Forest Ecology and Management 195 (2004) 291300

Effects of different pre-sowing seed treatments on germination of 10 Calligonum species

Jun Rena,b,*, Ling Taoa
a

Environmental and Municipal Engineering College, Institute of Environmental Ecology, Lanzhou Jiaotong University, Lanzhou 730070, PR China b Hexi Ecology Institute of Hexi University, Zhangye 734000, Gansu, PR China

Received 4 November 2002; received in revised form 10 September 2003; accepted 30 January 2004

Abstract A greenhouse experiment was conducted to study the effects of abrasion, sulphuric acid, boiling water, cold stratication and seed exudate treatments on the germination of 10 Calligonum species. That are dominant shrubs used for restoration of desert vegetation in mobile sand dunes and stabilized sand elds in the northern desert of China. Little is known about their germination characteristics. In AugustSeptember 1998, seeds of Calligonum were collected and were treated by ve pre-sowing treatments before the germination experiments. The results show that the germination response of seeds to the different pretreatments was more or less similar for all 10 Calligonum species. The germination percent of seed from the 10 species was lowest for exudate treatments and highest for abrasion treatments. The abrasion, sulphuric acid and cold stratication treatments signicantly promoted overall germination. Compared with the control, the exudate treatment signicantly hampered germination, rate of germination and bolstered dormancy for almost all species. The cold stratication treatment can break the dormancy of viable Calligonum seeds and increase the germination, but it has lethal effect on viable seeds probably as well as the boiling water treatments. Almost all germination parameters showed signicant difference between the pre-sowing treatments for all 10 Calligonum species. The speed and percent germination of the Calligonum species can be greatly increased by mechanical scarication or acid treatments. The results show that seeds of C. junceum have good germination potential. These conclusions are very important because Calligonum species can be propagated by seed in the arid desert regions. # 2004 Elsevier B.V. All rights reserved.
Keywords: Calligonum; Pre-sowing treatments; Germination; Dormancy

1. Introduction Calligonum species are dominant perennial shrubs in active sand dunes and stabilized sand elds in the northern desert of China (Mao and Pan, 1986; Tao, 2000). They can exist in mobile sand dunes in conditions of extreme drought (Liu, 19851990; Mao and
Corresponding author. Tel.: 86-931-4938017; fax: 86-931-4938017. E-mail address: renjun30@hotmail.com (J. Ren).
*

Pan, 1986). They have a reputation for high tolerance to water decit. They appear to be suitable for revegetation of desert (Mao et al., 1983; Mao and Pan, 1986; Zhang, 1992; Tao, 2000). They have great potential to provide different products and services such as forage, traditional medicine, halting desert encroachment and stabilizing sand dune (Liu, 1985 1990; Tao, 2000). Because of their importance in providing many uses and services, they have attracted some attention. However, there has been little experimental research dealing with the seed germination and

0378-1127/$ see front matter # 2004 Elsevier B.V. All rights reserved. doi:10.1016/j.foreco.2004.01.046

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seedling emergence of them (Yu and Wang, 1998). The available information about these species is their botany, cultivation method, taxonomy, genetic diversity, brief descriptions of their habitat conditions and the range of their geographical distribution (Mao et al., 1983; Mao, 1984; Liu, 19851990; Mao and Pan, 1986; Zhang, 1992; Yu and Wang, 1998; Tao, 2000; Tao et al., 2000). One of the problems with these species is the difculty of raising seedlings from seeds. The seeds have a hard impermeable testa which prevents imbibition of water and germination (Yu and Wang, 1998; Tao, 2000; Tao et al., 2000). Therefore, the seeds require seed treatments before sowing to obtain rapid, uniform and high germination (Demel, 1996, 1998; Demel and Mulualem, 1996; Schutz and Rave, 1999; Yang et al., 1999; Huang and Gutterman, 2000). The purpose of the present study was to investigate the germination responses of seeds of the 10 Calligonum species to different pre-sowing treatments. An understanding of these factors is crucial for successful regeneration and recruitment of these long-lived desert plant species.

Schrenk, C. arborescens Litv., C. alaschanicum A. Los. and C. potaninii A. Los.) were collected from at least 10 plants per species in August and September 1998 at Shapotou Desert Research and Experimental Station of the Chinese Academy of Sciences (378320 N, 1058020 E, 1339 m a.s.l.), Ningxia Province, China. Seeds were allowed to air-dry and were stored at room temperature (2326 8C) until May 1999, when experimental pretreatments were initiated (storage in this manner did not affect the dormancy or viability of the seeds). For each species, seeds were mixed and then allocated at random. Aborted and predated seeds were discarded. Intact plump seeds were surface sterilized with Na-hypochlorite prior to any experimental usage. Seed viability is variable, but is generally between 30 and 50% (Tao, 2000; Yu and Wang, 1998). 2.2. Pre-sowing seed treatments Abrasion of testa: The mechanical scarication treatment was carried out by grinding seeds in a mortar with a pinch of clean silica sand until all seta of testa are removed and the testa was broken. Sulphuric acid immersion: Although seeds from the 10 species varied apparently in size (Table 1) and degree of hardness of their testas (based on visual inspection), they received the same sulphuric acid treatments. Seeds were immersed in 96% sulphuric acid for 30 min, and then rinsed thoroughly in running water for 45 min. Hot water treatment: Each replicate of seeds was enclosed in a coffee lter bag which was then folded

2. Materials and methods 2.1. Seed collection and preparation Seeds of 10 Calligonum species (C. junceum (Fisch. Et Mey.) Litv., C. leucocladum (Schrenk) Bge., C. rubicundum Bge., C. densum Borszcz., C. mongolicum Turcz., C. chinense A. Los., C. caput-medusae

Table 1 Month and year of seed collection, as well as seed size and dry mass of 10 Calligonum species Species C. C. C. C. C. C. C. C. C. C. junceum leucocladum rubicundum densum mongolicum chinense caput-medusae arborescens alaschanicum potaninii Date of seed collection 24 August 1998 13 August 1998 3 September 1998 3 September 1998 24 August 1998 29 August 1998 10 August 1998 10 August 1998 1 September 1998 1 September 1998 Length of seed (mm) (mean S.D.) 11.07 12.53 16.70 16.09 13.32 13.11 21.66 20.32 19.78 14.73 0.76 0.97 1.62 2.05 1.35 0.90 2.31 4.42 2.44 1.93 Diameter of seed (mm) (mean S.D.) 9.24 9.95 14.17 14.77 9.32 11.85 18.18 16.34 16.27 12.23 0.99 1.81 1.43 1.83 1.48 1.12 3.05 4.01 2.17 2.12 Seed mass (mg) (mean S.D.) 0.08 0.07 0.16 0.10 0.10 0.12 0.15 0.15 0.11 0.11 0.01 0.01 0.02 0.01 0.02 0.01 0.01 0.03 0.02 0.01

Seed size and seed dry mass are mean S.D. of 10 randomly selected seeds from each species in Shapotou Desert Experiment and Research Station of Chinese Academy of Science (Tao, 2000).

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and fastened with paper clips to prevent seed loss. The seeds were then immersed in boiling water for 10 min. After immersion, they were removed from the boiling water and left to cool on a table for about 5 min. Cold stratication: Seeds were soaked in distilled water for 2 days, wrapped in paper bags, and then stored in plastic bags in a refrigerator (3 8C), for 25 days. During the period of imbibition, water was replaced twice a day. Exudate treatment: During preliminary studies it had been noted that a yellow, water-soluble material was exuded from soaking seeds that perhaps included water-soluble inhibitors from the testa (Yu and Wang, 1998). To examine the possible effect of this exudate on seed germination, a group of seeds was exposed to the exudate solution during the germination period, instead of clean distilled water. 2.3. Germination experiment The germination experiments were conducted during MayJune 1999. Controlled seeds and seeds that had undergone the pretreatments described above were placed on wet lter paper in glass Petri dishes (11.5 cm diameter 2 cm depth). These were then placed into temperature-controlled chambers for germination, set at 14 h of daylight at 25 8C and 10 h of darkness at 12 8C, to approximate general springtime eld conditions. To ensure no systematic effects due to position within the chamber, Petri dishes were rearranged at random every 2 days. All pre-sowing seed treatments consisted of ve replicates of 40 seeds for each species. Visible radicle growth was used to dene germination. Germination was recorded every 5 days and allowed to proceed for 9 weeks except where specied. Ungerminated seeds were soaked in water at 30 8C for 24 h. Testa was cut and the embri was soaked in 1% tetrazolium chloride for 24 h at 30 8C. Pink embryos were scored as alive. Germination was expressed as percentage of viable seeds germinated. 2.4. Data analysis The results of the germination experiments were analyzed for statistical signicance (ANOVA) from the STATISTICA software package for personal computer (Statsoft, 1993). All percent germination data

were arcsine-square-root transformed prior to analysis. Arcsine-transformed means and standard errors were backtransformed for graphic presentation. Multiple comparisons of means were made with Duncans tests at 95%. The following parameters were determined: number of germinating seeds germination 100 number of seeds initiated relative germination number of germinating seeds 100 number of viable seeds initiated dormancy number of ungerminated but viable seeds 100 number of seeds initiated relative dormancy number of ungerminated but viable seeds 100 number of viable seeds initiated number of inviable seeds mortality 100 number of seeds initiated XG index of germination rate : IGS t where G is the relative germination percentage at 5-day intervals, and t is the total germination period. This parameter characterizes the rate of germination for particular seed replicates.

3. Results 3.1. Seed quality of germination ecology Table 2 summarizes mean values for germination, relative germination, dormancy, relative dormancy, mortality and IGS, prior to any pretreatment. It was clear that there was no signicant correlation between these variables and seed size or weight (Tables 1 and 2). Germination and relative germination differed among the 10 species (one-way ANOVA: F9;36 14:94, P < 0:0001 for the former; F9;36 10:52, P < 0:0001 for the latter). C. leucocladum had the highest germination and relative germination (47 and 69.3%). C. leucocladum and C. mongolicum had signicant poorer germination and relative germination (11.5 and 28.8% for the former, 14.5 and 31.3% for the latter) than the other species. Inversely,

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Table 2 Overall seed germination qualities for 10 species of Calligonum following storage for 8 months at room temperature in Shapotou Desert Experiment and Research Station of Chinese Academy of Science Species C. C. C. C. C. C. C. C. C. C. junceum leucocladum rubicundum densum mongolicum chinense caput-medusae arborescens alaschanicum potaninii Germination percent S.D. 47.0 11.5 37.5 42.5 14.5 35.5 34.0 29.5 26.0 34.0 7.4 5.2 7.7 3.1 8.2 4.1 8.4 4.5 8.2 4.5 a b ac ac b ac ac ac c ac Relative germination percent S.D. 69.3 28.8 58.9 64.1 31.3 61.3 52.2 58.5 58.2 64.2 13.4 a 8.1 b 14.7 a 9.6 a 8.7 b 6.2 a 12.9 a 2.9 a 12.6 a 7.1 a Dormancy percent S.D. 21.5 27.5 27.0 24.5 30.0 22.5 31.5 21.0 19.5 19.0 10.2 a 7.7 a 11.0 a 8.7 a 10.3 a 4.7 a 9.9 a 3.8 a 8.9 a 4.2 a Relative dormancy percent S.D. 30.7 71.2 41.1 35.9 68.7 38.7 47.8 41.5 41.8 35.8 13.4 a 8.1 b 14.7 a 9.6 a 8.7 b 6.2 a 12.9 a 2.9 a 12.6 a 7.1 a Mortality percent S.D. 31.5 61.0 35.5 33.0 55.5 42.0 34.5 49.5 54.5 47.0 6.8 a 11.0 b 6.9 a 6.5 a 16.6 b 5.1 ab 9.7 a 7.8 ab 13.9 b 4.5 ab IGS S.D. 9.8 4.2 7.6 8.6 4.5 7.6 6.4 7.0 7.4 8.0 2.4 1.5 2.7 1.5 0.8 1.0 1.9 1.0 2.3 1.5 a bd abc a cd ac ab abc a ac

Different letters indicate signicant difference between the values of pairs of species in the same column (Duncans multiple comparison test, P < 0:05).

C. leucocladum and C. mongolicum had highest relative dormancy (71.2 and 68.7%). The one-way ANOVA showed that the percentage of dormant seeds was not signicantly affected by species (F9;36 2:12, P 0:53). There are signicant difference among species for seed mortality and IGS (oneway ANOVA: F9;36 7:70, P < 0:0001 for mortality; F9;36 6:87, P < 0:0001 for IGS). C. leucocladum, C. mongolicum and C. alaschanicum had signicantly greater mortality (61, 55.5 and 54.5%). The IGS were lowest for C. leucocladum (4.2) and C. mongolicum (4.5; Table 2). 3.2. Effect of pretreatment on germination For all 10 species, germination differed signicantly for seeds given the different pretreatments (one-way ANOVA: F5;20 35:57, P < 0:0001 for C. junceum; F5;20 30:40, P < 0:0001 for C. leucocladum; F5;20 38:46, P < 0:0001 for C. rubicundum; F5;20 49:41, P < 0:0001 for C. densum; F5;20 87:17, P < 0:0001 for C. mongolicum; F5;20 29:55, P < 0:0001 for C. chinense; F5;20 49:30, P < 0:0001 for C. caput-medusae; F5;20 65:21, P < 0:0001 for C. arborescens; F5;20 1 3:90, P < 0:0001 for C. alaschanicum; F5;20 48:26, P < 0:0001 for C. potaninii). For most species, the lowest germination percent occurred in the exudate treatment and the highest occurred in the abrasion treatment. Compared with the control, seeds of all species that had undergone the boiling water treatment

had poorer germination, the abrasion, sulphuric acid and cold stratication treatments signicantly increased the germination of all species (Fig. 1). For all species, the relative germination differed signicantly for seeds given different pretreatments (one-way ANOVA: F5;20 5:03, P 0:0038 for C. junceum; F5;20 24:50, P < 0:0001 for C. leucocladum; F5;20 6:85, P 0:0007 for C. rubicundum; F5;20 12:76, P < 0:0001 for C. densum; F5;20 32:22, P < 0:0001 for C. mongolicum; F5;20 3:32, P 0:0240 for C. chinense; F5;20 38:36, P < 0:0001 for C. caput-medusae; F5;20 26:58, P < 0:0001 for C. caput-medusae; F5;20 10:69, P < 0:0001 for C. alaschanicum; F5;20 12:68, P < 0:0001 for C. potaninii). Relative to the control and other treatments, seeds from all species showed an increase in relative germination after sulphuric acid and boiling water treatments. The relative germination was signicantly decreased by exudate treatment for C. leucocladum, C. densum, C. mongolicum, C. caputmedusae, C. arborescens and C. alaschanicum (Fig. 2). There were signicant differences in the percent seed dormancy between the different seed pretreatments in eight species (one-way ANOVA: F5;20 8:03, P < 0:001 for C. junceum; F5;20 16:51, P < 0:0001 for C. leucocladum; F5;20 3:41, P 0:0217 for C. rubicundum; F5;20 6:61, P < 0:001 for C. densum; F5;20 12:97, P < 0:0001 for C. caput-medusae; F5;20 7:09, P < 0:001 for C. arborescens; F5;20 19:19, P < 0:0001 for C. alaschanicum; F5;20 7:48,

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Fig. 1. Seed germination percent of 10 Calligonum species after ve seed pretreatments and one control. Values with the same superscript letters are not signicantly different at P < 0:05 according to Duncans multiple comparison test.

P < 0:001 for C. potaninii) and no signicant difference in three species (F5;20 2:37, P 0:0767 for C. mongolicum; F5;20 1:44, P 0:2533 for C. chinense). The percent dormancy responses of seeds to the yellow water-soluble exudate differed in between species. In the case of C. potaninii, the exudate treatment had no signicant effects on dormancy (Fig. 3). For C. leucocladum, C. densum, C. caputmedusae, C. arborescens and C. alaschanicum, the exudate treatment signicantly increased dormancy relative to the control and all other pretreatments. For C. leucocladum, C. densum, C. caput-medusae, C.

arborescens and C. alaschanicum, the exudate treatment signicantly increased dormancy relative to the control and all other pretreatments (Fig. 3). Relative dormancy was signicantly different between pretreatments in 10 species (one-way ANOVA: F5;20 11:24, P < 0:0001 for C. junceum; F5;20 29:66, P < 0:0001 for C. leucocladum; F5;20 12:4329, P < 0:0001 for C. rubicundum; F5;20 17:89, P < 0:0001 for C. densum; F5;20 18:89, P < 0:0001 for C. mongolicum; F5;20 4:64, P 0:0056 for C. chinense; F5;20 41:24, P < 0:0001 for C. caput-medusae; F5;20 30:87,

Fig. 2. Seed relative germination percent of 10 Calligonum species after ve seed pretreatments and one control. Values with the same superscript letters are not signicantly different at P < 0:05 according to Duncans multiple comparison test.

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Fig. 3. Seed dormancy percent of 10 Calligonum species after ve seed pretreatments and one control. Values with the same superscript letters are not signicantly different at P < 0:05 according to Duncans multiple comparison test.

P < 0:0001 for C. caput-medusae; F5;20 17:65, P < 0:0001 for C. alaschanicum; F5;20 12:87, P < 0:0001 for C. potaninii). Compared with the control, the abrasion treatment signicantly lowed overall dormancy except for C. chinense and C. potaninii. C. potaninii had signicantly lower relative dormancy than the control and other treatments following cold stratication. Cold stratication treatment signicantly reduced relative dormancy in C. leucocladum, C. rubicum, C. mongolicum, C. caput-medu-

sae, C. arborescens, C. potaninii relative to the control (Fig. 4). There were signicant differences in percent seed mortality between the different seed pretreatments in 10 species (one-way ANOVA: F5;20 24:92, P < 0:0001 for C. junceum; F5;20 6:41, P 0:0011 for C. leucocladum; F5;20 19:87, P < 0:0001 for C. rubicundum; F5;20 7:84, P 0:0003 for C. densum; F5;20 3:02, P 0:0342 for C. mongolicum; F5;20 27:96, P < 0:0001 for C. chinense; F5;20 5:84,

Fig. 4. Seed relative dormancy percent of 10 Calligonum species after ve seed pretreatments and one control. Values with the same superscript letters are not signicantly different at P < 0:05 according to Duncans multiple comparison test.

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Fig. 5. Seed mortality percent of 10 Calligonum species after ve seed pretreatments and one control. Values with the same superscript letters are not signicantly different at P < 0:05 according to Duncans multiple comparison test.

P 0:0017 for C. caput-medusae; F5;20 8:39, P 0:0002 for C. arborescens; F5;20 22:45, P < 0:0001 for C. alaschanicum; F5;20 36:82, P < 0:001 for C. potaninii). For C. junceum and C. rubicundum, the exudate treatments applied resulted in signicant effects on mortality than the control and other treatments (Fig. 5). Compared with the control, the abrasion treatment signicantly lowed overall mortality except for C. leucocladum and C. mongolicum. Cold stratication tended to increase mortality, but such increases were not always signicant (Fig. 5). The index of germination of seeds (IGS) differed signicantly between pretreatments for all species (oneway ANOVA: F5;20 5:61, P 0:0022 for C. jun-

ceum; F5;20 39:56, P < 0:0001 for C. leucocladum; F5;20 8:07, P 0:0003 for C. rubicundum; F5;20 22:49, P < 0:0001 for C. densum; F5;20 59:6, P < 0:0001 for C. mongolicum; F5;20 8:8, P 0 :0002 for C. chinense; F5;20 44:4, P < 0:0001 for C. caput-medusae; F5;20 57:73, P < 0:0001 for C. arborescens; F5;20 14:88, P < 0:0001 for C. alaschanicum; F5;20 21:26, P < 0:0001 for C. potaninii). For all species, highest IGS occurred following abrasion, sulphuric acid and boiling treatments. Compared with the control, the cold stratication and exudate treatments signicantly decreased the IGS in C. leucocladum, C. mongolicum, C. arborescens and C. potaninii, the abrasion, sulphuric acid and boiling

Fig. 6. Index of germination rate of seed (IGS) for 10 Calligonum species after ve seed pretreatments and one control. Values with the same superscript letters are not signicantly different at P < 0:05 according to Duncans multiple comparison test.

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water treatments signicantly increased the IGS in C. leucocladum and C. mongolicum (Fig. 6).

4. Discussion The hard testa of many desert plants has evolved to withstand unfavourable conditions such as heat caused by sunlight, the strong teeth of dispersing animals, severe drought and mechanical damage (Freas and Kemp, 1983; Washitani and Masuda, 1990; Fenner, 1991; Meyer and Monsen, 1991; Gutterman, 1993; Baskin and Baskin, 1998; Silvertown, 1999). In consequence, severe treatments are required to chance the testa permeability to water. Several pre-sowing treatments have been used to overcome hard testa imposed dormancy (Meyer et al., 1990; Ferasol et al., 1995; Broncano et al., 1998; Goslan and Gutterman, 1999; Rachel and Galatowitsch, 1999). The objective of all these treatments is to make the testa permeable to water by acting on specic weak spots of the hard testa. Of the different pre-sowing treatments, cold stratication, mechanical, acid and hot water treatments are widely used because they improve germination within a relatively short period of time. In the present study, the germination response of seeds to different pretreatments was more or less similar for all 10 Calligonum species. Germination of seeds of the 10 species was poorest with the exudate treatment and best with the abrasion treatment. Compared with the control, seeds given boiling water treatment had poorer germination, and the abrasion, sulphuric acid and cold stratication treatments signicantly improved germination. Exudates reduced seed germination except for C. alaschanicum. Seeds from all species showed an increase in relative germination after sulphuric acid and boiling water treatments relative to the control and other treatments. The exudate treatment had different effects on dormancy depending on species. Compared with the control, the exudate treatment signicantly reduced percent germination and increased dormancy for almost all species Germination inhibiting substance have been reported previously in the yellow water-soluble exudate of Calligonum seeds (Vleeshouwers et al., 1995; Schutz and Milberg, 1997; Yu and Wang, 1998). The abrasion treatment enhanced germination of all species. The cold stratication treatment can break the

dormancy of viable Calligonum seeds and increased the germination, but it has a little lethal effect on viable seeds (Fig. 5). According to our observation during germination experiments, many seeds treated with boiling water rotted and succumbed to mold attacked, indicating that they are very sensitive to high temperature. The positive responses of seeds to the pre-sowing scarication treatments (abrasion and sulphuric acid) suggests that the hard testa is responsible for the low percentage germination of untreated seeds by preventing imbibition of water. Prevention of germination by hard testa of Calligonum species has different ecological advantages (Tao, 2000). This feature favours the accumulation of persistent seed banks in the soil, spreads germination over time, suffers the extremely environmental condition and increases the chance that some seeds will germinate, survive and establish (Freas and Kemp, 1983; Fenner, 1991; Gutterman, 1993; Morgan, 1998; Wang et al., 1998). Overcoming dormancy, softening of the testa and water uptake are therefore, crucial matters in the life cycle of hardseeded species (Wang et al., 1997; Yang et al., 1999). The results from the present study provides evidence that almost all germination variables differed signicantly between the pre-sowing treatments for all 10 Calligonum species. The speed and percent germination of the Calligonum species investigated can be greatly increased by subjecting the seeds to either mechanical scarication or acid treatments. The results show that seeds of C. junceum have strong ability of germinating (Table 3). Improper seed pre-sowing treatment can lead to reduced seed germination. The implications of poor seed supply for restoration efforts are substantial. As viability rates are reduced, recommended seeding rates must be increased. Because the availability of Calligonum seeds is often limited by climate (rainfall), seasonal and site variability (Tao, 2000) obtaining sufcient seed may be difcult and the associated costs prohibitive. The formal standard recommended seed pre-sowing treatments for Calligonum species are sulphuric acid and cold stratication (Zhang, 1992). The results of the present study indicate that abrasion, cold stratication and sulphuric acid all are appropriate for the 10 Calligonum species under study. These conclusions are very meaningful because they indicated that Calligonum species can be propagated

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Table 3 Overall seed germination qualities for 10 species of Calligonum following storage for 8 months at room temperature after the control and ve pretreatments Species C. C. C. C. C. C. C. C. C. C. junceum leucocladum rubicundum densum mongolicum chinense caput-medusae arborescens alaschanicum potaninii Germination percent S.D. 50.1 13.4 41.2 46.6 19.4 38.7 37.3 33.0 30.3 37.7 14.2*** 9.7*** 13.1*** 12.2*** 11.2*** 11.8*** 13.2*** 12.2*** 12.0*** 12.5*** Relative germination percent S.D. 71.5 36.3 64.0 63.4 38.3 63.8 56.6 58.6 57.4 65.2 9.2** 20.7*** 11.4*** 13.1*** 17.7*** 9.9* 16.6*** 17.8*** 16.0*** 16.0*** Dormancy percent S.D. 17.9 25.8 20.6 24.9 27.8 19.6 25.3 20.3 18.7 17.3 7.6*** 13.0*** 8.1* 10.5*** 11.1 6.0 10.4*** 10.1*** 7.3*** 7.0*** Relative dormancy percent S.D. 16.0 58.6 33.5 34.5 59.4 33.9 40.5 37.8 38.5 32.0 10.2*** 25.8*** 13.0*** 14.7*** 21.2*** 11.2** 18.6*** 20.0*** 18.4*** 16.6*** Mortality percent S.D. 29.7 54.9 36.2 26.6 51.4 39.9 35.1 44.1 48.2 42.8 18.6*** 11.8** 15.4*** 11.8*** 13.4* 14.9*** 10.9** 12.3*** 13.3*** 13.2*** IGS S.D. 10.5 4.9 8.5 8.8 5.1 8.3 7.4 7.6 7.4 8.5 2.1** 3.2*** 2.3*** 2.2*** 2.8*** 1.7*** 2.7*** 2.8*** 2.7*** 2.6***

Statistical differences for each species among ve pretreatments and one control determined by ANOVA (d.f. 5, 20). * P < 0:05. ** P < 0:01. *** P < 0:001.

by seed in the arid desert regions. Further work is required to develop seeding criteria for good seed presowing practice to ensure germination in the eld.

Acknowledgements Financial support by Qing Lan Talent Engineering Funds by Lanzhou Jiaotong University and by The National Key Basic Research Special Foundation Project of China (G2000018602) and Shapotou Desert Experiment and Research Station of Chinese Academy of Science. References
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