Plant Ecology 136: 95103, 1998. c 1998 Kluwer Academic Publishers. Printed in Belgium.

95

Inuence of heat on seed germination of seven Mediterranean Leguminosae species

Jos M. Herranz, Pablo Ferrandis & Juan J. Martnez-S nchez e a
Department Plant Production & Agricultural Technology, E.T.S.I. Agr nomos, University of Castilla-La Mancha, o Campus Universitario s/n, 02071 Albacete, Spain
Received 23 June 1997; accepted in revised form 24 January 1998

Key words: Hardseedness, Heat, Leguminosae, Seed germination

Abstract The inuence of high temperatures (dry heat and hot water) on germination of seven Mediterranean Leguminosae species typical of re-prone ecosystems in southern Spain is analyzed, in order to know the response of seeds to wildres and the possible implications in their regeneration after this disturbance. Seeds were heated to a range of temperatures (50 150 C) and exposure times (160 min) similar to those registered in the upper soil layers during wildres. Germination tests were carried out in plastic Petri dishes over 60 days. In general, the degree of seed germination promotion by dry heat treatments showed a wide interspecic variation, although the nal germination level was increased in all the studied species except for Scorpiurus muricatus. The thermal pretreatment of 50 C, however, was not effective for germination in any species, and rising the temperature to 70 C only slightly enhanced the germination in Cytisus patens. The preheatings of 90 C (5 and 10 min), 120 C (5 and 10 min), and 150 C (1 min) were the most effective in promoting seed germination. Hot water (100 C) scarication also increased the nal germination level in all cases, with the exception of C. patens. The germination rates after preheating were much lower than in mechanically scaried seeds and closely resembled those of the untreated seeds, except for C. reverchonii, whose seed germination rate decreased with heat. The response of species to heat shock had no clear relationship with life trait or with the specic post-re regeneration strategy (obligate seeder or facultative resprouter). Those species coexisting in the same habitats had different heat optimal requirements for seed germination, an strategy suggested by some authors as minimizing interspecic competition in the secondary succession started after re. Introduction Fire has a great inuence in maintaining many plant communities throughout the world, especially in Mediterranean-type ecosystems (Naveh 1975; Trabaud 1983; Trabaud & Lepart 1981; Arianoutsou & Margaris 1982; Mazzoleni & Pizzolongo 1990; Hanes 1971; Purdie 1977), but also in African savannas, pine plain communities in southeastern USA, and heathlands in Europe (Bullock & Webb 1995). After re, most species in these communities are able to regenerate by vegetative regrowth, but there are a few others which regenerate solely from seeds (Trabaud 1987; Casal 1987; T rrega et al. 1992; Martneza S nchez et al. 1996). When plants are killed by re a (obligate seeders), they regenerate by seedling establishment. Where adult plants are capable of resprouting from belowground vegetative buds, no seedling recruitment may be needed to maintain a community after a single re. However, there may be some mortality of adult plants during the re and some senescence in interre periods, so that some successful seedling recruitment will be necesary in order to compensate adult plant losses over a long series of res (Auld & OConnell 1991). Moreover, it must be kept in mind that although sexual reproduction may not be quantitatively the most important way of regeneration with regard to biomass contribution, it can be of great interest since it is the only way of conserving the genetic variation of species which tend to regenerate

96 by sprouting (T rrega et al. 1992). Therefore, it is crua cial to know the factors affecting seed germination of main species in a community, in order to understand the post-re plant dynamics. Although there are many studies that demostrate the inuence of re on seed germination of species of Mediterranean-type ecosystems (Naveh 1974; Arianoutsou & Margaris 1981; Troumbis & Trabaud 1986; Keeley 1987, 1991; Trabaud & Oustric 1989; Thanos & Georghiou 1988; Thanos et al. 1992; Corral et al. 1990; Roy & Sonie 1992; Valbuena et al. 1992; Gonz lez-Rabanal & Casal 1995), those referring to a legumes are scarce (Cushwa et al. 1968; Martin et al. 1975; Cavanagh 1980; Mott et al. 1982; Pereiras et al. 1985; T rrega et al. 1992; Auld & OConnell 1991). a These studies suggest that the high temperatures generated in the re can stimulate germination in species with a hard seedcover, by inducing the breaking of seed coats, thereby facilitating the subsequent embryo imbibition and radicle expansion. In the present work, heat requirements for seed germination of seven Mediterranean Leguminosae species are studied. They are: Cytisus striatus (Hill.) Rothm., Cytisus reverchonii (Degen & Hevier) Bean, Cytisus patens L., Dorycnium pentaphyllum Scop., Argyrolobium zanonii (Turra) P.W. Ball, Scorpiurus muricatus L., and Psoralea bituminosa L. These species are an important component in shrublands established in regularly burnt areas in the southern half of the Iberian Peninsula, and can be frequently found growing side by side, as in the case of C. reverchonii and D. pentaphyllum, as well as P. bituminosa, S. muricatus, D. pentaphyllum, and A. zanonii (Martnez-S nchez a 1994; Martnez-S nchez et al. 1994, 1996). a The three Cytisus species are Iberian endemic medium-large shrubs (1.53 m), which behave as facultative resprouters after re. C. striatus forms mixed shrublands on siliceous substrata with Cistaceae and Ericaceae species in the western half of the Iberian Peninsula, C. patens is a typical plant of humid habitats in the eastern half, whereas C. reverchonii forms practically pure shrublands in the Betic mountain range (south Spain). The remaining studied species show a broad Mediterranean distribution (Tutin et al. 1964 1980). P. bituminosa, S. muricatus, and A. zanonii are obligate seeders, whereas D. pentaphyllum is a facultative resprouter. These four species usually form a dense vegetation during the rst post-re years, not only in Spain but also in other different Mediterranean countries, and show notable declines subsequently (Papavassilious & Arianoutsou 1993; Clemente et al. 1996). This phenomenom has been attributed by these authors to a re-induced seed germination, although until now no test of this hypothesis has been published. This study tries to make up for the lack of information in this eld on Leguminosae species. Besides, it is of great interest to know whether those species which coexist in the same habitats have similar heat requirements for the promotion of seed germination or not. As it has been suggested for Cytisus scoparius (L.) Link and Genista orida L. (T rrega et al. 1992), differena tial requirements for heat could have adaptative value, as such a mechanism could permit germination of each species in different points of the ground, according to the temperature reached with the re. In contrast, uniform heat requirements would cause simultaneous germination of these species and lead to a damaging competition between seedlings. Thus, the aim of this study is: (1) assessing the temperature intervals and dry heat exposure times which promote seed germination for the studied species, as well as the effect of hot-water treatments; (2) knowing the degree of similarity in heat requirements for the promotion of seed germination in those species tending to share the habitat.

Methods Plant material Names of taxa follow the nomenclature of Tutin et al. (19641980). Seed samples were collected from ripe fruits of as many plants as possible in healthy looking wild populations (so that high seed viability was assumed for all species) in several localities in the province of Albacete and Ciudad Real (southern half of Spain): C. striatus: Caba eros National Park, province n of Ciudad Real, M.T.U.: 30SUJ6860, 840 m of altitude, quartzite substratum, in a mixed maquis shrubland with Arbutus unedo L., Erica australis L., E. arborea L., Phillyrea angustifolia L., Cistus ladanifer L., and Halimium ocymoides (Lam.) Willk. The area was last affected by re in August 1970. Date of seed collection: 22 July 1994. C. patens: Ba os de Tus, Yeste, province of n Albacete, M.T.U.: 30SWH5246, 820 m of altitude, siliceous sand substratum, in a mixed maquis shrubland with Pistacia lentiscus L., Quercus coccifera L., Lonicera implexa Aiton, Arbutus unedo, and Phil-

97 lyrea angustifolia. The area was last affected by re in August 1979. Date of seed collection: 26 July 1994. C. reverchonii: Monte Ardal, Yeste, province of Albacete, M.T.U.: 30SWH5646, 1220 m of altitude, calcareous substratum, in a mixed shrubland with Rosmarinus ofcinalis L., Juniperus oxycedrus L., Cistus monspeliensis L., Brachypodium retusum (Pers.) Beauv., and Festuca nevadensis (Hackel) Markg. The area was last burnt in August 1985. Date of seed collection: 26 July 1994. D. pentaphyllum, A. zanonii, P. bituminosa, and S. muricatus: the surrounding area of Moropeche village, Yeste, province of Albacete, M.T.U.: 30SWH5560, 900 m of altitude, siliceous sand substratum, in gaps of a mixed shrubland with Quercus coccifera, Juniperus oxycedrus, Rosmarinus ofcinalis, Phillyrea angustifolia, and Cistus ladanifer. The area was last burnt in August 1989. Date of seed collection: 27 July 1994. The three mentioned localities in Yeste were subsequently affected by a great forest re, which destroyed 13 341 ha between 7 and 19 August 1994. Until February 1996, when germination tests were started, seed samples were stored in paper bags in a 5 C cold room. Temperature treatments Seeds were heated to a range of temperatures similar to those registered during wildres in the upper layer of the soil (depth of 05 cm), where the majority of seed banks are usually concentrated (Roberts 1981; Simpson et al. 1989). Measurements in wildres in the California chaparral (De Bano et al. 1977), in the garrigue of southern France (Trabaud 1979), and in Mediterranean-type ecosystems of southeastern Australia (Floyd 1966; Auld & OConnell 1991; Bradstock & Auld 1995), showed that temperature in the 05 cm deep soil layer usually ranges between 50150 C, with very variable heat exposure periods depending on depth and re intensity. Therefore, it was decided to test the treatment temperatures of 90 C, 120 C, and 150 C, for durations of 1, 5, and 10 min. Moreover, it was considered interesting to test the effect of lower temperatures during longer heat exposition periods: 50 C for 10, 15, 30, and 60 min, and 70 C for 5, 10, 15 and 30 min. Thus, the temperature interval used in this study was similar to those tested in other works on legumes seed germination of the same nature (Cushwa et al. 1968; Mott et al. 1982; Pereiras et al. 1985; Auld & OConnell 1991; T rrega et al. 1992). The prea heating was carried out on dry seeds spread on glass dishes in a mufe furnace and the temperatures were maintained stable within a range of 2 C. There were 4 replicates of 25 seeds for each treatment. Besides, one complete set of 100 untreated seeds was used as control in each species. Another set was submitted to mechanical scarication, that was achieved by abrasion between two pieces of sandpaper. This treatment has a high effectiveness in the removal of physical dormancy imposed by an impermeable hard seedcoat, both in Leguminosae (Pereiras et al. 1987; A orbe et al. 1990; T rrega et al. 1992) and n a in Cistaceae (Thanos & Georghiou 1988; Trabaud & Oustric 1989; Thanos et al. 1992) species (70100% germination level). It gives valuable information on seedlot viability and an important reference to compare the effectiveness of thermal treatments on seed germination promotion. In those species for which there were enough seeds (all of them except D. pentaphyllum and P. bituminosa), an additional treatment with hot water was carried out. This scarication technique has been previously used to reduce hardseedness in legumes (Cushwa et al. 1968), as well as in Cistaceae species with hard coat seed (P rez-Garca et al. 1995). Seeds were immersed e in a glass containig 50 ml hot distilled water (100 C), and then left to cool to room temperature. Treated seeds were kept in the water for 24 h, and subsequently sown and incubated in the same conditions as seeds submitted to the other treatments. Germination conditions Seeds were germinated in plastic Petri dishes (diam. 7 cm) over 2 caps of lter paper. They were kept saturated with distilled water throughout all the treatments, and laid at random on a temperature controlled chamber (model Radiber, AGP-600, 9000 LUX, 0.1 C of precision). Incubation was carried out in photoperiod conditions: fourteen hours in light at 18 C, and ten hours in darkness at 13 C. Radicle emergence was the criterion for deciding if germination had taken place (Come 1970). Recount of the germinated seeds was carried out every two days over a period of 60 days, and germinated seeds were removed from the plastic Petri dishes. Statistical analysis The effect of treatments on seed germination was studied by the analysis of two parameters: (i) the nal cumulative percentage of germination (i.e., germina-

98 tion level), and (ii) the rapidity of germination (i.e., germination rate; Keeley 1987; Thanos & Georghiou 1988; Thanos et al. 1992) expressed by the T 50 parameter (Thanos & Georghiou 1988; Escudero et al. 1997), which can be dened as the time required for manifestation of half of the nal germination level. For each species, germination level and germination rate data were submitted to a oneway ANOVA with treatment (dened by temperature and exposure time, besides mechanical scarication, control, and, in some cases, hot water scarication) as factor. Tukeys test (1949) was used to detect any signicant differences ( = 0:05) in the comparison between the pairs of treatments. Previously the sampling normality was checked by the David-test (1954) and the homogeneity of the variances by the Cochran-test (1941). Data of nal germination (percentages) needed an arcsin transformation. any studied species, irrespective of heat exposure duration, and the 70 C temperature only slightly promoted the germination of C. patens seeds. From this point, increases in temperature caused signicant increases of seed germination in the majority of cases, until a constant level. Preheatings at 90 C for 5 and 10 min enhanced germination in C. striatus, C. reverchonii, A. zanonii, and P. bituminosa. Temperatures of 120 C and 150 C were also very effective in promoting seed germination, although 150 C during exposure times equal and longer than 5 min resulted in a detriemental effect on seed viability of all the studied species. In both C. striatus and C. reverchonii there were several dry heat treatments which promoted germination as much as mechanical and hot water scarication. D. pentaphyllum had the highest heat requirements for promoting germination, and C. striatus showed the highest heat resistance. So, the preheating at 120 C for 10 min on seeds of this species gave a nal germination value of 76 8:48%, and even 2% of seeds were able to germinate after the 150 C for 10 min treatment.

Results The germinability of untreated seeds was generally low, ranging between 615%, with the exception of P. bituminosa whose control seeds reached a nal germination level of 27% (Figure 1). As deduced from the T50 values (Table 1), the germination rate of untreated seeds was considerably slower than that of mechanically scaried seeds, except for A. zannonii and P. bituminosa, and similar to the values recorded for preheated seeds. Only in the case of C. reverchonii the germination rate value of untreated seeds was higher than those recorded in thermal treatments. The mechanical scarication of the seed coat resulted in a pronounced increase of germinability, with nal germination levels including between 77.5 and 93% (Figure 1), and the T50 values decreased to 4.512 days (Tabla 1). Hot water scarication also increased the nal germination level in all tested species, except in the case of C. patens. The germination rate of hot water scaried seeds was similar to that recorded by mechanical scarication in C. patens, A. zanonii, and S. muricatus, but slower in C. striatus and C. reverchonii (Table 1). Dry heat scarication had also a strong effect on nal germination level, with the exception of S. muricatus. In the case of D. pentaphyllum, germination was increased signicantly over the control merely for one heat treatment: 120 C for 10 min. It can be said that germination followed a characteristic response. At low temperatures (50 C) germination was not enhanced in

Discussion All the species included in the study produce a substantial fraction (627%) of softcoated seeds which germinate without the necessity of any particular treatment (control seeds), similar in magnitude to those obtained in other Cytisus species and Genista orida (A orbe n et al. 1990; T rrega et al. 1992). This softcoated seed a fraction is responsible for the maintenance of population levels of these species during periods without re, and explains their colonizing role in disturbed areas as abandonned elds and vegetation gaps. The ability of D. pentaphyllum and the three Cytisus species for regenerating by vegetative regrowth, however, enables them to decrease their dependence on reproduction by seeds. Nevertheless, in all tested species, the proportion of hardcoated seeds which can be softened and, thus, promoted to germinate by mechanical and hot water scarication or thermal pretreatments has been higher. In nature there are many mechanisms producing the crack of the tegumentary barrier in legumes, as temperature oscillation and the alternance of dry and wet periods (Quinlivan 1971; Rolston 1978), bacteria and other soil microrganism action, and the chemical scarication suffered throughout the herbivore digestive system (Pereiras et al. 1985), although the heat generated by wildres is an important trigger factor (Casal 1982;

99

Figure 1. Effect of high temperature pretreatments (50 C150 C) upon the germination of Cytisus striatus, Cytisus reverchonii, Cytisus patens, A. zanonii, S. muricatus, D. pentaphyllum and P. bituminosa seeds. Anova F ratio value for each species is indicated under specic names (p 0:001 in all cases). Different lower case letters above columns indicate signicant differences ( p 0:05) between nal average germination percentages in the pretreatments carried out for each species. Vertical lines in the columns indicate standard deviation. C control seeds; S mechanical scaried seeds; W hot water scaried seeds. 10 , 50 , 100 , 150 , 300 , and 600 are the different dry heat exposure periods (in min) used in the experiment.

100
Table 1. Average T50 values ( standard deviation) recorded for untreated (control; C) seeds, mechanically (S) and hot water (W) scaried seeds, as well as for those thermal treatments which remarkably promoted germination. F Anova F ratio value ( : p 0.01; : p 0:001) for each species. Superscripts with different letters denote signicant differences (p 0:05) between pairs of cases within each species. Blank spaces correspond to absence of germination in a species.

C Cytisus striatus Cytisus reverchonii Cytisus patens Argyrolobium zanonii Scorpiurus muricatus Dorycnium pentaphyllum Psoralea bituminosa 22.00a ( 7.48) 24.00a ( 0.00) 37.50ab ( 0.86) 8.66a ( 3.77) 28.00a ( 4.00) 12.00a ( 1.63) 6.50 ( 0.86)

S 6.00b ( 0.00) 5.50b ( 0.96) 12.00c ( 1.41) 6.00a ( 0.00) 9.50b ( 0.86) 4.50b ( 0.86) 6.50 ( 0.86)

W 31.00a ( 3.00) 27.50a ( 0.86) 14.50c ( 0.86) 6.50a ( 0.86) 9.00b ( 0.00)

90 50 37.00a ( 1.00) 48.50c ( 0.86) 38.00bd ( 4.47) 15.00ab ( 3.60) 32.50a ( 1.65) 11.00a ( 1.00) 7.00 ( 1.00)

90 100 40.50a ( 2.95) 37.50d ( 5.17) 38.00bc ( 2.00) 15.00ab ( 1.00) 30.00a ( 1.63) 9.50ab ( 1.65) 7.50 ( 1.65)

120 10 32.00a ( 12.08) 40.00d ( 1.41) 28.50f ( 3.84) 22.50bc ( 7.92) 32.00a ( 1.63) 12.00a ( 4.69) 6.50 ( 0.86)

120 50 36.50a ( 2.17) 35.00d ( 1.00) 48.00c ( 0.00)

150 C 10 40.50a ( 5.17) 43.00c ( 2.23) 48.50c ( 2.95) 13.00ab ( 2.23) 29.50a ( 4.33) 11.00a ( 1.73) 7.50 ( 1.65)

F 7.08

98.78

15.67

7.22

35.00a ( 3.00) 14.50a ( 0.86) 6.00 ( 0.00)

25,84

3.91

0.77 (n.s.)

Baskin & Baskin 1989; T rrega et al. 1992). As other a plant families having hardseedness feature, in legumes heat cracks the seed coat thus allowing water uptake and removing those mechanical restrictions to radicle expansion (Rolston 1978; Vuillemin & Bullard 1981). Therefore, the hardcoated seed fraction, commonly associated with a persistent soil seed bank (Baskin & Baskin 1989), may be mainly responsible for massive germination of legumes recorded after some wildres, since soft seeds and some of the less hard ones are very likely to be destroyed by re. So, in several burnt stands adjacent to Moropeche (Yeste, province of Albacete), the average cover of P. bituminosa, A. zanonii and D. pentaphyllum in the rst spring after re occurred in August 1989 were 82.00%, 3.78% and 4.98% respectively; they showed values of 0.40%, 0.82% and 5.83% in the spring 1994 (Martnez-S nchez et al. 1996). A a suitable explanation for these post-re legume cover uctuations may be that hardcoated seeds stored in soil as a seed bank persistent component are promoted to germinate in a massive way by high temperatures reached during wildre, although the presence of A. zanonii and P. bituminosa tend to be reduced in subsequent years by competition with other bigger size

plants. Similar cover oscillations during the rst postre years has been described in Serra da Arr bida (Pora tugal) for the Leguminosae species P. bituminosa and Astragalus lusitanicus Lam. (Clemente et al. 1996). The practically total absence of germination recorded for all the studied species at higher temperatures during long exposure times (different among species), must be due to these heating treatments turning out to be fatal for the embryo, as has been suggested by T rrega et al. (1992) for C. scoparius and G. orida. a Results in this direction have been recorded for many legume species in other re-prone regions. So, Martin et al. (1975) showed 100110 C for 4 min as a lethal heat treatment for several Lespedeza, Cassia, Desmodium and Galactia herbaceous species of meadows in southeastern USA. Auld & OConnel (1991) recorded 120 C for 5 min treatment killing whole seedlots of several Acacia, Bossiaea, Daviesia, Glycine, Gompholobium, and Platylobium species of Australian ora. However, seed germination of other Acacia species, as well as of Pultenaea, and Sphaerolobium was stimulated, denoting a wide variation in the specic seed germination requirements.

101 The degree of stimulation of germination by thermal treatments obtained in the present work for some species (i.e., C. striatus, C. reverchonii, A. zanonii and P. bituminosa) show similar values to those obtained for G. orida and C. scoparius (T rrega et al. a 1992), and for Ulex europaeus L. (Pereiras et al. 1985) in Spain, for Acacia saligna (Labill.) Wendl., Podalyria calyptrata Wild., and Virgilia oroboides (Berg.) Salter in South Africa (Jeffery et al. 1988), and for several legume species in eastern Australia (Auld & OConnel 1991). The most effective temperature in promoting C. patens seed germination was 70 C, coinciding with records for six Acacia species in the east coast of Australia (Floyd 1966). Five species in the study had a broad interval of thermal treatments promoting seed germination. This interval is within the range of temperature conditions commonly encountered during res in the soil surface horizons (Trabaud 1979; T rrega 1992), so that any a heat conditions in the soil produced by re induce the softening and subsequent germination of a certain seed quantity. During wildres the heat conditions in soil are not uniform in space and time, because of the ground microrelief, the presence of big stones and rock emergence, and the uneven distribution of litter and standing biomass (Trabaud & Oustric 1989). Thus, when a re takes place, there is always a relatively high number of hardcoat seeds released from physical dormancy, irrespective of the intensity and the duration of the re heat (Thanos & Georghiou 1988). The range of temperatures in the soil allows the seeds of each legume, whose germination is promoted by thermal treatments, to nd their specic heat requirements. The high temperatures promoting germination in those studied species which often share the habitat were notably different. Mixed shrublands with D. pentaphyllum and C. reverchonii are frequent in Betic mountain ranges. Although there is certain risk in using results from laboratory experiments for the interpretation of eld observations, it may be thought that post-re germination in D. pentaphyllum will be only stimulated in those microhabitats where temperatures reach 120 C for 10 min, whereas for C. reverchonii there are a broad range of thermal treatments which promote its seed germination. A similar interpretation of results can be made for those communities where D. pentaphyllum, P. bituminosa, A. zanonii and S. muricatus are important components in the post-re vegetation, as commonly occurs in several localities in the river Tus valley (Yeste, Albacete province). The heat requirements of P. bituminosa and A. zanonii are similar, but very different to those for D. pentaphyllum. The preheating at 120 C for 5 min, however, was effective in promoting germination for P. bituminosa but not for A. zanonii, and the preheating at 90 C for 5 min produced a higher nal germination level in the second species than in the former. S. muricatus seed germination was not stimulated by any dry heat treatment used in the experiment. Its high post-re cover and density values recorded in several locations of river Tus valley (Martnez-S nchez 1994; Martnez-S nchez et al. a a 1996) could be explained taking into account that a considerable fraction of seeds are able to tolerate the re effect (i.e., seeds submitted to 120 C for 10 min treatment showed a nal germination value of 9%), the high seed production in this species, as well as the possible softening of its hardcoated seeds in nature. The strategy of heat requirement differentiation in coexisting species has been already described both in Leguminosae species, as G. orida and C. scoparius (T rrega et al. 1992), and several Cistus species (Traa baud & Oustric 1989), and interpreted as a mechanism which minimizes the intensity of interspecic competition, since it allows a differential establishment of species by occupying different points in the ground after re. The present work fails to nd a clear relationship between the response to heat shock and the life trait or the specic post-re recovery strategy. Moreno & Oechel (1991, 1994) have hypotetized and conrmed for Ceanothus greggii Gray and Adenostoma fasciculatum H. & A. in California chaparral, that seeds of obligate seeders are more resistant to elevate re intensity than facultative resprouters, because the former rely solely on their seeds to regenerate after re. Our results show an opposite pattern in the case of the facultative resprouter C. striatus (and even for D. pentaphyllum), which showed a higher resistance to elevate temperatures (120 C) than S. muricatus, A. zanonii and P. bituminosa, the three obligate seeders in the study. For the other two facultative resprouters, C. reverchonii and C. patens, no clear trend was detected. Future studies on the importance of each post-re regeneration strategy in these facultative resprouters may help to explain the different seed response to heat: a more active involvement of seed germination, instead of resprouting, in the post-re regeneration of C. striatus could be predicted, as observed by Herranz et al. (1996) for the facultative resprouters C. scoparius and G. orida in Tejera Negra Natural Park (central Spain). With the exception of P. bituminosa, the germination rates after thermal pretreatments were much

102 slower than in mechanically scaried seeds and almost similar to those of untreated seeds (for C. reverchonii was even slower than the control set). This behaviour has been previously recorded in Leguminosae species (T rrega et al. 1992) as well as in Cistaceae spea cies (Troumbis & Trabaud 1986; Thanos & Georghiou 1988; Trabaud & Oustric 1989; Thanos et al. 1992). These differences in germination rates has been interpreted as a result of the different effect of mechanical scarication and thermal pretreatments on the seed coat structures in species with hard coat seeds (Thanos et al. 1992). The slow germination rate of softened seeds by heat has been previously considered as an obvious ecological advantage in the summer-dry and re-prone Mediterranean climatic conditions (Thanos & Georghiou 1988; Thanos et al. 1992). It could be interpreted as a delaying mechanism which prevents germination under conditions of occasional rainfall during late summer and early autumn but which do not supply enough moisture for seedling establishment and growth.
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Acknowledgements We wish to thank I. Cuenca and R. Garca for their laboratory assistance, and four anonymous referees for their suggestions on the original manuscript. This work forms a part of a more extensive study focused on postre plant recovery processes in Mediterranean forests of southeastern Spain carried out in the AGF96-1151 project, supported by the Comision Interministerial de Ciencia y Tecnologa of Spain.

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