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Journal of Arid Land
Journal of Arid Land  2023, Vol. 15 Issue (6): 667-694    DOI: 10.1007/s40333-023-0014-7
Research article     
Do aeolian deposits and sand encroachment intensity shape patterns of vegetation diversity and plant functional traits in desert pavements?
M'hammed BOUALLALA1, Souad NEFFAR2,3, Lyès BRADAI4, Haroun CHENCHOUNI5,6,*()
1Laboratory of Saharan Natural Resources, Faculty of Sciences and Technology, University of Ahmed Draia, Adrar 01000, Algeria
2Department of Nature and Life Sciences, Faculty of Exact Sciences and Nature and Life Sciences, University of Tebessa, Tebessa 12002, Algeria
3Laboratory Water and Environment, University of Tebessa, Tebessa 12002, Algeria
4Laboratory of Saharan Bioresources Preservation and Valorization, Faculty of Nature and Life Sciences, University of Kasdi Merbah, Ouargla 30000, Algeria
5Department of Forest Management, Higher National School of Forests, Khenchela 40000, Algeria
6Laboratory of Natural Resources and Management of Sensitive Environments, University of Oum-El-Bouaghi, Oum-El- Bouaghi 04000, Algeria
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Abstract  

The effects of sand encroachment on composition, diversity, and functional patterns of vegetation in drylands are rarely studied, and yet addressing these aspects is important to deepen our understanding of the biodiversity conservation. This study aimed to investigate the effect of sand encroachment on plant functional biodiversity of desert pavements (gravel deserts) in the Sahara Desert of Algeria. Plants were sampled and analyzed in three desert pavements with different levels of sand encroachment (LSE) and quantity of aeolian deposits (low, LLSE; medium, MLSE; and high, HLSE). Within the sample-plot area (100 m2), density of every plant species was identified and total vegetation cover was determined. Plant taxonomic and functional diversity were analyzed and compared between LSE. Result showed that 19 plant species in desert pavements were classified into 18 genera and 13 families. Asteraceae and Poaceae were the most important families. The species Anabasis articulata (Forssk) Moq. characterized LLSE desert pavements with 11 species, whereas Thymelaea microphylla Coss. & Durieu ex Meisn. and Calobota saharae (C&D) Boatwr. & van Wyk were dominant species of desert pavements with MLSE (14 species) and HLSE (10 species), respectively. The highest values of species richness and biodiversity were recorded in desert pavements with MLSE, while low values of these ecological parameters were obtained in desert pavements with HLSE. Desert pavements with LLSE were characterized with the highest values of species abundances. Plant communities were dominated by chamaephytes, anemochorous, arido-active, and competitive stress-tolerant plants. The increase in LSE along the gradient from LLSE to HLSE induced significant changes in plant community variables including decreases in plant density, plant rarity, lifeform composition, morphological type, and aridity adaptation. Desert pavements with HLSE favor the degradation of vegetation and trigger biodiversity erosion.

Key words： desert pavements      hot and arid rangeland      plant diversity      land degradation      sand encroachment      plant functional trait      Sahara Desert     
Received: 25 December 2022      Published: 30 June 2023
Corresponding Authors: * Haroun CHENCHOUNI (E-mail: chenchouni@gmail.com; chenchouni.haroun@ensf.dz)
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M'hammed BOUALLALA
Souad NEFFAR
Lyès BRADAI
Haroun CHENCHOUNI
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M'hammed BOUALLALA, Souad NEFFAR, Lyès BRADAI, Haroun CHENCHOUNI. Do aeolian deposits and sand encroachment intensity shape patterns of vegetation diversity and plant functional traits in desert pavements?. Journal of Arid Land, 2023, 15(6): 667-694.

URL:

http://jal.xjegi.com/10.1007/s40333-023-0014-7     OR     http://jal.xjegi.com/Y2023/V15/I6/667

Fig. 1 Geographic location and elevation (a1 and a2) of the El-Guerrara region (Ghardaia Province, Algeria), and monthly meteorological data for mean temperature, precipitation, and potential evapotranspiration (PET; b). Bars are standard errors.
Family Species Level of sand encroachment (LSE)
Low (LLSE) Moderate (MLSE) High (HLSE) Overall
Amaranthaceae Anabasis articulata (Forssk.) Moq. 29.8±5.1 [149] (100%) 10.0±9.5 [50] (100%) 5.0±2.5 [25] (100%) 14.9±12.6 [224] (100%)
Apiaceae Deverra scoparia subsp. scoparia Coss. & Durieu - - 0.2±0.4 [1] (20%) 0.1±0.3 [1] (7%)
Asteraceae Mecomischus halimifolius (Munby) Hochr. 1.0±1.4 [5] (40%) 1.2±1.1 [6] (60%) 2.6±2.1 [13] (80%) 1.6±1.6 [24] (60%)
Asteraceae Pulicaria undulata subsp. undulata (L.) C.A. Mey. 0.6±0.9 [3] (40%) 0.6±0.5 [3] (60%) - 0.4±0.6 [6] (33%)
Asteraceae Rhanterium suaveolens Desf. - 0.2±0.4 [1] (20%) 0.6±0.9 [3] (40%) 0.3±0.6 [4] (20%)
Boraginaceae Moltkiopsis ciliata (Forssk.) I.M. Johnst. 2.2±3.3 [11] (60%) 0.6±0.5 [3] (60%) - 0.9±2.1 [14] (40%)
Brassicaceae Moricandia nitens (Viv.) E.A. Durand & Barratte 0.2±0.4 [1] (20%) 2.8±2.6 [14] (60%) - 1.0±1.9 [15] (27%)
Caryophylla-
ceae		 Polycarpaea repens (Forssk.) Asch. & Schweinf. 2.8±2.6 [14] (60%) 1.6±2.3 [8] (40%) - 1.5±2.2 [22] (33%)
Cistaceae Helianthemum lippii (L.) Dum. Cours. - 0.2±0.4 [1] (20%) 0.6±0.9 [3] (40%) 0.3±0.6 [4] (20%)
Ephedraceae Ephedra alata subsp. alenda (Stapf) Trab. - - 1.6±2.1 [8] (60%) 0.5±1.4 [8] (20%)
Euphorbia-
ceae		 Euphorbia guyoniana Boiss. & Reut. - 2.0±1.0 [10] (100%) - 0.7±1.1 [10] (33%)
Fabaceae Calobota saharae (C&D) Boatwr. & van Wyk - - 14.2±5.2 [71] (100%) 4.7±7.5 [71] (33%)
Fabaceae Retama raetam (Forssk.) Webb - 0.2±0.4 [1] (20%) - 0.1±0.3 [1] (7%)
Poaceae Centropodia forsskaolii (Vahl) Cope 1.4±3.1 [7] (20%) 0.8±1.3 [4] (40%) 2.0±3.5 [10] (40%) 1.4±2.6 [21] (33%)
Poaceae Stipagrostis acutiflora (Trin. & Rupr.) De Winter 0.4±0.9 [2] (20%) - - 0.1±0.5 [2] (7%)
Poaceae Stipagrostis pungens (Desf.) De Winter - - 0.2±0.4 [1] (20%) 0.1±0.3 [1] (7%)
Scrophularia-
ceae		 Kickxia aegyptiaca (L.) Nábelek 2.0±2.9 [10] (60%) 0.2±0.4 [1] (20%) - 0.7±1.8 [11] (27%)
Scrophularia-
ceae		 Scrophularia syriaca Benth. A. DC. 2.4±1.3 [12] (100%) 3.6±1.7 [18] (100%) - 2.0±1.9 [30] (67%)
Thymelaea-
ceae		 Thymelaea microphylla Coss. & Durieu ex Meisn. 2.0±2.1 [10] (60%) 19.4±11.3 [97] (100%) 3.0±1.6 [15] (100%) 8.1±10.3 [122] (87%)
Table 1 Taxonomic list, density, and occurrence frequency of plant species sampled at desert pavements under different levels of sand encroachment in the Sahara Desert of Algeria
Family No. of
genus		 Level of sand encroachment (LSE)
Low (LLSE) Moderate (MLSE) High (HLSE) Overall
S (RF) N' (RF) S (RF) N' (RF) S (RF) N' (RF) S (RF) N' (RF)
Amaranthaceae 1 1 (9.1%) 149 (66.5%) 1 (7.1%) 50 (23.0%) 1 (10.0%) 25 (16.7%) 1 (5.3%) 224 (37.9%)
Apiaceae 1 - - - - 1 (10.0%) 1 (0.7%) 1 (5.3%) 1 (0.2%)
Asteraceae 3 2 (18.2%) 8 (3.6%) 3 (21.4%) 10 (4.6%) 2 (20.0%) 16 (10.7%) 3 (15.7%) 34 (5.8%)
Boraginaceae 1 1 (9.1%) 11 (4.9%) 1 (7.1%) 3 (1.4%) - - 1 (5.3%) 14 (2.4%)
Brassicaceae 1 1 (9.1%) 1 (0.4%) 1 (7.1%) 14 (6.5%) - - 1 (5.3%) 15 (2.5%)
Caryophyllaceae 1 1 (9.1%) 14 (6.3%) 1 (7.1%) 8 (3.7%) - - 1 (5.3%) 22 (3.7%)
Cistaceae 1 - - 1 (7.1%) 1 (0.5%) 1 (10.0%) 3 (2.0%) 1 (5.3%) 4 (0.7%)
Ephedraceae 1 - - - - 1 (10.0%) 8 (5.3%) 1 (5.3%) 8 (1.4%)
Euphorbiaceae 1 - - 1 (7.1%) 10 (4.6%) - - 1 (5.3%) 10 (1.7%)
Fabaceae 2 - - 1 (7.1%) 1 (0.5%) 1 (10.0%) 71 (47.3%) 2 (10.4%) 72 (12.2%)
Poaceae 2 2 (18.2%) 9 (4.0%) 1 (7.1%) 4 (1.8%) 2 (20.0%) 11 (7.3%) 3 (15.7%) 24 (4.1%)
Scrophulariaceae 2 2 (18.2%) 22 (9.8%) 2 (14.3%) 19 (8.8%) - - 2 (10.5%) 41 (6.9%)
Thymelaeaceae 1 1 (9.1%) 10 (4.5%) 1 (7.1%) 97 (44.7%) 1 (10.0%) 15 (10.0%) 1 (5.3%) 122 (20.6%)
Total 18 11 (100.0%) 224 (100.0%) 14 (100.0%) 217 (100.0%) 10 (100.0%) 150 (100.0%) 19 (100.0%) 591 (100.0%)
Table 2 Species richness (S) and density (N') with relative frequencies (RF) of plant families at desert pavements under different levels of sand encroachment in the Sahara Desert of Algeria
Fig. 2 Partitioning of plant species richness (S) recorded at desert pavements in the Sahara Desert of Algeria using nested Venn diagrams. Values of S reported between round brackets represent the total number of species recorded at each level of sand encroachment. Number of species shared exclusively between habitats with different levels of sand encroachment (LSE) and sampling sites are designated between square brackets and within the five-set diagrams, respectively.
Fig. 3 Observed and estimated plant species richness at desert pavements under different levels of sand encroachment in the Sahara Desert of Algeria. Species richness was estimated using first- and second-order Jackknife's (SJack1 and SJack2) and Chao (SChao1 and SChao2) estimators. Vertical bars represent standard deviations. Sest, estimated species richness.
Fig. 4 Sample-based rarefaction (solid line) and extrapolation (dashed line) curves of estimated plant species richness at desert pavements under different levels of sand encroachmentin (LSE) in the Sahara Desert of Algeria. Black solid circles indicate species richness estimated based on reference samples, whereas white solid circles refer to extrapolation to 150 samples. Dark colored areas indicate standard deviations, and light colored areas represent lower and upper bounds of 95% confidence intervals for the estimated values. (a), low LSE (LLSE); (b), moderate LSE (MLSE); (c), high LSE (HLSE); (d), overall.
Fig. 5 Variation of diversity parameters at desert pavements under different levels of sand encroachment in the Sahara Desert of Algeria. Boxes indicate the IQR (interquartile range, 75th to 25th of the data). The median value is shown as a line within the box. White circle is shown as mean. Outlier is shown as black circle. Whiskers extend to the most extreme value within 1.5×IQR. (a), vegetation cover (VC); (b), plant abundance; (c), species richness (S); (d), individual cover (VC:N° ratio), N°, density; (e), species cover (VC:S ratio); (f), species abundance (N°:S ratio); (g), Shannon's diversity index; (h), maximum Shannon's diversity index (H°max); (i), Evenness index; (j), Simpson's reciprocal index (SRI); (j), SRI:S ratio; (k), H°:SRI ratio. The abbreavitaions are the same in the following figures and tables.
Similarity estimator Levels of sand encroachment (LSE)
Low (LLSE) (S=11)
Moderate (MLSE) (S=14)		 Low (LLSE) (S=11)
High (HLSE) (S=10)		 Moderate (MLSE) (S=14)
High (HLSE) (S=10)
Shared species observed 10.0 4.0 6.0
Estimated Chao-shared 10.7 4.0 7.6
Classic Jaccard index (%) 66.7 23.5 33.3
Classic Sørensen index (%) 80.0 38.1 50.0
Raw Chao-Jaccard index (%) 93.2 37.2 39.4
Estimated Chao-Jaccard index (%) 97.2 37.2 42.3
Raw Chao-Sørensen index (%) 96.5 54.2 56.5
Estimated Chao-Sørensen index (%) 98.6 54.2 59.4
Morisita-Horn index (%) 50.3 32.5 31.8
Bray-Curtis index (%) 44.0 25.1 28.3
Table 3 Qualitative and abundance-based similarity scores between plant communities at desert pavements under different levels of sand encroachment in the Sahara Desert of Algeria
Fig. 6 Correlation matrices exposing correlations between plant characteristics (vegetation cover, plant abundance, and diversity parameters) at desert pavements under different levels of sand encroachment (LSE) in the Sahara Desert of Algeria. (a), low LSE (LLSE); (b), moderate LSE (MLSE); (c), high LSE (HLSE).
Plant species Lifeform Morp type Dispersal
type		 Noy-Meir's
type		 Grime's strategy Phyt
type		 Rarity
status		 Ecology
Anabasis articulata Cham Perennial Anemo Arido-active CS SA C Psa, Sax
Centropodia forsskaolii Hemi Perennial Anemo Arido-passive CS SA CC Psa
Ephedra alata Phan Perennial Anemo Arido-active Str SA CC Psa
Euphorbia guyoniana Hemi Perennial Baro Arido-passive Str S° CC Psa
Calobota saharae Cham Perennial Baro Arido-active CS S° AR Psa
Helianthemum lippii Cham Perennial Anemo Arido-passive CSR M-SA CCC Sax, gyps-sax
Kickxia aegyptiaca Cham Perennial Anemo Arido-active CS M-SA C Sax, gyps-sax
Mecomischus halimifolius Cham Perennial Anemo Arido-active CS End RR Psa
Moltkiopsis ciliata Cham Perennial Anemo Arido-active CS SA C Psa
Moricandia nitens Cham Perennial Baro Arido-passive Str S° AR Rud, gyps
Deverra scoparia Cham Perennial Zoo Arido-active Str M-SA C Sax
Polycarpaea repens Ther Annual Anemo Arido-passive SR SA RR Psa
Pulicaria undulata Cham Perennial Anemo Arido-active Str SA AC Gypv
Retama raetam Phan Perennial Baro Arido-active Str M-SA C Psa
Rhanterium suaveolens Cham Perennial Zoo Arido-active CS SA R Sax, gyps-sax
Scrophularia syriaca Cham Perennial Baro Arido-passive Str M-SA CC Psa
Stipagrostis acutiflora Hemi Perennial Anemo Arido-active CS SA R Psa
Stipagrostis pungens Hemi Perennial Anemo Arido-active CS SA CC Psa
Thymelaea microphylla Cham Perennial Baro Arido-active Str M-SA CC Sax, gyps-sax
Table 4 Lfeform, morphological (morp) type, dispersal type, Noy-Meir's category, Grime's strategy, phytogeographic (phyt) type, rarity status, and ecology of plant species at desert pavements under different levels of sand encroachment in the Sahara Desert of Algeria
Fig. 7 Species- and abundance-based biological spectra of plant functional traits (Raunkiaer's lifeform (a), morphological type (b), dispersal type (c), Noy-Meir's strategy (d), Grime's strategy (e), and phytogeographic type (f)) characterizing the vegetation associated to desert pavements under different levels sand encroachment in the Sahara Desert of Algeria. S, species richness; N', density.
Plant status Levels of sand encroachment (LSE)
Low (LLSE) Moderate (MLSE) High (HLSE) Overall
S (RF) N' (RF) S (RF) N' (RF) S (RF) N' (RF) S (RF) N' (RF)
Ecological category
Psammophyte 12 (48.0%) 200 (50.8%) 12 (48.0%) 100 (26.2%) 12 (48.0%) 128 (65.3%) 12 (48.0%) 428 (44.1%)
Saxicolous 6 (24.0%) 169 (42.9%) 6 (24.0%) 150 (39.4%) 6 (24.0%) 47 (24.0%) 6 (24.0%) 366 (37.7%)
Gypsophyte 5 (20.0%) 21 (5.3%) 5 (20.0%) 114 (29.9%) 5 (20.0%) 21 (10.7%) 5 (20.0%) 156 (16.1%)
Gypsovag 1 (4.0%) 3 (0.8%) 1 (4.0%) 3 (0.8%) 1 (4.0%) - 1 (4.0%) 6 (0.6%)
Ruderal 1 (4.0%) 1 (0.3%) 1 (4.0%) 14 (3.7%) 1 (4.0%) - 1 (4.0%) 15 (1.5%)
Rarity/abundance status
CCC - - 1 (7.1%) 1 (0.5%) 1 (10.0%) 3 (2.0%) 1 (5.3%) 4 (0.7%)
CC 3 (27.3%) 29 (12.9%) 4 (28.6%) 129 (59.4%) 4 (40.0%) 34 (22.7%) 6 (31.6%) 192 (32.5%)
C 3 (27.3%) 170 (75.9%) 4 (28.6%) 55 (25.3%) 2 (20.0%) 26 (17.3%) 5 (26.3%) 251 (42.5%)
AC 1 (9.1%) 3 (1.3%) 1 (7.1%) 3 (1.4%) - - 1 (5.3%) 6 (1.0%)
AR 1 (9.1%) 1 (0.4%) 1 (7.1%) 14 (6.5%) 1 (10.0%) 71 (47.3%) 2 (10.5%) 86 (14.6%)
R 1 (9.1%) 2 (0.9%) 1 (7.1%) 1 (0.5%) 1 (10.0%) 3 (2.0%) 2 (10.5%) 6 (1.0%)
RR 2 (18.2%) 19 (8.5%) 2 (14.3%) 14 (6.5%) 1 (10.0%) 13 (8.7%) 2 (10.5%) 46 (7.8%)
Total 11 (100.0%) 224 (100.0%) 14 (100.0%) 217 (100.0%) 10 (100.0%) 150 (100.0%) 19 (100.0%) 591 (100.0%)
Table 5 Species richness (S) and density (N') with relative frequencies (RF) of ecological groups and rarity-abundance status for plants at desert pavements under different sand encroachment levels in the Sahara Desert of Algeria
Fig. 8 Vegetation pattern associated to desert pavements under different levels of sand encroachment in the Sahara Desert of Algeria. L1, low LSE; L2, moderate LSE; L3, high LSE. The vegetation of each sand encroachment level was projected on MFA (multiple factor analysis) dimensions using ten vectors representing different plant characteristics.
Fig. S1 Pearson's correlation matrix between sub-levels of all plant variables clustered in the bi-plot of multiple factorial analysis (MFA). CSR, competition-stress-ruderality; CS, competition-stress; SR, stress-ruderality; Str, stress-tolerant; NA, North African chorological type; M-SA, Mediterranean-Saharo-Arabian; Sah, Saharan chorological type; SA, Saharo-Arabian; CCC, widespread; CC, very common; C, common; AC, fairly common; AR, quite rare; R, rare; RR, very rare; VC, vegetation cover; N°, density; S, species richness; H°, Shannon's diversity index; H°max maximum Shannon's diversity index; E, Evenness index; SRI, Simpson's reciprocal index.
Climatic information Value/class
Latitude 32°46′N
Longitude 04°33′E
Altitude (m) 280
Koeppen's climate classification BWh (B=Arid climate; W=Desert; h=hot)
Budyko's climate Desert
Radiational index of dryness 35.662
Budyko's evaporation (mm/a) 40
Budyko's runoff (mm/a) 0
Budyko's evaporation (%) 100
Budyko's runoff (%) 0
Aridity Hyper-arid
Aridity index 0.02
Moisture index (%) -98
de Martonne's index 1
Precipitation deficit (mm/a) 1639
Climatic NPP (g DM/(m2•a)) 79
NPP (Temperature) 2363
NPP (Precipitation) 79
NPP is precipitation limited
Gorczynski's continentality index 51.7
Table S1 Climatic information of the El-Guerrara region at the Ghardaia Province in the Sahara Desert of Algeria
Parameter Jan Feb Mar Apr May Jun Jul
Mean temperature (°C) 11.1±1.38 13.6±1.20 16.8±1.50 21.0±0.95 25.7±1.63 31.2±1.46 33.7±1.54
Maximum temperature (°C) 17.2±3.66 20.0±3.94 23.2±4.15 28.8±4.31 32.7±4.22 39.4±4.40 42.7±4.64
Minimum temperature (°C) 4.4±3.24 6.6±3.64 9.3±4.29 13.8±4.95 17.7±4.99 23.2±5.80 25.5±5.62
Precipitation (mm) 7±3.44 5±2.11 8±2.10 1±3.85 1±2.62 0±3.16 0±1.20
Potential evapotranspiration (mm) 57±16.38 74±17.90 120±25.69 147±21.64 193±25.24 219±22.40 234±20.27
Water vapor pressure (hPa) 8.2±2.00 8.8±3.15 8.2±3.97 12.3±4.46 13.9±6.43 16.5±8.07 20.5±7.52
Wind speed (km/h) 7.2±4.19 7.9±4.83 7.2±5.33 9.0±5.14 9.0±4.79 8.6±4.61 7.2±3.14
Sunshine frequency (%) 67±9.64 73±7.44 74±9.13 72±9.94 70±4.57 67±4.65 80±1.85
Day length (h) 9.6 12.0 12.0 12.0 14.4 14.2 13.9
Sunshine hours (h) 7.2 7.2 9.6 9.6 9.6 9.6 11.3
Ground frost frequency (%) 13 5 0 0 0 0 0
Effective rainfall (mm) 7 5 8 1 1 0 0
Effective rainfall percentage (%) 99 99 99 100 100 100 100
Number of rainy days (d) 1 1 1 0 0 0 0
Solid precipitation ratio 2 1 0 0 0 0 0
Parameter Aug Sep Oct Nov Dec Average
Mean temperature (°C) 34.0±1.34 28.6±1.29 22.2±1.02 15.8±1.21 11.1±0.95 22.07±1.29
Maximum temperature (°C) 41.7±4.39 37.2±3.92 30.5±4.34 23.2±3.86 18.2±3.51 29.6±4.11
Minimum temperature (°C) 25.0±5.20 22.2±4.81 16.1±4.59 10.0±3.73 6.0±3.28 15.0±4.51
Precipitation (mm) 0±0.86 4±2.45 3±2.53 6±3.06 5±6.30 3±5.31
Potential evapotranspiration (mm) 228±20.63 172±15.29 125±18.01 62±15.80 47±12.78 140±19.34
Water vapor pressure (hPa) 21.8±8.21 19.7±5.11 15.3±2.66 13.7±3.25 9.6±1.97 14.0±4.73
Wind speed (km/h) 7.2±3.05 7.9±2.81 8.6±3.31 7.2±2.77 7.2±3.24 7.9±3.93
Sunshine frequency (%) 78±4.70 73±9.62 71±7.29 66±9.58 67±12.16 72±7.53
Day length (h) 13.2 12.0 12.9 9.6 9.6 12.0
Sunshine hours (h) 10.3 9.6 7.2 7.2 7.2 9.6
Ground frost frequency (%) 0 0 0 0 7 2
Effective rainfall (mm) 0 4 3 6 5 40
Effective rainfall percentage (%) 100 99 100 99 99 99
Number of rainy days (d) 0 0 0 1 1 5
Solid precipitation ratio 0 0 0 1 2 1
Table S2 Long-term monthly climatic data near the El-Guerrara region at the Ghardaia Province in the Sahara Desert of Algeria
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