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Journal of Arid Land  2020, Vol. 12 Issue (6): 1071-1082    DOI: 10.1007/s40333-020-0023-8
Research article     
Germination strategies of annual and short-lived perennial species in the Arabian Desert
Arvind BHATT1, David J GALLACHER2,*(), Paulo R M SOUZA-FILHO3
1Lushan Botanical Garden, Chinese Academy of Sciences, Jiujiang 332900, China
2School of Life and Environmental Sciences, The University of Sydney, Narrabri NSW 2390, Australia
3Multidisciplinary Center of Barra, Federal University of Western Bahia, Barra 47100-000, Brazil
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Germination timing is highly regulated in short-lived plant species since it strongly influences recruitment success of vegetation. In deserts, the spatiotemporal distribution of plant-available water is highly episodic and unpredictable, making winter months more favorable for seed germination when other abiotic conditions co-occur. We hypothesized that changes in photoperiod and thermoperiod would impact germination more in seeds that had undergone in situ storage. We assessed 21 annual and short-lived perennial species in the Arabian Desert to find (1) if seeds were dormant at maturity, (2) if in situ seed storage increased germination percentage compared with no storage, (3) if photoperiod and thermoperiod germination requirements were influenced by in situ storage, and (4) if a phylogenetic association in seed germination could be observed. Seeds of each species collected in early 2017 were divided into two batches. One was tested for germination within one week (fresh seeds). The other was stored in situ at the maternal location (stored seeds) until October 2017 and tested for seed germination in the first week of November. Seed germination was conducted in incubators at two thermoperiods (15°C/20°C and 20°C/30°C; 12 h/12 h), and two photoperiods (12 and 0 h light per day). Results indicated that seed germination percentages of 13 species were significantly enhanced by in situ storage. A thermoperiod response was exhibited by stored, but not fresh seeds. Light exposure increased germination of fresh seeds but had only a minimal effect on stored seeds. Germination traits exhibited no phylogenetic correlation. This result indicated that selection pressure for germination strategy was stronger than that for taxonomic traits of these desert species.

Key wordsdesert species      dormancy      germination      light      temperature      phylogeny     
Received: 09 March 2020      Published: 10 November 2020
Corresponding Authors: J GALLACHER David     E-mail:
About author: *David J GALLACHER (E-mail:
Cite this article:

Arvind BHATT, David J GALLACHER, Paulo R M SOUZA-FILHO. Germination strategies of annual and short-lived perennial species in the Arabian Desert. Journal of Arid Land, 2020, 12(6): 1071-1082.

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Family/species Life form Mohth Location Latitude Longitude
Suaeda vermiculata Forssk. Ex J.F.Gmel. Perennial April Coast 28°54′19″N 48°13′22″E
Calendula arvensis L. Annual April Peri-urban 29°09′47″N 47°41′28″E
Moltkiopsis ciliata (Forsk.) I. M. Johnst. Perennial May Peri-urban 29°09′31″N 47°38′09″E
Horwoodia dicksoniae Turrill Annual May Peri-urban 29°09′29″N 47°38′11″E
Savignya parviflora (Delile) Webb Annual March Peri-urban 29°09′46″N 47°41′28″E
Brassica tournefortii Gouan Annual April Peri-urban 29°09′41″N 47°41′35″E
Gypsophila capillaris (Forsk.) C. Chr. Annual May Peri-urban 29°09′44″N 47°41′26″E
Silene arabica Boiss. Annual March Peri-urban 29°09′47″N 47°41′28″E
Silene villosa Forsk. Annual May Peri-urban 29°09′35″N 47°41′32″E
Helianthemum lippii (L.) Dum. Courset Perennial April Peri-urban 29°09′51″N 47°41′43″E
Asphodelus tenuifolius Cav. Annual March Peri-urban 29°09′45''N 47°41′34″E
Malva parviflora L. Annual May Peri-urban 29°09′34″N 47°41'34″E
Eragrostis curvula (Schrad.) Nees Perennial May Inland 29°09′26″N 47°09′26″E
Panicum acuminatum Sw. Perennial May Peri-urban 29°09′46″N 47°36′28″E
Polypogon monspeliensis (L.) Desf. Annual May Peri-urban 29°09′46″N 47°36′37″E
Digitaria sanguinalis (L.) Scop. Annual May Peri-urban 29°09′51″N 47°41′32″E
Eragrostis barrelieri Daveau Annual May Coast 28°56′23″N 48°11′35″E
Rumex vesicarius L. Annual May Peri-urban 29°09′50″N 47°41′32″E
Reseda arabica Boiss. Annual March Peri-urban 29°09′46″N 47°41′28″E
Haplophyllum tuberculatum (Forsk.) Ad. Juss. Perennial May Coast 28°52′31″N 48°17′30″E
Peganum harmala L. Perennial June Peri-urban 29°09′50″N 47°41′05″E
Table 1 Species collected, and date and location of seeds collection
Factor Treatment Estimate Standard error P
Intercept Fresh, dark, 15°C/20°C -6.213 1.129 <0.001
Storage Stored 5.305 0.887 <0.001
Thermoperiod 20°C/30°C, 12 h/12 h -0.064 0.136 0.638
Photoperiod 12 h light per day 0.660 0.137 <0.001
Interaction Stored×20°C/30°C -0.759 0.172 <0.001
Interaction Stored×Light -0.430 0.172 0.013
Table 2 Coefficients of generalized linear mixed model of in situ storage, thermoperiod, photoperiod and their interactions
Group and species Fresh seeds (%) In situ stored seeds (%)
15°C/20°C 20°C/30°C 15°C/20°C 20°C/30°C
12 h light 0 h light 12 h light 0 h light 12 h light 0 h light 12 h light 0 h light
Group I
P. acuminatum 89±3ab 76±4b 93±2a 83±3ab 95±2a 95±2a 94±2a 95±2a
P. harmala 90±2a 85±3a 61±4b 50±5b 90±2a 83±3a 62±4b 45±4b
S. vermiculata 71±4b 49±4c 82±3a 64±4b 67±4b 64±4b 78±3b 76±3b
Group II
G. capillaris 4±1b 0b 4±1b 0b 42±4a 37±4a 42±4a 38±4a
H. dicksoniae 15±3b 12±2b 28±4ab 24±3ab 62±4a 57±4a 17±3b 15±3b
H. lippi 21±3c 6±1d 28±4c 9±2d 88±2a 64±4b 84±3a 56±4b
H. tuberculatum 7±2c 5±2c 4±2c 3±1c 18±3b 14±3b 45±4a 38±4a
M. ciliata 5±2b 6±2b 10±2b 12±3b 29±4a 31±4a 13±3b 14±3b
M. parviflora 4±2a 5±2a 3±1a 4±2a 12±3a 12±3a 9±2a 8±2a
R. vesicarius 29±4b 10±2d 12±2d 3±1e 57±4a 26±3c 31±4b 10±2d
Group III
R. arabica 0 0 0 0 0 0 0 0
Group IV
A. tenuifolius 0a 0a 0a 0a 0a 0a 7±2a 3±2a
D. sanguinalis 0b 0b 0b 0b 1±1b 1±1b 15±3a 9±3a
E. barrelieri 0a 0a 0a 0a 1±1a 3±2a 1±1a 1±1a
S. parviflora 0a 0a 0a 0a 9±3a 8±3a 0a 0a
Group V
C. arvensis 0c 0c 0c 0c 53±0a 40±5a 9±2b 6±2b
B. tournefortii 0b 0b 0b 0b 5±0b 31±4a 4±1b 28±4a
E. curvula 0c 0c 0c 0c 18±0b 51±5a 1±1c 7±2b
P. monspeliensis 0b 0b 0b 0b 88±0a 78±4a 82±3a 69±4a
S. arabica 0c 0c 0c 0c 75±0a 72±4a 26±4b 22±4b
S. villosa 1±1c 0c 1±1c 0c 75±0a 67±4a 23±4b 17±3b
Table 3 Seed germination percentage for each species, ordered by clustered groups
Fig. 1 Classification of species using clustering of seed germination under different seed storage (fresh and in situ stored seeds) and germination conditions of light (12 and 0 h per day) and temperature (15°C/20°C and 20°C/30°C)
Fig. 2 Number of days to 50% germination of fresh and in situ stored seeds under 15°C/20°C and 20°C/30°C temperatures
Fig. 3 Principal component analysis (PCA) of germination parameters of the species under different seed storage (fresh and in situ stored seeds) and germination conditions of light (12 and 0 h per day) and temperature (15°C/20°C and 20°C/30°C). Group results of plant species can be found in Figure 1.
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