
Plumbaginaceae

876
Appendix 1 Continued
No.

Species name

Family

234
235
236
237
238
239

Acantholimon reexifolium Bokhari

Plumbago europaea L.
Polygala anatolica Boiss. & Heldr.
Urtica dioica L.
Valeriana alliariifolia Adams
Valerianella vesicaria (L.) Moench

Plumbaginaceae
Plumbaginaceae
Polygalaceae
Urticaceae
Valerianaceae
Valerianaceae

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