Rend. Fis. Acc.

Lincei (2013) 24:349–367

DOI 10.1007/s12210-013-0246-0

Vegetation of Thumamah Nature Park: a managed arid land site

in Saudi Arabia
Mohamed A. El-Sheikh • Jacob Thomas • Abdulrahman A. Alatar •
Ahmed K. Hegazy • Ghanim A. Abbady • Ahmed H. Alfarhan • Mohamed I. Okla

Received: 25 April 2013 / Accepted: 27 June 2013 / Published online: 17 July 2013
 Accademia Nazionale dei Lincei 2013

Abstract Thumamah Nature Park is located at about group III with Acacia gerrardii–Panicum turgidum, group
100 km north of Arriyadh (Saudi Arabia), having an area IV with Panicum turgidum and group V with Acacia eh-
of 170 km2. The Park was established since 30 years ago. renbergiana–Lasiurus scindicus. Vegetation in the upland
The aim of this study is to analyze the vegetation structure plateau was represented by the remaining two groups;
in relation to the environmental factors in different habitat group VI with Helianthemum lippii and group VII with
types. The phenological activities around the year of the 20 Helianthemum kahiricum. The environmental variables
dominant species were monitored. 119 species were iden- that affect the species distribution and diversity in the park
tified, of which 51 (43 %) annuals and 68 (57 %) peren- include the altitude, soil texture, pH, EC, Ca, Mg and Mn.
nials after 30 years of exclusive human impact. The The increased plant species richness, turnover, evenness
Saharo-Arabian component species were the highest and cover were mostly due to the increase of the herba-
among the monoregional species (64 %) in most life forms, ceous species. Plant populations showed interspecific
while the Sahelien-Somali Masai attained the highest variations in their relative timing of phenological phases
among the biregionals (46 %). The TWINSPAN, DCA and with reproductive activity period ranged between 3 and
CCA analyses separated seven vegetation groups. The first 6 months with unimodal flowering peak. Three flowering-
two groups were dominated by psammophytic species, fruiting activities were recorded during late winter–spring,
which occupy the lower sandy plain as shown in group I summer and late autumn–early winter. In an attempt to
with Rhanterium epapposum–Rhazya stricta and group II explain the vegetation dynamics after 30 years conserva-
with Pennisetum divisum–Haloxylon salicornicum. The tion, the progressive succession varied among the different
escarpment habitat was characterized by three groups, viz., habitat types, including the lower sandy sites, the escarp-
ment and the upland rocky habitats, which reflect the
relationship between altitude, edaphic factors and the type
M. A. El-Sheikh  J. Thomas  A. A. Alatar  of vegetation units in each habitat type after exclusion of
A. K. Hegazy  A. H. Alfarhan  M. I. Okla the human impact.
Botany and Microbiology Department, College of Science, King
Saud University, P.O. Box 22455, Riyadh 11451, Saudi Arabia
Keywords Keystone species  Landscape  Najd Plains 
M. A. El-Sheikh (&) Psammophytic species  Succession
Botany Department, Faculty of Science, Damanhour University,
Damanhour, Egypt
e-mail: el_sheikh_eg@yahoo.co.uk
1 Introduction
A. K. Hegazy
Department of Botany, Faculty of Science, Cairo University, Spatial variations in the vegetation composition of the
Giza, Egypt desert and the composition of life forms reflect the
response of vegetation to certain environmental factors
G. A. Abbady
Biological Science Department, Faculty of Science, Kuwait such as topography, climate and human impact (Zohary
University, P.O. Box 5969, Safat 13060, Kuwait 1973; El-Demerdash et al. 1995; Shaltout and Mady 1996;

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350 Rend. Fis. Acc. Lincei (2013) 24:349–367

Hosni and Hegazy 1996; Hegazy et al. 1998; Hutchinson recreation activities and conservation of wild plant
et al. 2000; Alday et al. 2010; Kürschner and Neef 2011; diversity against human impacts. Thumamah Nature Park
El-sheikh et al. 2012; Alatar et al. 2012). The relationship (Saudi Arabia) had been designed in 1983, owing to the
between vegetation dynamics and disturbance has long unique and variable habitat types including escarpments,
been a central focus in ecology. In desert ecosystems, cli- foothills, wadi systems, rock and gravel deserts, and
mate fluctuates markedly and water is the dominant lim- sandy plains. The diversity and complexity of the flora
iting factor that controls the plant growth. Nevertheless, and vegetation are exceptionally high. Owing to the
like many other vegetation types, the dynamics of desert management of the area since 30 years ago this mini-
perennial vegetation can be affected by site history, cli- mized the recreational and grazing impacts. Thumamah
matic factors, life history of the dominant species and Nature Park has unique macro and micro-habitats vary-
recovery after disturbances (Brown et al. 1997; Qinfeng ing from sandy plain lowland to escarpment and upper
2004). Most early studies in arid ecosystems addressed land plateau. This variation and heterogeneity in the
climatic effects on annuals rather than perennial plants, habitats increase the species diversity (Abbady and El-
mainly because the latter required longer observations. The Sheikh 2002; El-Sheikh et al. 2010). Therefore, land-
greater longevity of many perennials, especially woody scape designers’ were taken into their consideration in
species, may lead to greater time lags in their response to the conservation of wild plant diversity and created the
disturbances or climatic changes than observed for annuals camp in the empty areas far away from the major plant
(e.g., Cody 2000; Valone et al. 2002; Qinfeng 2004). associations.
Desert ecosystems are fragile ecosystems and nearly Hitherto, there are no thorough vegetation studies on
always regeneration is slow, since primary production is Thumamah Nature Park except the first unpublished
low (Batanouny 1983; Qinfeng 2004). report (Baierle and Kürschner 1985) describing the veg-
Arabian Desert is subjected to major environmental etation in the early stages of the Park establishment. The
disturbances that often leave the landscape with no vege- objectives of this study are to: (1) study the vegetation
tation and poor soil-forming material on which an eco- structure, in relation to the prevailing environmental
system can develop. In recent years, Saudi Arabia has variables affecting the floristic composition and species
suffered from the effects of desertification, mainly due to diversity in the landscape habitat types after 30 years
climatic fluctuation resulting in drought, soil erosion and protection, (2) monitor the phenology of the keystone
inappropriate land practices that disturb the ecological species dominating the habitat types in the Park, and (3)
balance of ecosystems. In many cases, the human impacts propose and explain the potential progressive succession
and the ill-advised land-use are the main reasons in the in different habitat types in response to the park conser-
deterioration of many ecosystem types (Hajara and Bata- vation management.
nouny 1977; El-Sheikh et al. 2010). In desert habitats of
central Saudi Arabia, water and temperature are the limit-
ing ecological factor for vegetation establishment. Sporadic 2 Materials and methods
rainfall occurs in low amounts, often at irregular intervals
and with large inter-annual variability (Batanouny 1973; 2.1 Study site
Springuel et al. 2006). The desert vegetation in Najd region
is found mainly in runnels, wadis, rangelands, sandy plains Thumamah Park is a unique natural ecosystem and popular
and raudhas (depressions), that possess variations of spe- entertainment center due to its scenic surroundings and
cies abundance and diversity (Kürschner and Neef 2011; proximity to the Arriyadh City. It is about 100 km from the
Alatar et al. 2012). Vegetation is generally of xeromorphic central Arriyadh, covering an area of approximately
type and consists of an approximately continuous herb- 170 km2. The study area is located between 25140 02.5300
layer, which is dominated by annual and perennial grasses, N–46370 12.1500 E and 25180 14.3300 N–46 410 27.5100 E
and varying densities of shrubs and trees (Vesey-Fitzgerald (Fig. 1). The Park is a large and shallow depression, which
1957; Schulz and Whitney 1986; Van der Merwe and receives runoff water several times the amount of the actual
Kellner 1999; El-Sheikh and Abbadi 2004; Alatar et al. rainfall. It is flanked by Aramah escarpment and adjoining
2012). wadis. Aramah escarpment, which is capped with upper
Camping and picnicking have significantly increased Cretaceous limestone, extends, 250 km northeast of Arri-
over the past few decades and helped to accelerate the yadh with maximum elevation of 805 m a.s.l. and 120 m
deterioration of the once intact and ecologically balanced above the nearby sandy plains (Al-Nafie 2008).
desert habitats (Al-Farraj 1989; Hajara and Batanouny The Thumamah Park is divided into different zones,
1977). Whereby, establishment of natural parks was each with a specific purpose, from limited camping area to
taken much attention to support management of the ‘high energy activity areas’. A large portion of the study

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Rend. Fis. Acc. Lincei (2013) 24:349–367 351

sloping and vertical cliffs interrupted by well developed

wadi system, sand and gravel areas, and small mosaic
depressions (Brown 1960). The upland plateau is mostly
bare gravel land interrupted by small depressions
‘‘grooves’’ which support few plant growths (Fig. 1).
The rainfall in the area is highly variable. The mean
annual precipitation is \20 mm; of which 80 % is received
from December through April and the prevailing winds
from north. The mean minimum and maximum annual
temperature is 15 and 37 C. The maximum temperature
often reaches 47 C during July–August, while the lowest
minimum temperature touches below 3 C in some
extreme cold days during winter (Fig. 2).

2.2 Sampled stands

A total of 23 stands were distributed to cover the various

landforms and succession stages (5 in sandy plain outlet, 15
in the escarpment and 3 in the upland plateau). The sam-
pling process was carried out during spring when most
species were expected to be growing. In each stand listing
of all species and their life forms and chorotypes were
determined. Species nomenclature followed Chaudhary
(1999, 2001) and Collenette (1999). In each stand, species
present were recorded and plant cover was estimated
quantitatively using the line intercept method (Canfield
1941), with five parallel lines each of 50 m long. The
lengths of interceptions by each species in each stand were
measured in centimeter, summed up and related to the total
length of lines stretched in that stand. The cover was
expressed as m 100-1 m. Phenological observations were
recorded on randomly chosen individuals of each plant
population. Permanent plots each of (10 9 10 m) were
placed at the sampled stands in the nature park. Periodic
observations were carried out every 10 days during spring
2011 (the peak growing season of most species) and
Fig. 1 a Map of the study area and landscape of the sampled stands
(1–23: •). b Landscape units of the study area. The vegetation groups monthly during the other seasons. Five phenological phases
after identified by TWINSPAN are: VG I: Rhanterium epapposum– were defined: vegetative growth, flowering, fruiting, seed
Rhazya stricta, VG II: Pennisetum divisum–Haloxylon salicornicum), dispersal and dormancy. For each species, at least 20
VG III: Acacia gerrardii–Panicum turgidum, VG IV: Panicum marked individual plants were observed. Mean values of
turgidum, VG V: Acacia ehrenbergiana–Lasiurus scindicus, VG VI:
Helianthemum lippii and VG VII Helianthemum kahiricum individuals of every species were determined.

area is also left unattended which is now regarded as a 2.3 Soil analysis
protective desert zone and currently serving as a land bank
for future recreation use. Geomorphologically its land- Three soil samples, down to 50 cm depth, were collected
scape, is identified into three fundamental components: from each stand and mixed as one composite sample for
lower plain (lowland), escarpment and upland plateau. The each stand. Soil texture was determined by hydrometer
average highest point on the site is 805 m a.s.l and the method (Allen et al. 1989). Total organic matter was
average lowest point is 560 m a.s.l giving altitudinal range determined based on loss-on-ignition at 450 C. Soil water
of 245 m. The lowland is composed of flatlands including: extract was prepared (1:5), by dissolving 100 g air dried
depressions of various areas, crossing wadis and run off soil in 500 ml distilled water for estimation of pH and
channels, sandy and gravel plains. The escarpment is the electrical conductivity (EC) as mS cm-1. Soil nutrient
dominant component of the site and characterized by both elements (Ca, K, Na, Mg, Fe, N and P) were determined

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352 Rend. Fis. Acc. Lincei (2013) 24:349–367

Fig. 2 Clima-diagram of Riyadh (623 m) Average: 24.8 °C, 11 mm

Arriyadh Region (1985–2008: 1985-2008 (23 yr)
23 years) 40 80

35 70

30 60

Temperature
25 50

Rainfall
20 40

15 30

10 20

5 10

0 0
J F M A M J J A S O N D
Temp. °C Rainfall mm

using Spectrophotometer (model ICP MSEOS 6000 Ser- and soil variables on the other hand were tested using
ies). All procedures are outlined by Allen et al. (1989). Pearson’s simple linear correlation coefficient (r). The
variation in the species diversity, stand traits and soil
2.4 Data analysis variables in relation to plant community were assessed
using one-way analysis of variance (SAS 1989, 1996).
The cover estimates of 119 species were recorded in 23
stands, and were subjected to multivariate analysis. Two
data matrices of common species cover data were created: 3 Results
(1) matrix of 23 stands 9 119 common species cover value
and (2) matrix of 23 stands 9 48 of the most common 3.1 Floristic diversity and chorotypes
species cover values 9 soil variables. The first matrix was
subjected to a numerical classification using two-way Total of 119 species (51 annuals and 68 perennials),
indicator species analysis ‘‘TWINSPAN’’ (Hill 1979a). belonging to 95 genera and 37 families were recorded from
TWINSPAN produces a hierarchical classification of veg- various stands, of which the most represented families are
etation groups (i.e. plant communities). Plant communities Asteraceae (17 %) and Poaceae (11 %). Therophytes con-
were named after their dominant species. The Detrended stituted 46 species (41 %) followed by 33 Chamaephytes
Correspondence analysis ‘‘DCA’’ (Hill 1979b) was applied (30 %) and 18 hemicryptophytes (15 %). Species of rud-
to the same first matrix data sets in order to obtain an eral habitats were recorded, such as Cynodon dactylon,
efficient graphical representation of ecological structure of Malva parviflora, Senecio glaucus and Calotropis procera,
the vegetation groups identified using TWINSPAN. To and parasitic species Cuscuta planiflora and Cistanche
detect correlations of the derived vegetation associations tubulosa (Appendix and Fig. 3). Regarding the chorotypes,
with environmental data, Canonical Correspondence the monoregionals were highly represented in most life
Analysis ‘‘CCA’’ according to (Ter Braak and Smilauer forms except for the phanerophytes where the biregionals
2002) was conducted with very common species cover, were the highest. Saharo-Arabian components were the
stands and soil variables ‘‘Second matrix’’. highest among the monoregionals (about 64 %), while the
Gamma species diversity (c-diversity) was calculated as Sahelien-Somali Masai were the highest among the bireg-
the total species number in each landscape or vegetation ionals (46 %).
group. Species richness (a-diversity) of the vegetation
cluster was calculated as the average number of species per 3.2 Multivariate analysis of plant communities
stand. Species turnover (b-diversity) are calculated as
P
(a c-1). Shannon–Wiener index Ĥ = - si¼1 pi log pi for TWINSPAN dendrogram divided the data set into seven
P
the relative evenness, and Simpson index C = si¼1 p2i for major vegetation groups (i.e., plant communities) at level
the relative concentration of dominance were calculated for 3. These communities were named after the dominant and
each stand on the basis of the relative cover pi of the ith subdominant species as follows: the first two groups (VG I:
species (Pielou 1975; Magurran 1988). Relationships Rhanterium epapposum–Rhazya stricta, and VG II: Pen-
between the ordination axes on one hand, and community nisetum divisum–Haloxylon salicornicum) only dominated

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by psammophytic species, which occupied the lower sandy turgidum) and VG V (Acacia ehrenbergiana–Lasiurus
plain. The escarpment habitat was dominated by VG III scindicus). The remaining two groups occupied the upland
(Acacia gerrardii–Panicum turgidum), VG IV (Panicum plateau (VG VI: Helianthemum lippii and VG VII: He-
lianthemum kahiricum). The application of DCA and CCA
0 50 100
100 Phanerophytes confirmed the separation between these communities and
indicated the relationships between some environmental
and topographic gradients of the park area (Fig. 4a, b).
Hemicryptophytes

Chamaephytes 3.3 Plant communities–environment relationships

50 CCA ordination was used to verify the correlation analysis

Therophytes between the dominant environmental factors and CCA axes
(Fig. 5a, b; Table 1). Correlation analysis indicated that the
separation of the species along the first axis is positively
affected by altitude, EC and Mn content (r = 0.79, 0.40
and 0.40, respectively). On the other hand, organic matter
0
(r = 0.64) and K (r = 0.44) are positively correlated with
Monoregional Biregional Multiregional the second axis, while Ca is negatively correlated (r =
-0.53) (Table 1). Steppe species type in communities I
Fig. 3 Layer diagram of the life form and chorotype spectra of and II (e.g., Rhanterium epapposum, Rhazya stricta, Pen-
Thumamah Park nisetum divisum, Haloxylon salicornicum, Stipagrostis
Fig. 4 Relationship between a
the seven plant communities
after the application of
TWINSPAN (a) and DCA (b)

b
400
Axis 2 VGI
350
VG II
300
250 VG III

200 VG IV

150 VG V
100
VG VI
50
Axis 1 VG VII
0
0 100 200 300 400 500 600

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plumosa) on the negative axis 1 are correlated with fine of upland plateau. The grass-shrubby species in community
sand and Fe content. These combinations are typical in the V (e.g., Acacia ehrenbergiana, Lasiurus scindicus, Plan-
runnels and depressions of lowland plain. The cliff plant tago boissieri, Stipagrostis drarii, Convolvulus austro-
species of communities VI and VII (e.g. Helianthemum aegyptiacus, Salvia aegyptiaca) on the positive axis 2 is
species, Fagonia bruguieri, Farsetia depressa, Salvia correlated with organic matter. These species are typical in
spinosa) on the positive axis 1 are correlated with altitude, the deep depressions at the foothills of the escarpment. On
EC and Mn. These combinations are typical in the fissures the other hand, the species of pseudo-savanna communities

Fig. 5 CCA biplot with

environmental variables
(arrows), the stands a abundant
species b. For complete species
name, see ‘‘Appendix’’

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Rend. Fis. Acc. Lincei (2013) 24:349–367 355

III and IV (e.g., Acacia gerrardii, Panicum turgidum, without clear distribution. These combinations are typical
Aristida adscensionis, Ziziphus nummularia, Lycium shawii in the main wadi channels and narrow runnel of the
and Hyparrhenia hirta) on the center of the CCA diagram escarpment. (Fig. 5a, b).
According to correlation analysis of species diversity
Table 1 Interset correlations of environmental variables with CCA
and edaphic variables (Table 2), the species richness,
axes
turnover and evenness are positively correlated with total
Variable AX1 AX2 cover (r = 0.56, 0.30 and 0.40, respectively) and nega-
Altitude m-1 0.79*** -0.05 tively with pH (r = -0.47, -0.45 and -0.41, respectively)
EC mS cm-1 0.35* 0.19 and Mn (r = -0.40, 0.30 and -0.41, respectively). On the
pH 0.01 0.04 other hand, the species dominance correlated negatively
Bulk soil (%) with cover (r = -0.30) and positively with pH, coarse
O.M. -0.20 0.64***
sand, and Mn (r = 0.31, 0.34 and 0.30, respectively). The
Coarse sand -0.28 0.28
altitude correlated positively with EC, silt and Mn
(r = 0.46, 0.54 and 0.46, respectively), but negatively with
Fine sand -0.13 -0.24
coarse sand (-0.43).
Silt 0.29 -0.08
Clay 0.11 -0.09
3.4 Species diversity
Minerals soil (ppm)
Ca 0.04 -0.53**
The lowland habitat demonstrated the high value of con-
K 0.26 0.44*
centration of dominance (0.33) and lower species relative
Na -0.03 -0.13
evenness (0.73) (Table 3). The escarpment habitats are
Fe -0.26 0.07
characterized by high values of species cover, gamma
Mg 0.11 0.09
diversity, species richness, species turnover and species
Mn 0.40* 0.14
relative evenness (163.6, 111.0, 25.0, 4.4 and 1.09,
Significant at * p B 0.05, ** p B 0.01 and *** p B 0.001 respectively) and low value of species dominance (0.17).

Table 2 Correlation analysis (Pearson correlation coefficient = r) of the species diversity and environmental variables
Variable Species Species Species rel. Species Species Altitude
richness turnover evenness dominance cover

Species richness (a) (sp stand-1) 1.00

-1
Species turnover (b) (c a ) 0.52** 1.00
Species relative evenness Shannon evenness (Ĥ) 0.78*** 0.36 1.00
Species dominance Simpson dominance (C) -0.50** -0.04 -0.68*** 1.00
-1
Cover (m 100 ) 0.56** 0.30* 0.40* -0.30* 1.00
Altitude m-1 0.01 0.05 0.02 -0.23 -0.14 1.00
EC mS cm-1 -0.12 -0.06 0.07 -0.20 -0.31 0.46*
pH -0.47* -0.45* -0.41* 0.31* -0.19 -0.04
Bulk soil (%)
Org. matter -0.19 -0.12 -0.11 0.12 -0.15 0.02
Coarse sand 0.01 -0.02 -0.03 0.34* -0.20 -0.43*
Fine sand 0.17 0.02 -0.14 0.05 0.13 -0.08
Silt 0.01 0.31* 0.05 -0.16 0.07 0.54**
Clay -0.09 -0.24 0.06 -0.22 0.07 -0.05
Minerals soil (ppm)
Ca -0.21 -0.21 -0.19 -0.05 0.07 0.02
K -0.28 0.02 -0.22 0.11 -0.03 0.26
Na -0.05 -0.25 -0.11 0.07 0.05 0.35
Fe 0.08 0.52** 0.15 -0.03 0.40* -0.24
Mg -0.14 0.12 0.01 -0.17 -0.03 0.37
Mn -0.40* -0.30* -0.41* 0.30* -0.13 0.46*
Significant at * p B 0.05, ** p B 0.01 and *** p B 0.001, and significant values are in bold

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On the other hand, the upland habitat showed low value of fragrantissima, Teucrium oliverianum, Pancratium tortuosum
species cover, gamma diversity, species richness and spe- and Echium arabicum; and (c) species with accidental flow-
cies turnover (73.33, 32.0, 14.0 and 2.28, respectively). ering including Fagonia bruguieri, F. glutinosa, Farsetia
aegyptia, Lycium shawii and Zilla spinosa. During the acci-
3.5 Edaphic variables dental flowering period, the fruits generally do not produce
viable or mature seeds, but act as photosynthetic organs. All
The plant communities in the low altitude habitats support species exhibited reproductive activity during period
soils with high value of fine sand (22.8 %), Ca 3–6 months with either unimodal or bimodal peaks for
(1171.8 ppm), Na (1646.8 ppm) and Fe (6752.6 ppm), low flowering and fruiting.
salinity (120.2 mS cm-1) and organic matter (0.66 %)
(Table 4). The escarpment habitat is characterized by high
values of coarse sand (26.23 %), and low values of pH, silt, 4 Discussion
clay, Ca, K, Mg and Mn (7.17, 30.1 %, 23.6 %, 669.8 ppm,
705.5 ppm, 1,607.2 ppm and 40.6 ppm, respectively) Variations of the landscape in the study area reflect the
(Table 4). On the other hand, the upland habitat has low vegetation and floristic differences among sites. The con-
value of coarse and fine sand (7.2 and 11.8 %), and high tribution of annual plants, that represented 43 % of the
values of salinity, pH, silt, clay, K, Mg and Mn (266.7 mS total flora in the region, is slightly lower than the woody
cm-1, 7.27, 50.8 %, 30.0 %, 2750 ppm, 3,056.7 ppm, plants. These results are comparable with those of Man-
118.3 ppm, respectively). daville (1990) who estimated the annuals as 46 %, but not
As for Rhanterium epapposum–Rhazya stricta commu- comparable with Alatar et al. (2012) on Wadi Al-Jufair
nity (I) that inhabits the sandy sheet of lowland had the (51 %). This variation in the life form reflects the differ-
highest values of fine sand (24.6 %) and Ca (1237 ppm); ence in topography, human impacts, and aridity due to high
but with the lowest of organic matter (0.63 %) and Fe temperature, low rainfall with unpredictable duration and
(324 ppm) (Table 4). Panicum turgidum community (IV) distribution and hence soil instability (see Hutchinson et al.
which inhabits the slope runnels of escarpment attained the 2000; Kürschner and Neef 2011; Alatar et al. 2012;
high value of Na (2,069 ppm) and the lowest of pH (7.12), El-Sheikh et al. 2012).
clay (16.8 %) and Mn (28.4 ppm). Acacia ehrenbergiana– Among the chorotypes, the Saharo-Arabian elements in
Lasiurus scindicus community (V) that inhabits the the monoregional category constitute the major floristic
depression of escarpment had the highest of organic matter component. This can be explained by the fact that the study
(2.4 %) and coarse sand (42.7 %), but lower values of area is located in the center of Saharo-Arabian region
altitude, EC, silt, Ca, and Mg (582.7 m, 97.0 mS cm-1, (Zohary 1973; Kürschner 1986; Ghazanfar and Fisher
23.6 %, 171.3 and 1,348.0 ppm, respectively). On the other 1998; Hegazy et al. 1998; El-Ghanem et al. 2010;
hand, Helianthemum lippii (VI) that inhabits the upland Kürschner and Neef 2011; Alatar et al. 2012). The Sahe-
grooves had high contents of EC, silt, K, Mg and Mn (310.0 lien-Somali Masai elements were the highest among the
mS cm-1, 53.5 %, 3,261.50, 3,580.00 and 144.00 ppm, bi-regionals for phanerophytes. Due to most of the south-
respectively), but lower values of the fine sand (10.5 %). western and southern part of Arabia are strongly influenced
Helianthemum kahiricum community (VII) dominated the by a tropical climate and monsoonal rainfalls. Zohary
upland habitats at 782.0 m, which is characterized by neu- (1973) proposed a Sudanian floral region for this area also,
tral pH (7.31) and (36.0 %) clay content, with low values of which already belongs to the Palaeotropic floristic king-
coarse sand (3.6 %) and Na (1,009 ppm) contents. dom. As a meeting point, the flora in the region is the
product of historical and former phytogeographical
3.6 Phenology of common species migration routes (Zohary 1973; Mandaville 1990; White
and Leonard 1991; Hegazy et al. 1998; Ghazanfar and
The peak period of flowering and fruiting of most species Fisher 1998; Alfarhan 1999; Kürschner and Neef 2011;
in the study occurs between January and May where Alatar et al. 2012).
maximum fruiting occurs during late spring and early Vegetation in the lowland habitat in the outlet of sandy
summer. A diagrammatic representation of the phenologi- wadis are colonized by two pseudo-steppe clusters, Rhan-
cal stages of the most common plants in the study area is terium epapposum–Rhazya stricta community (I) on the
presented in (Fig. 6). Three major phenological categories deep sandy plain sheets and Pennisetum divisium–Halox-
feature the vegetation; (a) species vegetative all the year ylon salicornicum community (II) on the shallow gravel
round, e.g., Acacia gerrardii, Calotropis procera, Lycium runnels depression. These communities are almost similar
shawii and Rhazya stricta; (b) species dormant during to the communities described 30 years ago (Baierle and
summer/autumn months, e.g., grasses, Achillea Kürschner 1985) in the study area. These communities

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 Table 3 Presence and cover percentage of the characteristic species and the diversity variables (mean ± standard deviation) of seven vegetation units in different habitats of Thumamah Park
Landscape habitat (Macro habitat) Lowland Escarpment Upland plateau Total Mean F-value
Microhabitat Sandy Depres. Mean Wadi cliffs Slope Depres. Mean Grooves Grooves Mean
sheets channel runnels
Community unit VG I VG II VG III VG IV VG V VG VI VG VII
Stand No. 10,13,14 1,15 8,11,12, 4,5,6,7,20 2,3,9 21,22 23
16,17,18,19)
G/P (%) 13 9 22 31 22 13 65 9 4 13
-1
Percentage % and (cover m 100 ) of Dominant species in landscape habitat
I- Rhanterium epapposum- Rhazya 100 (61) – 60 (37) 72 (10) 80 (11) – 60 (8) – – –
Rend. Fis. Acc. Lincei (2013) 24:349–367

stricta 100 (12) 100 (49) 100 (27) 14 (2) – – 7 (0.7) – – –

II- Pennisetum divisum - 100 (16) 100 (30) 100 (26) 29 (10) 20 (0.2) – 20 (5) – – –
Haloxylon salicornicum 100 (9) 100 (23) 100 (15) 85 (13) 80 (5) – 67(8) 50 (0.5) – 25 (0.3)
III- Acacia gerrardii- – – – 86 (22) 40 (6) – 54 (13) – – –
Panicum turgidum – 50 (1) 20 (0.4) 86 (22) 100 (21) – 73 (18) – – –
IV-Panicum turgidum – 50 (1) 20 (0.4) 86 (22) 100 (21) 73 (18) – – –
V-Acacia ehrenbergiana- – – – – 20 (6) 67 (13) 20 (5) – – –
Lasiurus scindicus – 100 (28) 40 (12) 43 (3) – 100 (24) 40 (6) – – –
VI-Helianthemum lippii 66 (1) – 40 (0.6) 43 (0.7) 20 (0.2) – 27 (0.4) 100 (17) – 50 (9)
VII- Helianthemum kahiricum – – – – – – – – 100 50 (5)
(10)
Community or diversity variables
Cover (m 100-1) 116.0 187.0 144.4 215.9 ± 74.9 116.4 120.0 163.6 93.0 34.0 73.33 147.6 2.61*
± 15.5 ± 130.1 ± 76.5 ± 56.4 ± 28.1 ± 77.4 ± 26.8 ± 39.0
Gamma diversity (c) Total species 16.0 25.0 34.00 51.0 42.0 29.0 111.0 24.0 7.0 32.00 119.0
no.
Species richness (a) (sp stand-1) 10.67 16.50 13.00 29.28 ± 7.1 24.4 16.0 25.00 17.00 8.0 14.00 21.0 6.59***
± 1.1 ± 9.1 ± 5.6 ± 3.8 ± 3.4 ± 7.3 ± 2.82 ± 5.5
-1
Species turnover (b) (c a ) 1.49 1.52 ± 0.5 2.60 1.74 ± 0.23 1.72 1.81 4.44 1.41 0.88 2.28 3.1 0.01
± 0.12 ± 29.3 ± 0.15 ± 0.21 ± 19.4 ± 0.16 1.0 ± 0.12
Species relative evenness 0.57 0.96 0.73 1.16 1.10 0.88 1.09 0.97 0.83 0.93 0.99 5.91**
Shannon evenness (Ĥ) ±0.2 ±0.19 ±0.27 ±0.07 ±0.09 ±0.28 ±0.17 ±0.17 ±0.15
Species dominance 0.45 0.15 0.33 0.11 0.12 0.43 0.17 0.18 0.15 0.18 0.21 5.84**
Simpson dominance (C) ±0.23 ±0.04 ±0.23 ±0.03 ±0.05 ±0.17 ±0.15 ±0.08 ±0.06
G/P the percentage of the stands representing each vegetation group in relation to the total sampled stand.* p B 0.05, ** p B 0.01 and *** B 0.001, according to one-way ANOVA. Max. and
min. values are in bold
357

123
 Table 4 The edaphic variables of the seven vegetation groups (mean ± standard deviation) as identified after TWINISPAN: VG I-Rhanterium epapposum–Rhazya stricta, VG II-Pennisetum
358

divisum–Haloxylon salicornicum, VG III-Acacia gerrardii–Panicum turgidum, VG IV-Panicum turgidum, VG V-Acacia ehrenbergiana–Lasiurus scindicus, VG VI-Helianthemum lippii and
VII-Helianthemum kahiricum

123
Landscape macro Lowland Escarpment Upland plateau Total F-value
habitat Mean
Microhabitat Sandy Depr. Mean Wadi cliffs Slope Depr. Mean Grooves Grooves Mean
sheets channel runnels
Community unit VG I VG II VG III VG IV VG V VG VI VG VII
Stand no. 10,13,14 1,15 8,11,12, 4,5,6,7,20 2,3,9 21,22 23
16,17,18,19
G/P (%) 13 9 22 31 22 13 65 9 4 13

Edaphic (soil) variables

Altitude m 618.7 596.5 609.80± 21.2 648.9 ± 40.9 619.6 582.7 625.87 753.0 782.0 762.67 ± 27.6 640.22 7.35***
± 15.0 ± 27.5 ± 50.4 ± 17.2 ± 46.5 ± 31.1
EC mS cm-1 123.33 115.50 120.19± 25.2 138.57 ± 65.2 108.4 97.0 120.20 310.0 180.0 266.67 ± 150.1 139.30 2.84*
± 18.9 ± 41.7 ± 22.0 ± 29.1 ± 50.7 ± 182
pH 7.23 7.30 ± 0.14 7.26 ± 0.13 7.18 ± 0.13 7.12 7.20 7.17 ± 0.11 7.25 7.31 7.27 ± 0.05 7.20 0.77
± 0.15 ± 0.11 ± 0.10 ± 0.07
Bulk soil (%)
OM (%) 0.63 0.72 ± 0.28 0.66 ± 0.23 0.97 ± 0.61 0.61 2.40 1.13 ± 0.92 1.35 1.24 1.31 ± 0.12 1.06 3.40*
± 0.26 ± 0.41 ± 1.15 ± 0.15
Coarse sand 18.02 13.68 16.29 ± 8.12 14.75 ± 15.6 32.43 42.67 26.23 9.00 ± 7.5 3.64 7.21 ± 6.1 21.59 3.54*
± 5.3 ± 13.6 ± 10.3 ± 7.0 ± 16.7
Fine sand 24.60 20.02 ± 4.2 22.77 ± 4.10 22.29 ± 11.3 19.61 15.71 20.08 ± 9.2 10.51 14.9 11.97 ± 5.1 19.61 0.80
± 3.4 ± 8.9 ± 2.5 ± 6.2
Silt 34.71 35.30 34.94 ± 13.2 32.11 ± 13.3 31.16 23.61 30.09 53.49 45.46 50.81 ± 9.2 33.85 1.46
± 12.7 ± 19.2 ± 9.4 ± 6.7 ± 10.9 ± 11.3
Clay 22.67 31.00 26.00 ± 11.2 30.86 ± 25.5 16.8 18.0 ± 5.3 23.60 27.0 36.0 30.00 ± 10.3 24.96 0.52
± 14.4 1.4 7.8 ± 18.7 ± 12.7
Minerals soil ppm
Ca 1,237.3 1,073.5 1,171.80 831.57 ± 407 742.4 171.33 669.80 877.0 833.0 862.33 ± 519 804.04 2.87*
± 100 ± 347 ± 219.2 ± 149.9 ± 97.2 ± 383.8 ± 732
K 773.0 2,420.0 1,431.80 572.71 ± 286 538.2 1,294.0 705.47 3,261.5 1,727.0 2,750.00 1,130.04 2.05
± 90.1 ± 2,231 ± 1,436 ± 428 ± 1,238 ± 634.0 ± 3,602 ± 2,697
Na 1,644.7 1,650.0 1,646.80 ± 511 1,341.1 ± 836 2,069.2 1,278.7 1,571.33 1,068.5 1,009.0 1,048.67 ± 181 1,519.57 0.88
± 519 ± 711 ± 795 ± 342 ± 794 ± 252
Fe 324.3 16,395 6,752.60 1,227.3 450.8 3,224.7 1,367.93 331.0 2,432.0 1,031.33 2,494.61 2.05
± 346.2 ± 2,2920 ± 1,4452 ± 2,071 ± 358 ± 2,505 ± 2,163 ± 62.2 ± 1,213
Mg 1,696.7 2,898.0 2,177.20 1,615.9 ± 570 1,750.6 1,348.0 1,607.20 3,580.0 2,010.0 3,056.67 1,920.17 2.81*
± 299 ± 1,548 ± 1,037 ± 519 ± 997 ± 616 ± 1,131 ± 1,208
Mn 74.67 92.00 81.60 ± 49.8 49.00 ± 27.6 28.4 41.33 40.60 144.0 67.0 118.33 ± 44.4 59.65 3.64**
± 59.1 ± 50.9 ± 12.1 ± 35.6 ± 25.4 ± 1.4
Rend. Fis. Acc. Lincei (2013) 24:349–367
Rend. Fis. Acc. Lincei (2013) 24:349–367 359

25.59***

G/P the percentage of the stands representing each vegetation group in relation to the total sampled stand. * p B 0.05, ** p B 0.01 and *** B 0.001, according to one-way ANOVA. Max. and
have a mosaic like pattern with dominant shrubby vege-

8.96***
F-value tation that have a well developed root system, capable of
regeneration and that enables plants to survive period of
burial (Kürschner 1986; El-Sheikh et al. 2010). The asso-
ciated psammophytic species survive densely around these
Mean
Total

2.50

2.09
dominant shrubs which can make small phytogenic
mounds, where aeolian of the fine particles deposits
covered the land surface (i.e., wind protected areas).
3.17 ± 1.2

3.83 ± 2.1
Therefore, these communities are characterized by high
species dominance and moderate species diversity. In such
Mean

13

cases, most of the plant cover is accounted by one or two

dominant species that capable to use the available recour-
Grooves

VG VII

ses, due to their high competitive capacities under envi-

3.65

6.25
23

ronmental stress. Our results are similar to the effect of

4
Upland plateau

hummock formations on the reduction of species evenness

2.94 ± 1.7

2.62 ± 0.7

and diversity, which also detected in Central Saudi Arabia,

Grooves

VG VI

Kuwait and North Africa (Shaltout and Mady 1996; Shal-

21,22

tout and El-Ghareeb 1992; El-Sheikh et al. 2010; Wale

9

et al. 2012). Moreover, these habitats have little substrate

2.10 ± 0.29
2.38 ± 1.0

heterogeneity and low pronounced relies and dominated by

pseudo-steppe vegetation (Baierle and Kürschner 1985).
Mean

The escarpment habitat types are colonized by three

65

community types (III, IV and V): Acacia gerrardii–Pani-

cum turgidum (III) inhabits the main wadi channels of the
± 0.65

± 0.20

escarpment cliffs, Panicum turgidum (IV) inhabits the

VG V
Depr.

2,3,9

0.40

2.25

narrow runnel of the escarpment slopes, and Acacia

13

ehrenbergiana–Lasiurus scindicus (V) inhabits the deep

4,5,6,7,20

± 0.20

± 0.38

depressions of the foothills in the escarpment. These are

runnels
VG IV
Slope

wadi and runnel communities of vein-like structure which

2.92

1.98
22

represent a different size of the catchment areas and

dominated by prominent component of the native range
2.13 ± 0.26
16,17,18,19
Escarpment

Wadi cliffs

flora and scattered arboreal and scrubby-grass vegetation

2.85 ± 0.4
8,11,12,

(i.e., extremely xeromorphic woodlands or pseudo-savan-

VG III

nas of thorn woodlands). Deep wadi cliffs and deep

31

depressions canyon are dominated by Acacia spp. which

represents the climax stage of succession of the xerophytic
vegetation in Arabia (Zohary 1973; Kürschner 1986;
2.46 ± 1.0

0.99 ± 0.9

Shaltout and Mady 1996; Aref et al. 2003; El-Sheikh 2005;

Mean

Kürschner and Neef 2011). The habitat types have alluvial

22

fan, fluvial coarse sand, gravels with moderately drained

1.45 ± 0.01

1.81 ± 0.02

soil and characterized by high organic matter content and

low salinity, pH, and minerals. The soils are often derived
channel
VG II
Depr.

from weathered calcareous formations by rugged relief,

1,15

rapid runoff and pronounced water erosion (karstic rocks).

Hence, the deriving heavy coarse sand particles from cal-
± 0.71

± 0.74
Lowland

10,13,14

careous escarpment formations was fallen firstly in the

Sandy
sheets
VG I

3.13

0.46

deep wadis or gullies and rocky basin depressions, whereas

min. values are in bold
13

the light fine sand particles will travel by wind to long

Table 4 continued
Landscape macro

distance and drops in the lower sand plain land of the park,
Community unit

which characterized by the fine soil particles. The coarse

DCA axis1

DCA axis2
Microhabitat

Stand no.

sand particles accumulate with large amounts of the plant

G/P (%)
habitat

debris, which shed at the soil surface of these deep

depressions or deep canyon-like structure.

123
360 Rend. Fis. Acc. Lincei (2013) 24:349–367

Fig. 6 Phenology of the common selected species in Thumamah Park

In addition, the community III which inhabits the deep In general, the escarpment microhabitats are charac-
main wadi channel of the escarpment cliffs has high plant terized by special type of ‘‘biotopes’’, which mostly
cover, species richness, species relative evenness and lower formed by water erosion and formed deep and narrow
species dominance. Acacia gerrardii dominates this wadi canyon-like biotopes with the heterogeneity of the soil
and is considered as one of the two large-trunked Acacias substrates. These represent protected gaps with small
in Arabia that indicates high ground water sources (Man- sized biotope, providing high amount of ecological niches
daville 1990; Aref et al. 2003). This may explain the high for different species with good water supply that can
species cover and attainment the final stage of pseudo- hardly be found in the rest of the Park and therefore the
savannas climax. Moreover, the well established popula- species diversity increase in these habitats (Baierle and
tion of Acacia gerrardii plays a secondary role in the Kürschner 1985). For these reasons and since most of the
enhancement of the ecological niche conditions such as runnels, wadis and drainage systems being major com-
favorable microclimate under trees by providing a shelter ponent in the landscape, the area has an ecological key
or nourishment to other species, or even act as a key plant functional role for the whole park and its surroundings
for many birds and animals and thereby increase the plant (Alday et al. 2011; Wale et al. 2012). It is an important
species diversity through the dispersal of their seeds and regeneration center and acts as distribution center or
fruits (Baierle and Kürschner 1985; Kürschner 1986; nucleus area for regeneration of plants and animals for the
Batanouny 1987; El-Sheikh 2005). whole area.

123
Rend. Fis. Acc. Lincei (2013) 24:349–367 361

The communities inhabiting the upland plateau are recover fully from the effects of prolonged grazing and
dominated by Helianthemum lippii (VI) and Helianthemum other large-scale human disturbances.
kahiricum (VII). Within both clusters, vegetation is Altitude is correlated with the increase of salinity, silt,
restricted to the small fissures or grooves in colline rock of Na, Mg, Mn and had low species diversity. Because the
the upland plateau. These communities are representing increasing salinity and minerals content, usually resulted
habitat types characterized by high salinity, pH and clay from rock erosion, where the direct action of wind and
contents and lower values of plant cover and species heating effect of the sun which increases the evaporation
diversity (i.e., xerodrymlum community with poor species rates from exposed fissures soil, hence the water table
productivity and diversity). These habitat types are char- decrease and salinity and minerals become concentrated.
acteristic to the highest altitude of our study area. These Correspondingly, our results show that species richness
vegetation groups are adapted to low nutrient availability, decreased markedly, with increased dosage of fertile soil,
narrower local distribution in the grooves or fissures agree with the general trend and therefore, upland plateau
between rocks and represent the earlier stages of xerarch are restricted by dwarf ‘‘xerosere’’ plant community
succession ‘‘xerosere’’. As described by Alatar et al. (Kürschner 1986; Batanouny 1987; El-Sheikh and Abbadi
(2012), Wale et al. (2012), these communities are charac- 2004; Wale et al. 2012).
terized by their low productivity. In general, the Park vegetation in the present study has
The correlation analysis indicates that species richness, higher species cover and species diversity indices than the
evenness and turnover are correlated positively with total vegetation of the other regions (see Table 5). Moreover,
plant cover and negatively with pH and Mn, whereas the study of Zahran and Younes (1990) on a fully protected
species dominance correlated positively with the coarse area in the region of Khamis Mushait with the mean annual
sand and negatively with total plant cover. Similar corre- rainfall of 242 mm year-1 indicated that the total cover
lations were reported in Pysek et al. (2004) and El-Sheikh was 96 % inside the protected area and 45 % in the free
et al. (2012) in late stage of succession of the desert veg- grazing areas. In the region of Abha, the mean annual
etation. Because many different life forms, the tree, shrub rainfall is 332 mm year-1, whereby Abulfatih et al. (1989)
and grass species have different functional behavior. The reported an increase of phytomass that reached 267 % after
interspecific competition increases at the medium and later 5 years of protection. The increase of species richness and
successional stages than in the early stages, which in typ- total plant cover was mainly due to emergence of palatable
ical plots, most of the species cover was accounted for by herbs and grasses, which was usually decreased in response
one or two species. Meanwhile, the species dominance to heavy grazing before fencing the Park (Hajara 1993;
increases and species richness decreases at the first stage of Shaltout et al. 1996; Assaeed 1997; Havstad et al. 1999;
succession (Fynn et al. 2011; El-Sheikh et al. 2012). Our Floyd et al. 2003; Qinfeng 2004; El-Sheikh et al. 2006;
results are coincided with those of Qinfeng (2004) who Wale et al. 2012).
pointed out that a continuing trend towards the increasing A variation of plant phenology in space and time
species richness and plant cover with time after protection explains the phenological diversity among perennial plant
from disturbance. He concluded that it would take between species in Thumamah Nature Park. Beside the importance
30 and 50 years for the desert perennial vegetation to of environmental factors in such phenological diversity, it

Table 5 Comparison between the species diversity indices of vegetation of the present study and that of the raudhas (depressions) in Arriyadh
and coastal lowland in the eastern region of Saudi Arabia
Diversity variables Present study 2012 Shaltout and Mady 1996 Shaltout et al. 1996
Protected Thumamah Unprotected raudhas Protected Coastal Unprotected Coastal
Park (stand size near the study area vegetation (stand vegetation (stand size
10 9 10 m) (stand size 10 9 10 m) size 20 9 20 m) 20 9 20 m)

Species cover m 100 m-1 147.6 54 66.5 39.7

Total species no ( -diversity) 119 107 68 61
-1
Species richness (a-diversity) spp. stand 21.0 3.3 18.9 14.2
Species turnover (b-diversity)  a-1 3.1 3.10 3.96 5.1
Relative evenness (Ĥ) Shannon-Wiener index 0.99 0.39 0.82 0.62
Conc. of dominance (C) Simpson index 0.21 0.56 0.28 0.38

123
362 Rend. Fis. Acc. Lincei (2013) 24:349–367

a On the lower land outlet habitats of shallow sand sheets and sand accumulations and
shallow gravel runnels depression, the progressive succession proceeds as the following:

Annual on bare sand Few perennials on sandy Rhanterium epapposum on deep

sheet and fine gravel sheet and gravelly runnels
Rhayza stricta with sand sheets and Pennisetum
desert runnel habitats habitat, (e.g. Fagonia
(e.g. Ifloga spicata, bruguieri, Launaea perennial grasses divisum on gravelly runnels with
capitata, Polycarpaea psammophytic stands rich in Haloxylon salicornicum.and
Erucaria hispanica
Plantago boissieri). repens) grasses as pseudo-steppe community

b On the escarpment habitat of the deep sandy wadis depression and small runnels, the
progressive succession proceeds as the following:

Acacia gerrardii-A.
Annual (Neurada Rhazya stricta – Zilla Pennisetum divisum Acacia ehrebegiana- ehrebegiana with
procumbens) on bare spinosa and annual with Astragalus Panicum turgidum grasses of pseudo-
sandy, gravel wadi community spinosus community shrubby community savanna as climax
and runnel habitats community

c On the upland habitat of rocky fissures, the progressive succession proceeds as the
following:

Pioneer annuals on silty of Annual with perennial Helianthemum lippii Ephedra foliata with
rock fissures: Anastatica herbs (e.g Asteriscus with Farsetia depressa Gymnocarpos decandrum
heirochuntica - Emex heirochunticus-Fagonia community xerosere community
spinosa bruguieri) community

0_____________5______________10_____________15______________20_____________25____________30 years

Fig. 7 The hypothesis of the main processes involved in the progressive succession of Thumamah Nature Park vegetation during the prolonged
conservation of 30 years a–c

may be a product of competition among plants for polli- (Penuelas et al. 2004). Soil type plays a role in water
nation activities (Stone et al. 1998). The phenological availability for plants and may produce various phenolog-
diversity around the year results from variable biotic and ical niches under same environmental variables (Hegazy
abiotic factors, particularly, the plant life forms, rainfall et al. 2012). The species having accidental flowerings
events and temperature (Marques et al. 2004; Hegazy et al. during autumn, e.g., Ochradenus baccatus, Zilla spinosa
2012). This is considered as important reproductive and and Lycium shawii do not reach into fruiting stage or
adaptive plant strategies in arid deserts (Ward 2009). All produce viable seeds. In this case flowers function as
study species exhibited a unimodal peak for flowering and photosynthetic organs rather than reproductive organs
fruiting around the year. (Kassas 1966).
The phenological behavior of Ochradenus baccatus in When a park is fenced against human impacts, the
the study area demonstrated unimodal flowering–fruiting success of the initial revegetation treatment provides a
peak covered spring and summer seasons. This contrasts starting species pool for the subsequent succession. Once
the species behavior in another arid region, wadi Degla in this pool is established, other processes come into play
Egypt, where many individuals of the species flower over including species assembly rules, rates of species coloni-
most of the year with two major peaks (bimodal) in spring zation and extinction and both positive and negative spe-
and autumn (Hegazy et al. 2012). This behavior was cies interactions; all of which interact to produce species
reported also in Ein Gedi, Palastine (Wolfe and Burns turnover and vegetation structure change. Therefore by
2001). Topographic features, historical and geographical following the course of succession those spatial and
changes in rainfall are important drivers that cause com- temporal changes in the vegetation structure and species
plex phenological changes among species and communities patterns that determine the outcome of succession can be

123
Rend. Fis. Acc. Lincei (2013) 24:349–367 363

identified (Walker and del Moral 2009; Alday et al. 2011). community of Acacia gerrardii–Acacia ehrenbergiana
Hence, as succession proceeds, there is an opportunity to (Fig. 7b). In the upland plateau, Anastatica heirochuntica–
study the dynamics of vegetation development in these Emex spinosa of pioneer annuals community establishes
disturbed ecosystems (Alday et al. 2010, 2011). The plant first and followed by annual and perennial herbs of Aste-
succession has a practical importance for future ecological riscus heirochunticus–Fagonia bruguieri community. Then
restoration and theoretical importance for understanding Helianthemum lippii and Farsetia depressa community are
the fundamental ecological theory and the initial phases of established with Ephedra foliata and Gymnocarpos
vegetation establishment (Robbins and Mathews 2009). decandrum as a dwarf xerosere community (Fig. 7c).
We can hypothesized the main processes involved the In conclusion, the community types representing
progressive succession of Thumamah Nature Park vegeta- common stages of the progressive succession reflect the
tion during the prolonged conservation of 30 years relationship between altitude, edaphic factors and the type
(Fig. 7a–c): On the bare sand sheets and fine gravel runnels of vegetation type in each habitat type. The simultaneous
of the lowland, annuals are established first followed by invasion of both early and late succession species and their
perennial herbs and grasses of psammophytic species successful establishment depends on the specific nutrient
dominated by Rhazya stricta. Finally, Rhanterium epap- and water availability, and the life cycles of the taxa. The
posum on deep sand sheets and Pennisetum divisum on findings of this study highlight that desert vegetation under
gravel runnels colonize the lowlands in association with exclusion of anthropogenic effects and mixed prevailing
Haloxylon salicornicum and grasses as pseudo-steppe environmental conditions may enhance establishment of
succession community (Fig. 7a). The progressive succes- diverse plant communities. Plant populations showed
sion in the escarpment habitats ‘‘wadi and runnels’’ pro- interspecific variations in their relative timing of pheno-
ceeds from pioneer annual herbs, e.g., Neurada logical phases with varied flowering/fruiting activity
procumbens community, then perennial herbs and shrubs periods.
of Rhazya stricta–Zilla spinosa community appear and
modify the substrate before the next succession stages of Acknowledgments This research was supported by the National
Plan for Sciences and Technology (NPST), King Saud University,
Acacia ehrenbergiana–Panicum turgidum shrubby com- Saudi Arabia, Riyadh (Project grant number 11-ENV1754-02).
munity and finally to pseudo-savanna as a diverse climax

123
364 Rend. Fis. Acc. Lincei (2013) 24:349–367

Appendix

Synoptic table of the vegetation groups in Thumamah Park after identifying by TWINSPAN: VG I-Rhanterium epapposum–Rhazya stricta, VG
II-Pennisetum divisum–Haloxylon salicornicum, VG III-Acacia gerrardii–Panicum turgidum, VG IV-Panicum turgidum, VG V-Acacia eh-
renbergiana–Lasiurus scindicus, VG VI-Helianthemum lippii and VII-Helianthemum kahiricum
I II II IV V VI VII
Spp. Life Choro- 1111 1 111 121 222
no. Species form
type 03451468812957607239123
39 Echinops erinaceus Kit-Tan Ch IT ------312233223-------2
2 Acacia gerrardii Zoh. Ph SA ------5554554-4--------
18 Calligonum comosum L'Her. Ph SA-IT -------5---------------
20 Carthamus oxyacantha M. Bieb. Th IT ----------4---1--------
71 Lycium shawii Roem. & Schult. Ph SA-SM -------5-355--5--------
101 Savignya parviflora (Del.) Webb Th SA -------2---3--1--------
119 Ziziphus nummularia (Burm.f.) Wight & Arn. Ph SH-SM -------5---------------
13 Astragalus spinosa (Forssk.) Muschl. Ch IT -------2-3------2------
29 Convolvulus buschiricus Bornm. Ch SA -----------3----2------
33 Cucumis prophetarum L. He SA ------2-----1----------
34 Cynondon dactylon (L.) Pers. He TR -------122-3--3--------
85 Pergularia tomentosa L. Ch SH-SM ------4--------2-------
104 Scorzonera musillii Vel. He IT ------212-2---1-2------
113 Teucrium oliverianum Ging. ex Benth. He SA -----------21----------
9 Artemisia monosperma Del. Ch SA ------33----53---------
19 Calotropis procera (Ait.) Ait.f. Ph SH-SM ------4-----4-2--------
32 Cuscuta planiflora Tenore Th AM -------1---21-111------
54 Gastrocotyle hispida (Forssk.) Bunge Th SA-IT --------------11-------
63 Horwoodia dicksoniae Turril Th SA ------1------1—2-------
64 Ifloga spicata (Forssk.) Sch.- Bip. Th SA ----------------5------
76 Monsonia nivea (Decne.) Decne. ex Webb. Th SH-SM -------------1—1-------
80 Pancratium tortuosum Herb. He Med ----------------2------
102 Schimpera arabica Hochst. & Steudel Th SA -------1-----1—2-------
103 Sclerocephalus arabicus Boiss. Th SA -----------3—222-------
106 Senna italica Miller Ch SH-SM ------------1---2------
11 Astenatherum forsskalii Vahl Ch SA-IT -----1--1---11-1-------
49 Farsetia aegyptiaca Turra Ch SH-SM ---2-1223-342-442------
73 Medicago laciniata (L.) Mill. Th SA -----12-------2-2------
81 Panicum turgidum Forssk. Ch SA-SM ----244442545455-------
25 Citrullus colocynthis (L.) Schrader He SA ----334-222-----2------
48 Fagonia glutinosa Del. Ch SA -----1--1—2----2-------
75 Moltkiopsis ciliata (Forssk.)I.M. John. Ch SA ----2-2---21----1------
7 Aristida adscensionis L. He Med-IT-SA -----1-------1—2-------
15 Bassia eriophora (Schrader) Asch. Th SA-SM ----2------------------
16 Blepharis ciliaris (L.) B.L. Burtt. Th SA-IT ----21-2----1-222------
26 Cleome amblyocarpa Barr. & Murb. Th SA-SM -----2----------1------
42 Ephedra alata Decne. Ph SH-SM ----23----------1------
45 Erodium laciniatum (Cav.) Willd. Th Med ----3--1-----1---------
51 Farsetia stylosa R.Br. Ch SH-SM ----312--2--11---------
60 Heliotropium arbainense Fresen. Ch SA -----2------1-2--------
69 Launaea angustifolia Boiss. Th SA -----1---------1-------
114 Tribulus macropterus Boiss. He SH-SM ----2-----------1------
115 Tribulus terrestris L. Th EU-Med-IT ----2---1-----1-2------
78 Neurada procumbens L. Th SA 2---5--2222331—5-------
91 Polycarpaea repens (Forssk.) Asch.. & Sch. He SH-SM 2----111112--1-12------
31 Convolvulus oxyphyllus ssp. oxycladus Rech. f. Ch SA 2----------------------
83 Pennisetum divisum (Gmel.) Henr. Ch SA 52245---4-5----1-------
96 Rhazya stricta Decne Ch SA 53345----4-------------
111 Stipagrostis plumosa (L.) Munro ex T. Anders. He SA-IT 222----1--------2------
36 Deverra triradiata (Hochst.) Aschers Ch SA 5-----34—34------------
46 Erucaria hispanica Druce Th Med -22-21-12—111421-------
95 Rhanterium epapposum Oliv. Ch SA 555--221--554-354------
56 Haloxylon sallicornicum (Moq.) Bunge Ch SH-SM 5225422435-5-3442-2-1--

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Rend. Fis. Acc. Lincei (2013) 24:349–367 365

continued
22 Centaurea pseudosinaica Czerp. Th SA ------21-23---1------2-
43 Ephedra foliata Boiss. ex C.A. May Ph SH-SM ------222-54--4-4----5-
21 Cenchrus ciliaris L. Ch SA ----3--13--21-22-2---2-
62 Heliotropium bacciferum Forssk. Ch SA-SM 1---4-12233--22-2—222--
88 Plantago boissieri Hausskn. & Bornm. Th SA -11-5--21--------2-----
118 Zilla spinosa (L.) Prantl. Th SA ---4—3-22543-3--32-----
79 Ochradenus baccatus Del. Ph TR AF ----2--35—4------2-----
47 Fagonia bruguieri DC. Ch SA ---12--2----1-52----2-3
97 Rumex vesicarius L. Th SA ----212-3---213223—22--
17 Calendula arvensis L. Th Med-IT ------1111----1-----12-
50 Farsetia burtoniae Oliver Ch SH-SM -------2---2---------3-
70 Leontodon laciniatus (Bertol) Widder Th SA-IT -----152---------4-----
82 Paronychia arabica (L.) DC. Th SA ------------1---2---2--
77 Moricandia sinaica (Boiss.) Boiss. Ch SA -----5--------------3-3
89 Plantago ovata Forssk. Th SA-IT -------2--2-----2—21-2-
8 Arnebia hispidissima (Lehm.) DC. Th SA-SM ---------1--------1----
41 Emex spinosa (L.) Campd. Th Med 1------2-221-1--225-1--
65 Lasiurus scindicus Henr. He SA-SM ---45--233-------255---
84 Periploca aphylla Decne Ph TR AF ----------5------4-----
59 Helianthemum lippii (L.) Doum.- Cours. Ch SA-SM -12----2----1-1-2---25-
14 Atractylis mernephethae Asch. Schweinf. Th SA -22---------------11---
61 Heliotropium aegyptiacum Lam. Ch SA ---2--1------------2---
66 Launaea capitata (Spreng.) Dandy Th SA 111--------------3-----
87 Plantago amplexicaulis Cav. Th SA -----------2--3—4-2----
5 Anisosciadium lanatum Boiss. Th SA -------1------11-4-----
28 Convolvulus austro-aegyptiacus Abdallah & Sa'ad He SA ------------12----3----
55 Gymnocarpos decander Forssk. Ch SA ---------------1----1--
72 Malva parviflora L. Th Med-IT ---------2-21-11-3-222-
1 Acacia ehrenbergiana Hayne Ph SH-SM ------------5----45----
3 Althaea ludwigii L. Th SA ---------2-------1-24--
4 Anastatica heirochuntica L. Th SA ---------------1-3-252-
6 Anvillea garcinii (Burm.f.) DC. Ch SA --------------2—2113--
30 Convolvulus pilosellifolius Desr. He IT -----------------2-1---
67 Launaea mucronata (Forssk.) Muschl. Th SA ------------------21---
68 Launaea nudicaulis (L.) Hook.f. Ch SA --------1--------3----1
94 Pulicaria undulata (L.) C.A. May Ch SH-SM ---2-------------3-----
112 Tamarix aphylla (L.) Karst. Ph SH-SM ------------------4----
117 Trigonella stellata Forssk. Th SA -----------------3-2---
12 Astragalus schimperi Boiss. Th SA --------------------2--
27 Cleome rupicola Vicary Ch SH-SM ---------------1-----2-
38 Diplotaxis harra (Forssk.) Boiss. Ch SH-SM ---------------1----2--
52 Farsetia depressa Kotschy Ch SH-SM ---------------1----22-
74 Microchloa kunthii Desv. He SA ---------------------2-
105 Senecio glaucus L. Th SA-IT --------------1------2-
10 Asteriscus heirochunticus (Michon) Wiklund Th SA ---------------1------2
44 Erodium glaucophyllum (L.) L'Her He SA ----------------------3
58 Helianthemum kahiricum Del. Ch SA ----------------------4

The cover levels are coded as follows: 1, B 10 %; 2, 10–20 %; 3, 20–30 %; 4, 30–40 %; 5, 40–50 %; 6, 50–60 %; 7, 60–70 %; 8, 70–80 %;
9, C 90 %. The life forms are: Th therophyte, Ch chamaephyte, Ph phanerophyte, He hemicryptophyte. The chorotypes are: IT Irano-Turanian,
SA-IT Saharo-Arabian–Irano-Turanian, SH-SM Sahelian-Somali Masai, TR tropical, SA Saharo-Arabian, SA-SM Saharo-Arabian-Somali Masai,
TR AF tropical African, TR AM tropical American, Med-IT Mediterranean–Irano-Turanian, EU-Med-IT Euro-Siberian–Mediterranean–Irano-
Turanian, Med Mediterranean, Med-IT-SA Mediterranean–Irano-Turanian–Saharo-Arabian, Cosm cosmopolitan, TR AF-SA Tropical Africa–
Saharo-Arabian

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