Reduced germination success of temperate grassland seeds sown in dung: consequences for post‐dispersal seed fate

• Endozoochory is one of the main drivers shaping temperate grassland communities by maintaining plant populations of its constituents and enabling plants to colonize new habitats. Successful endozoochorous dispersal implies that seeds not only get consumed and survive the digestive tract but are also able to develop into viable seedlings in a dung environment.
• We experimentally assessed the germination probability and timing of 15 annual and perennial temperate European grassland species in cattle and horse dung and in different climatic conditions (greenhouse and outdoor conditions).
• Interspecific variation in germinability and germination timing are found, while life strategy had only an effect on germination timing. We found adverse effects of both cattle and horse dung on the germination characteristics of all tested grassland species, but the effects of cattle dung were more pronounced. In comparison with the control treatment, fewer seeds emerged in dung and more time was needed to germinate. Also, germination metrics clearly differed between the artificial greenhouse and outdoor conditions, with generally a lower germinability in outdoor conditions.
• According to our results, a large cost seems to be associated with endozoochorous dispersal in this stage of the life cycle, as seed dispersal effectiveness strongly depends on the quality of the deposition site with a lowered survival and germination probability when seeds are deposited in dung.

     

热激对大兴安岭三种松科树种种子萌发的影响

研究了热激对大兴安岭地区兴安落叶松、樟子松、红皮云杉种子萌发的影响,热激温度分别为80℃、100℃、120℃、150℃,每个温度下热激时间分别为1min、3min、5min、10min。研究结果表明:兴安落叶松种子在100℃热激10min,120℃热激5min,10min,150℃热激5min处理下萌发率较对照显著下降(P%0.05),除80℃热激1min、3min、5min、120℃热激1min、150℃热激1min外,其他处理萌发速率指数较对照均显著下降(P%0.05);樟子松种子在高于80℃的所有热激处理萌发率较对照均显著下降(P%0.05),甚至没有萌发,除80℃热激1min外,所有处理下萌发速率指数较对照均显著下降;红皮云杉种子萌发率在80℃热激5min和100℃热激1min处理下较对照显著提高(P%0.05),萌发速率指数在80℃热激1min、3min、5min,100℃热激1min处理下较对照显著提高(P%0.05)。兴安落叶松种子可以在短时间高强度(150℃)热激下维持萌发率,但不能承受长时间热激;樟子松种子对高于80℃的热激反应敏感,热激降低了樟子松种子的萌发能力;热激可以提高红皮云杉种子的萌发能力,短时间(1min、3min、5min)低强度(80℃)热激(包括100℃热激1min)提高红皮云杉种子萌发率和萌发速率指数,红皮云杉种子也可以承受高强度(150℃)瞬时(1min)热激。     

Seed longevity and fire: post‐germination responses of Rumex acetosella L. in northwest Patagonian grasslands (Argentina)

The cosmopolitan herb Rumex acetosella forms persistent soil seed banks and increases in cover after fire. We investigated how the interaction between seed age and fire affects seedling growth by exposing different‐aged seeds to heat, smoke, charcoal, and ash treatments. We measured growth of germinated seedlings that were transplanted and allowed to grow for 65 days in a greenhouse. Seedlings from seeds >8 years old did not reach an appropriate radicle length for transplantation. Seedling growth decreased with increasing temperature of the heat treatment. As seed age increased, growth decreased with smoke and charcoal, and increased with ash treatment. Height was negatively correlated with seed age. Our results suggest that fire and seed age could affect demographic responses of R. acetosella seedling populations. Post‐fire recruitment could be partially favored by the positive effect of nutrient input from ash on seedling growth. High fire intensities, however, would be detrimental to seedling vigor.     

Response surfaces for the combined effects of heat shock and smoke on germination of 16 species forming soil seed banks in south‐east Australia

Abstract There is limited understanding of how fire‐related cues such as heat shock and smoke can combine to affect the germination response of seeds from fire‐prone vegetation because combinations of multiple levels of both cues have rarely been investigated. Germination response surfaces were determined for the combination of heat shock and smoke by applying factorial combinations of temperature (up to 100°C) and aerosol smoke (0–20 min) to 16 species that form soil seed banks in the Sydney region of south‐eastern Australia. Duplicate populations of three species were also examined to assess the constancy of a species response surface. Of the 19 populations examined, 16 showed a germination response to both the fire cues, which combined interactively in 14 populations, and independently in two. No population responded only to a single cue; however, seeds of 11 populations responded to heat in the absence of smoke, and nine responded to smoke in the absence of heat. Heat applied in the absence of smoke negatively affected germination in seven populations, either progressively as temperature increased, or above a set temperature. Negative germination responses over part of the temperature range were fully reversed at higher temperatures for unsmoked seeds of four populations (curvilinear heat response). Smoke effects were most frequently positive over all or part of the range of durations used, and when combined with heat frequently fully or partially reversed negative heat effects. Three populations required the obligatory combination of smoke and heat. A novel response to the cues was observed for three species, with smoke reversing negative heat effects at 75°C, being supplanted by a positive heat response of unsmoked seed at 100°C. The response surface for duplicate populations of two of the three species examined was variable. Heat shock and smoke frequently combined to affect germination, in both positive and negative ways. Consequently, to gain an accurate assessment of the response of seeds to fires, an experimental design that samples within the potential response zones of germination cues is essential.     

The role of avian ‘seed predators’ as seed dispersers

Seed dispersal is a central process in plant ecology with consequences for species composition and habitat structure. Some bird species are known to disperse the seeds they ingest, whereas others, termed ‘seed predators’, digest them and apparently play no part in dispersal, but it is not clear if these are discrete strategies or simply the ends of a continuum. We assessed dispersal effectiveness by combining analysis of faecal samples and bird density. The droppings of seed dispersers contained more entire seeds than those of typical seed predators, but over a quarter of the droppings of seed predators contained whole seeds. This effect was further magnified when bird density was taken into account, and was driven largely by one frequent interaction: the Chaffinch Fringilla coelebs, a typical seed predator and the most abundant bird species in the area and dispersed seeds of Leycesteria formosa, a non‐native plant with berry‐like fruits. These results suggest the existence of a continuum between seed predators and seed dispersers.     

Interactive influences of fire intensity and vertical distribution of seed banks on post‐fire recolonization of a tall‐tussock grassland in Argentina

Abstract Recolonization patterns of a tall‐tussock grassland differ between low‐frequency and high‐intensity fires. A series of laboratory and field experiments were performed on the seed bank and on seeds of the main colonizing species (Carduus acanthoides, Cirsium vulgare, Lotus glaber) to determine and compare the effects of fire frequency and fire intensity on their recolonization potential. Seed‐bank size of colonizing species did not show a significant variation with contrasting fire frequencies, so overall interval‐dependent effects do not seem to affect their propagule pool at the time of fire. However, frequency of fire modified the proportion of viable seeds and their dormancy state according to species. Accumulated emergence from experimentally buried soft‐seeds of Lotus was lower after high‐intensity fires (following wind direction) than after back‐fires (against wind direction), but no significant effects of fire‐front direction on seedling emergence were observed for hard‐seeds of Lotus nor Carduus and Cirsium. Seedling emergence from buried seeds of Cirsium was increased with doubling fuel loads, particularly from deepen soil layers, where Cirsium had most of their viable seeds after a low frequency of fire. Emergence from hard‐seeds of Lotus was less promoted than Cirsium by doubling fuel loads, and according to analysis of excavated seedlings, it was completely inhibited from upper soil layers of the seed bank. Integration of species‐specific and burial depth‐specific responses of Cirsium and Lotus seeds according to fuel load, vertical distribution of the seed banks, seed dormant states and mean emergence depths, resulted in expected emergence values which agree with their previously observed patterns of recolonization. Thus, event‐dependent mechanisms had a better predictive value on recolonization success of the studied species than the observed cumulative effects of fire history on their seed‐bank size.     

Fire‐related cues and germination from the soil seed bank of senescent remnants of mallee vegetation on Eastern Kangaroo Island

Plant communities dominated by narrow‐leaved mallee (Eucalyptus cneorifolia) are almost entirely confined to north‐eastern Kangaroo Island, South Australia, an area which has been extensively cleared for agriculture. Consequently, surviving examples consist mostly of small remnants which are thought to be senescent due to the exclusion of fire. This senescence is associated with the loss of many native understory species. Prescribed burns have been suggested as a management tool to stimulate the restoration of native plants from the soil seed bank; however, no seed bank studies have previously been conducted on Kangaroo Island and the seed bank literature usually focuses on particular species rather than on plant communities. We conducted an experiment to investigate the effects of the fire‐related cues heat and smoke on the germination of plants from the seed bank in soil sampled from 10 long‐ungrazed narrow‐leaved mallee sites on Kangaroo Island. Eighty trays of soil were monitored in a controlled glasshouse for five months after being subjected to heat and/or smoke treatments. The overall number of native, but not exotic, plant species germinating from the soil seed bank was significantly increased by all three fire‐related treatments (heat, smoke and heat plus smoke) compared with the control (no fire‐related treatment). Different plant life forms exhibited varying responses to heat and smoke treatments. The results of this study illustrate that the application of fire‐related treatments to soil seed banks in controlled glasshouse conditions can stimulate the recruitment of native species, including several species of conservation concern. These findings also indicate the potential of using these treatments for the ex situ germination of fire dependent species for revegetation purposes and indicate aspects of prescribed burns that may be important for restoring different components of native vegetation.     

Interactive effects of heat shock and smoke on germination of nine species forming soil seed banks within the Sydney region

Abstract The germination response of seeds from fire‐prone vegetation to fire‐related cues such as heat shock and smoke has usually been studied by applying the cues singly. The few studies that have applied the cues in combination have shown that interactions between the cues are possible. Here, the response of seeds from a number of species to combined heat shock and smoke is reported. Heat shock (25, 50, 75 and 100°C) and aerosol smoke (0, 5, 10 and 20 min) were applied factorially to nine species that form soil seed banks in the Sydney region of south‐eastern Australia. These species were from Epacridaceae (four species), Myrtaceae (four species) and Cyperaceae (one species) and ranged from fire‐sensitive obligate seeders to fire‐tolerant facultative resprouters. Germination of Dracophyllum secundum R. Br and Sprengelia monticola (A. Cunn. ex DC.) Druce was low and did not respond to the germination cues. The positive response of Gahnia sieberiana Kunth and Kunzea ambigua (Sm.) Druce to heat shock and smoke was independent and additive. The positive response of Kunzea capitata Rchb. to the interaction between heat shock and smoke was synergistic, and the response of Baeckea diosmifolia Rudge and Baeckea imbricata (Gaertn.) Druce was unitive, with germination increase only occurring following combined heat and smoke application. Epacris coriacea A. Cunn. ex DC. and Epacris obtusifolia Sm. had low levels of dormancy and hence it was not possible to find a fire response. Gahnia sieberiana and K. capitata responded differently to the combination of heat shock and smoke than has previously been reported. Germination of species from habitats that are infrequently burnt was not affected by heat shock or smoke. Low‐intensity fire or patches within fire may be important for seedling recruitment as the 50°C heat shock stimulated germination in four of the five species that responded to the heat cue, and germination of Baeckea imbricata declined within the 100°C heat shock treatment. Germination of one species, Baeckea imbricata, was only stimulated by a specific combination of cues, indicating that regeneration niches may be narrow for some species and that the application of a range of heat and smoke doses is required to find such responses. Of the species positively responding to heat shock and smoke, a requirement for both cues was prevalent, therefore the response to these cues in isolation cannot be relied upon to give a true indication of the fire response of a species.     

Seed production and germination in two rare and three common co‐occurring Acacia species from south‐east Australia

Abstract Seed set, size, viability and germination requirements were investigated for two rare (Acacia ausfeldii and A. willianisonii) and three common (A. pycnantha, A. genistifolia and A.paradoxa) co‐occurring congeners in box‐ironbark eucalypt forests near Bendigo, south‐east Australia to investigate correlates of rarity. Seed size was significantly smaller for the two rare species and germinants were less able to emerge from deeper sowing depths than were the larger seeded common congeners. All species had a strong heat‐stimulated germination response. While the rare A. ausfeldii showed strong germination only at the highest temperature treatment (100°C), the common and widespread A.pycnantha showed strong germination across a broad range of temperatures (60‐100°C), likely to be experienced by soil‐stored seeds during a fire. Seed viability, number of seeds per plant, and number of firm, aborted and eaten seeds per pod varied between species, but the pattern of variation was not related to rarity. Small seed size and a very specific temperature requirement for germination may help to explain rarity in A. ausfeldii, and to a lesser extent in A. willianisonii. Fires are often patchy and heating of the soil is likely to be highly spatially variable, so species with germination responses to a broad range of temperatures have an advantage over those that respond only to a narrow range. A narrower range of soil depths from which seeds can emerge will further reduce the proportion of the seed bank that might recruit following fire. Human impacts on species habitats, such as fragmentation, loss of topsoil through mining, timber harvesting, grazing and urbanization, and consequent reduction in fire intensity, are likely to have further contributed to rarity in these species. The role of pollination and other factors in relation to population size is the subject of further investigation.     

Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

Costs and benefits of non‐random seed release for long‐distance dispersal in wind‐dispersed plant species

The dispersal ability of plants is a major factor driving ecological responses to global change. In wind‐dispersed plant species, non‐random seed release in relation to wind speeds has been identified as a major determinant of dispersal distances. However, little information is available about the costs and benefits of non‐random abscission and the consequences of timing for dispersal distances. We asked: 1) to what extent is non‐random abscission able to promote long‐distance dispersal and what is the effect of potentially increased pre‐dispersal risk costs? 2) Which meteorological factors and respective timescales are important for maximizing dispersal? These questions were addressed by combining a mechanistic modelling approach and field data collection for herbaceous wind‐dispersed species. Model optimization with a dynamic dispersal approach using measured hourly wind speed showed that plants can increase long‐distance dispersal by developing a hard wind speed threshold below which no seeds are released. At the same time, increased risk costs limit the possibilities for dispersal distance gain and reduce the optimum level of the wind speed threshold, in our case (under representative Dutch meteorological conditions) to a threshold of 5–6 m s^–1. The frequency and predictability (auto‐correlation in time) of pre‐dispersal seed‐loss had a major impact on optimal non‐random abscission functions and resulting dispersal distances. We observed a similar, but more gradual, bias towards higher wind speeds in six out of seven wind‐dispersed species under natural conditions. This confirmed that non‐random abscission exists in many species and that, under local Dutch meteorological conditions, abscission was biased towards winds exceeding 5–6 m s^–1. We conclude that timing of seed release can vastly enhance dispersal distances in wind‐dispersed species, but increased risk costs may greatly limit the benefits of selecting wind conditions for long‐distance dispersal, leading to moderate seed abscission thresholds, depending on local meteorological conditions and disturbances.     

Dispersal by rodent caching increases seed survival in multiple ways in canopy‐fire ecosystems

Seed‐caching rodents have long been seen as important actors in dispersal ecology. Here, we focus on the interactions with plants in a fire‐disturbance community, specifically Arctostaphylos species (Ericaceae) in California chaparral. Although mutualistic relationships between caching rodents and plants are well studied, little is known how this type of relationship functions in a disturbance‐driven system, and more specifically to systems shaped by fire disturbance. By burying seeds in the soil, rodents inadvertently improve the probability of seed surviving high temperatures produced by fire. We test two aspects of vertical dispersal, depth of seed and multiple seeds in caches as two important dimensions of rodent‐caching behavior. We used a laboratory experimental approach to test seed survival under different heating conditions and seed bank structures. Creating a synthetic soil seed bank and synthetic fire/heating in the laboratory allowed us to have control over surface heating, depth of seed in the soil, and seed cache size. We compared the viability of Arctostaphylos viscida seeds from different treatment groups determined by these factors and found that, as expected, seeds slightly deeper in the soil had substantial increased chances of survival during a heating event. A key result was that some seeds within a cache in shallow soil could survive fire even at a depth with a killing heat pulse compared to isolated seeds; temperature measurements indicated lower temperatures immediately below caches compared to the same depth in adjacent soil. These results suggest seed caching by rodents increases seed survival during fire events in two ways, that caches disrupt heat flow or that caches are buried below the heat pulse kill zone. The context of natural disturbance drives the significance of this mutualism and further expands theory regarding mutualisms into the domain of disturbance‐driven systems.     

RopGEF2 is involved in ABA‐suppression of seed germination and post‐germination growth of Arabidopsis

The involvement of Rho of Plants (ROP) GTPases in abscisic acid (ABA) signalling in Arabidopsis has been demonstrated in many studies. However, the roles of RopGEFs (Rop guanine nucleotide exchange factors), which modulate ROP activities in ABA signalling, are poorly understood. Here, we demonstrate that RopGEF2 may play a negative role in ABA‐suppressed seed germination and post‐germination growth. We show that disruption of RopGEF2 enhances sensitivity to exogenous ABA in seed germination assays and that RopGEF2pro‐GUS is mainly expressed in developing embryos and germinating seeds. Interestingly, YFP‐RopGEF2 is located in both the cytoplasmic region and in mitochondria. Notably, the PRONE2 (plant‐specific ROP nucleotide exchanger 2) domain of RopGEF2 is detected in mitochondria, whereas the N‐terminus of RopGEF2 is shown to be in the cytosol. After ABA treatment, degradation of RopGEF2 is triggered in the cytosol through the ubiquitin‐26S proteasome system. The binding of RopGEF2 to ROP2, ROP6 or ROP10, which has been demonstrated to be involved in ABA signalling, not only alters the localization of RopGEF2 but also enables RopGEF2 to escape degradation in the cell. Thus, in this study, we deduce a sophisticated mechanism of ABA‐mediated RopGEF2‐ROP signalling, which potentially implicates the inactivation of ROPs in responsiveness to ABA.     

Fire effects on the soil seed bank and post‐fire resilience of a semi‐arid shrubland in central Argentina

Soil seed bank is an important source of resilience of plant communities who suffered disturbances. We analysed the effect of an intense fire in the soil seed bank of a semi‐arid shrubland of Córdoba Argentina. We asked if the fire affected seed abundance, floristic and functional composition of the soil seed bank at two different layers (0–5 cm and 5–10 cm), and if fire could compromise the role of the soil seed bank as a source of resilience for the vegetation. We collected soil samples from a burned site and from a control site that had not burned. Samples were installed in a greenhouse under controlled conditions. During 12 months, we recorded all germinated seedlings. We compare soil seed bank with pre‐fire vegetation in terms of floristic and functional composition. The high‐intensity fire deeply affected the abundance of seeds in the soil, but it did not affect its floristic or functional composition. Floristic and functional composition of soil seed banks – at burned and unburned sites‐ differed markedly from that of the pre‐fire vegetation, although a previous study at the same site indicated high resilience after fire of this plant community. Our results indicate that resilience of this system is not strongly dependent on direct germination from seeds buried in the soil. Other sources of resilience, like colonization from neighbouring vegetation patches and resprouting from underground organs appear to gain relevance after an intense fire.     

Gut passage effect of the introduced red‐whiskered bulbul (Pycnonotus jocosus) on germination of invasive plant species in Mauritius

In Mauritius, many of the worst invasive plant species have fleshy fruits and rely on animals for dispersal. The introduced red‐whiskered bulbul (Pycnonotus jocosus) feeds on many fleshy‐fruited species, and often moves from invaded and degraded habitats into higher quality native forests, thus potentially acting as a mediator of continued plant invasion into these areas. Furthermore, gut passage may influence seed germination. To investigate this, we fed fleshy fruits of two invasive plant species, Ligustrum robustum and Clidemia hirta, to red‐whiskered bulbuls. Gut passage times of seeds were recorded. Gut‐passed seeds were sown and their germination rate and germination success compared with that of hand‐cleaned seeds, as well as that of seeds in whole fruits. Gut passage and hand‐cleaning had significant positive effects on germination of both species. Gut‐passed seeds of both C. hirta and L. robustum germinated faster than hand‐cleaned seeds. However, for L. robustum, this was only true when compared with hand‐cleaned seeds with intact endocarp; when compared with hand‐cleaned seeds without endocarp, there was no difference. For overall germination success, there was a positive effect of gut passage for C. hirta, but not for L. robustum. For both C. hirta and L. robustum, no seeds in intact fruits geminated, suggesting that removal of pulp is essential for germination. Our results suggest that, first, the initial invasion of native forests in Mauritius may not have happened so rapidly without efficient avian seed dispersers like the red‐whiskered bulbul. Second, the bulbul is likely to be a major factor in the continued re‐invasion of C. hirta and L. robustum into weeded and restored conservation management areas.     

Stimulation of weed seed germination by 1–(3-chlorophthalirnido) cyclohexanecarboxamide (AC 94377)

The N-substituted phthalimide, AC 94377, promoted the germination of dry-stored seeds of 17 out of 24 weed species in Petri-dish tests maintained at 22°C. In a further test it was able to substitute for light and/or alternating temperatures in promoting the germination of some species. AC 94377 was more active than GA, in stimulating Solanum nigrum seed to germinate. When mixed with soil in the laboratory, AC 94377 again promoted germination of S. nigrum but the level and persistence of activity differed between soils. In the glasshouse, with soil containing a natural weed seed population, more seedlings emerged from treated than from untreated soil but this number represented fewer than half of the apparently viable seeds present. Field applications of AC 94377 promoted the germination of hand-sown seeds of certain weed species but did not increase the number of seedlings emerging from the natural weed seed bank.     

Implications of nonrandom seed abscission and global stilling for migration of wind‐dispersed plant species

Migration of plant populations is a potential survival response to climate change that depends critically on seed dispersal. Biological and physical factors determine dispersal and migration of wind‐dispersed species. Recent field and wind tunnel studies demonstrate biological adaptations that bias seed release toward conditions of higher wind velocity, promoting longer dispersal distances and faster migration. However, another suite of international studies also recently highlighted a global decrease in near‐surface wind speeds, or ‘global stilling’. This study assessed the implications of both factors on potential plant population migration rates, using a mechanistic modeling framework. Nonrandom abscission was investigated using models of three seed release mechanisms: (i) a simple drag model; (ii) a seed deflection model; and (iii) a ‘wear and tear’ model. The models generated a single functional relationship between the frequency of seed release and statistics of the near‐surface wind environment, independent of the abscission mechanism. An Inertial‐Particle, Coupled Eulerian‐Lagrangian Closure model (IP‐CELC) was used to investigate abscission effects on seed dispersal kernels and plant population migration rates under contemporary and potential future wind conditions (based on reported global stilling trends). The results confirm that nonrandom seed abscission increased dispersal distances, particularly for light seeds. The increases were mitigated by two physical feedbacks: (i) although nonrandom abscission increased the initial acceleration of seeds from rest, the sensitivity of the seed dispersal to this initial condition declined as the wind speed increased; and (ii) while nonrandom abscission increased the mean dispersal length, it reduced the kurtosis of seasonal dispersal kernels, and thus the chance of long‐distance dispersal. Wind stilling greatly reduced the modeled migration rates under biased seed release conditions. Thus, species that require high wind velocities for seed abscission could experience threshold‐like reductions in dispersal and migration potential if near‐surface wind speeds continue to decline.     

Temporal and spatial patterns of seed dispersal in two Cistus species (Cistaceae)

Cistus species are obligate seeding, early colonizers that follow disturbance, particularly fire, in Mediterranean ecosystems. We studied seed release, seed dispersal and soil seed populations in stands of Cistus ladanifer and C. libanotis. Seed release started in mid- to late summer (C. ladanifer) or in early autumn (C libanotis), and continued for a very extended period: 8-10 months in C. ladanifer, and for a mean of 16 months in C. libanotis. The xerochastic capsules of both species released seeds by successive dehiscence of the locules. All capsules begin to dehisce simultaneously at the start of the seed release period, but in C. libanotis capsule fragmentation replaced dehiscence early in the seed release period. In plants of both species, seed shadows were characterized by a peak of density beneath the plant canopy and a very short tail of much lower densities, indicating that seeds are concentrated beneath mother plants when dispersed. Nevertheless, in late May, at the onset of the fire season, soil seed densities beneath plant canopies were low compared with densities expected from seed shadows, but were apparently high enough to allow recovery of the stands if a disturbance, such as fire, had taken place. Seed-eating Bruchidae in summer and granivorous ants during the seed release period were apparently the main causes of seed losses. Results suggest that in both Cistus species, the staggered seed release could constitute an efficient risk-reducing trait. The plant pool of seeds existing throughout most of the year could be a relevant component of Cistus seed banks.