Soil warming increases plant species richness but decreases germination from the alpine soil seed bank

Global warming is occurring more rapidly above the treeline than at lower elevations and alpine areas are predicted to experience above average warming in the future. Temperature is a primary factor in stimulating seed germination and regulating changes in seed dormancy status. Thus, plant regeneration from seed will be crucial to the persistence, migration and post disturbance recruitment of alpine plants in future climates. Here, we present the first assessment of the impact of soil warming on germination from the persistent alpine soil seed bank. Contrary to expectations, soil warming lead to reduced overall germination from the soil seed bank. However, germination response to soil temperature was species specific such that total species richness actually increased by nine with soil warming. We further explored the system by assessing the prevalence of seed dormancy and germination response to soil disturbance, the frequency of which is predicted to increase under climate change. Seeds of a significant proportion of species demonstrated physiological dormancy mechanisms and germination of several species appeared to be intrinsically linked to soil disturbance. In addition, we found no evidence of subalpine species and little evidence of exotic weed species in the soil, suggesting that the soil seed bank will not facilitate their invasion of the alpine zone. In conclusion, changes in recruitment via the alpine soil seed bank can be expected under climate change, as a result of altered dormancy alleviation and germination cues. Furthermore, the alpine soil seed bank, and the species richness therein, has the potential to help maintain local species diversity, support species range shift and moderate species dominance. Implications for alpine management and areas for further study are also discussed.     

DELLA-mediated cotyledon expansion breaks coat-imposed seed dormancy

Seed dormancy is a key adaptive trait in plants responsible for the soil seed bank. The long established hormone-balance theory describes the antagonistic roles of the dormancy promoting plant hormone abscisic acid (ABA), and the germination promoting hormone gibberellin (GA) in dormancy control. Light, temperature, and other dormancy-breaking signals function to modulate the synthesis and perception of these hormones in the seed. However, the way in which these hormones control dormancy in the imbibed seed remains unknown. Here, we show that the DELLA protein regulators of the GA response are required for dormancy and describe a model through which hormone signal integration and dormancy regulation is achieved. We demonstrate that cotyledon expansion precedes radicle emergence during Arabidopsis seed germination and that a striking correlation exists between final seedling cotyledon size and seed dormancy in the DELLA mutants. Furthermore, twelve previously characterized seed-dormancy mutants are also defective in the control of cotyledon size in a manner consistent with their effect on germination potential. We propose that DELLA-mediated, light-, temperature-, and hormone-responsive cotyledon expansion prior to radicle emergence overcomes dormancy imposed by the seed coat and underlies seed-dormancy control in Arabidopsis.     

Seed bank assembly follows vegetation succession in dune slacks

Question: Is the seed bank in dune slacks during the whole successional range mainly composed of early successional species or does it vary according to the changing vegetation? Location: Belgium, western part of the coast. Methods: We investigated the soil seed bank composition using a seedling germination method in a chronosequence of 20 dune slacks, ranging in age from five to 55 yr. Results: Both seed density and species richness in the seed bank were very low in the first successional stages and increased with age, mainly as a result of increasing seed production. The similarity between seed bank and vegetation was higher in older slacks. A comparison of characteristics between seed bank and vegetation showed that the seed bank was, to a large extent, composed of later successional species. Occurrence patterns of individual species also showed that seeds become incorporated in the soil after the species has established in the vegetation. Conclusion: The seed bank is not likely to be the driving force for successional changes in the vegetation, and successional changes rely on dispersal. Some early successional species persist in the seed bank, but generally only in low numbers. The results also confirm that most typical dune slack species do not form persistent seeds, so that re‐establishment from the seed bank is not to be expected when the species has disappeared from the vegetation.     

Persistence of seed bank under thick volcanic deposits twenty years after eruptions of Mount Usu, Hokkaido Island, Japan

The topsoil that contained the seed bank became buried under thick tephra after the eruptions of Mount Usu during 1977 and 1978. To determine the seed bank potential of the topsoil 20 yr after the eruptions, i.e., in 1998, 408 100-cm(3) samples were excavated under 115-185 cm of volcanic deposits. The topsoil was collected at 10-cm intervals along the horizontal scale and was divided into a 0-5 cm deep upper layer and a 5-10 cm deep lower layer. The seed bank was estimated by both the germination (GM) and flotation (FM) methods. In total, 23 species with an average seed density of 1317 seeds/m(2) were identified by GM, and 30 species with a density of 2986 seeds/m(2) were extracted by FM. The dominant species was Rumex obtusifolius, and perennial herbs, such as Carex oxyandra, Viola grypoceras, and Poa pratensis, were common. For nine species this study provided the first records for field seed longevity >20 yr. The seed density in the upper layer was double that in the lower layer, and the horizontal distribution was heterogeneous even at 10-cm intervals. We concluded that the seed bank has retained the original structure of the seed bank under the tephra and will persist longer with soil water content between 20 and 40%, no light, and low temperature fluctuations (±0.17°C of standard deviation in a day).     

Seasonal variation in seed bank composition and its interaction with nutrient enrichment in the Everglades wetlands

In the Florida Everglades, nutrient enrichment from agricultural outflow and the change in hydrology have collectively contributed to the expansion of cattails (Typha spp.). To assess the effectiveness of prescribed fire in controlling cattails and to predict vegetation dynamics after the fire, it is important to understand the seasonal variation of the soil seed bank and how the seed bank is affected by nutrient enrichment and fire. This paper investigates the effects of season, nutrient enrichment, and fire on soil seed bank species composition, richness, and density along a nutrient gradient in Water Conservation Area 2A (WCA 2A) of the Florida Everglades. Species richness was significantly affected by nutrient enrichment and season but not their interaction. Total seed density, however, was significantly affected by the interaction between nutrient enrichment and season. Yet, at species level, the relationship between seed density, nutrient enrichment and season varied. The highest seed density of cattail occurred in summer at highly enriched sites, but that of sawgrass occurred in fall regardless of enrichment; the seed density of water lily was very low regardless of season and nutrient enrichment, and the highest Amarathus seed density occurred at highly enriched sites year round. Moreover, germination timing differed greatly among species. While cattail seeds had a short incubation period and started to germinate 2–3 days after initiation of the germination assay, sawgrass seeds generally started to germinate 4 weeks later. Further, both the prescribed summer fire at the highly enriched site and the natural winter fire at the moderately enriched site reduced the seed density of cattail but not of sawgrass. Our results suggest that fire application for vegetation recovery in WCA 2A would benefit from explicitly considering seasonal dynamics of the seed bank.     

准噶尔盆地梭梭土壤种子库基本特征

土壤种子库是种子的安全岛,是干旱荒漠区植被更新与恢复的基础。以梭梭为主的荒漠植被是准噶尔盆地绿洲区重要的生态屏障,但是,对于与梭梭种群自然更新紧密联系的土壤种子库特征的研究却相对缺乏。本研究通过野外采样、物理筛分并结合室内实验分析,对准噶尔盆地梭梭土壤种子库基本特征进行了研究。结果表明：（1）梭梭种群平均土壤种子库密度从71~696 粒·m^-2,局部小环境甚至达到7 534 粒·m^-2;（2）变异系数分析表明,各种群土壤种子库密度变异系数均大于1,说明梭梭土壤种子库的数量分布是非常不均匀的;（3）梭梭种群土壤种子库中种子萌发率从2.08%到47.62%不等,平均约18%;（4）变异函数分析表明,梭梭土壤种子库中种子分布的空间变异较大。综上,准噶尔盆地梭梭土壤种子库密度大且种子萌发能力强,同时其时空分布差异明显,这些特征均将影响梭梭种群的分布格局和种群更新。     

The underground phase: a special challenge in studies of terrestrial orchid populations

Orchid seeds are minute and the first seedling stages of terrestrial species are underground, so there is little knowledge about the biology of the early life history and the size of seed/seedling populations relative to the number of plants that eventually emerge above ground. A recently developed field sowing technique makes it possible to explore a variety of topics such as the length of time that seeds spend as part of the soil seed bank, the phenology of seed germination, substrate requirements for germination, dependence of germination on fungal availability, substrate requirements and growth rate of seedlings, seedling phenology and development, and seedling-fungus specificity patterns in the soil. Several of these applications are relevant for conservation issues. Studies of Goodyera pubescens, Corallorhiza odontorhiza and Liparis lilifolia show considerable variation in germination strategies, patterns of seedling development and establishment, substrate requirements, and levels of specificity with the fungi.     

Global asymptotic stability of plant-seed bank models

Many plant populations have persistent seed banks, which consist of viable seeds that remain dormant in the soil for many years. Seed banks are important for plant population dynamics because they buffer against environmental perturbations and reduce the probability of extinction. Viability of the seeds in the seed bank can depend on the seed’s age, hence it is important to keep track of the age distribution of seeds in the seed bank. In this paper we construct a general density-dependent plant-seed bank model where the seed bank is age-structured. We consider density dependence in both seedling establishment and seed production, since previous work has highlighted that overcrowding can suppress both of these processes. Under certain assumptions on the density dependence, we prove that there is a globally stable equilibrium population vector which is independent of the initial state. We derive an analytical formula for the equilibrium population using methods from feedback control theory. We apply these results to a model for the plant species Cirsium palustre and its seed bank.     

Can spatial variation and inter-annual variation in precipitation explain the seed density and species richness of the germinable soil seed bank in a tropical dry forest in north-eastern Brazil?

The interaction between microhabitat and inter-annual variation in precipitation has an important role on the dynamics of the seed bank and can play a crucial role in survival and maintenance of plant populations in semi-arid environments. We hypothesized that the type of microhabitat and the inter-annual variability in precipitation can explain the richness and density of the seed bank in a semi-arid region in Brazil. The study was conducted in an area of tropical dry forest with shrub-tree physiognomy, locally called caatinga. We collected 35 soil samples in three distinct microhabitats, at the end of rainy and dry seasons, respectively, over three years, totalling 630 samples. The seed bank (richness and seed density) were determined by seedling emergence method. Over the three years, 79 species emerged from the seed bank, 64, 45 and 42 in riparian, non-riparian and rocky microhabitats, respectively. We recorded differences in species richness and average density between microhabitats and between years, with significant statistical interaction between them. Inter-annual precipitation explained 48% and 5% of the variation in richness and seed density, respectively. Spatial variation explained 7% of the species richness and 31% of the density. Our results show that the interaction between spatial variation and precipitation has an important role on the spatial and temporal heterogeneity of the richness and density of seed banks in dry environments.     

Spatial relationships between the standing vegetation and the soil seed bank in a fire-prone encroached dehesa in Central Spain

Postfire vegetation regeneration in many fire-prone ecosystems is soil seed bank dependent. Although vegetation and seed bank may be spatially structured, the role of prefire vegetation patterns and fire in determining postfire vegetation patterns is poorly known. Here, we investigated the spatial patterning of species abundance and richness in the vegetation and seed bank of a Mediterranean encroached dehesa in Central Spain. The seed bank was studied with and without a heat shock simulating a spatially homogeneous fire. Semivariograms and cross-semivariograms showed that species richness in the vegetation was aggregated in patches, mainly of herbs, with highest values corresponding to high herb cover and low tree cover. Species richness in the seed bank was also structured in patches, but the spatial pattern was weak. Seedling density of germinates in the seed bank also showed weak spatial pattern. Heating increased overall germination and species richness, and the intensity of the spatial pattern of species richness, particularly of herbaceous species. However, seed bank density patterns disappeared after heat shock because of increased germination of shrubs without spatial pattern. Our results document that the spatial structure of plant richness in the vegetation may persist after fire due to the spatial patterns of herbaceous species in the seed bank, and that postfire species richness patterns can arise independently of fire intensity patterns. However, the spatial structure of the vegetation after fire can be altered by the feedback between shrub encroachment and an eventual fire because of the ubiquitous germination of shrubs.     

Effect of fire on hard-coated Cistaceae seed banks and its influence on techniques for quantifying seed banks

The impact of fire on hard-coated Cistaceae (Halimium ocymoides, Cistus ladanifer, and C. salvifolius) soil seed banks in a Mediterranean 'maquis' shrubland, and its effect on seed germinability were studied. The study also contrasts the effectiveness of two widely used techniques for quantifying seed banks, the seedling emergence and the physical separation methods, in relation to fire. The null hypothesis that a massive enhancement of physically-dormant Cistaceae seed germination by fire would make use of the time-consuming physical separation technique unnecessary was tested. Fire reduced Cistaceae seed banks in the 0–2 cm deep soil layer by both seed fire-consumption and lethal temperatures, revealed by the significant decreasing of the seed bank density and by the increase of apparently-intact but soft-unviable seeds, respectively. In contrast, no damage was recorded in the 2–5 cm soil layer. A dramatic seed bank depletion (> 90%) in both soil layers was recorded one year after fire in the burnt area, coinciding with a significant increase of seedling density confined to the first post-fire year. The ecological consequences of this massive post-fire seed bank input are discussed. A germinability test revealed that germination of surviving Cistaceae seeds was significantly enhanced in all cases except for the C. salvifolious seed bank in the deeper soil layer. However, final germination levels (60–75%) did not correspond to the magnitude of seed bank depletion, especially for C. salvifolious, which suggests that other environmental factors not exclusively associated with fire may also be important in softening Cistaceae seeds. Germination enhancement by fire soil-heating was not high enough to reject the physical separation technique, at least in the deeper soil layer. The simultaneous use of both seedling emergence and physical separation is recommended for reliable seed bank estimates when a physically-dormant hard-seeded component can be expected in the soil, as in many Mediterranean ecosystems, regardless of fire occurrence.     

天山中部天山云杉林土壤种子库年际变化

通过定点采样,采用萌发法对天山中部天山云杉(Picea schrenkiana Fisch.et Mey.)近熟林(101-120a)和成熟林(121-160a) 2004-2011年(8a)土壤种子库物种组成、种子密度的年际变化和不同间隔年限土壤种子库物种组成的相似性进行了分析.结果表明:(1)土壤种子库中共萌发鉴定出种子植物87种,隶属29科70属,其中乔木种子植物2种,灌木种子植物2种,草本种子植物83种,土壤种子库中草本植物种子密度远远大于木本植物种子密度;8个采样年份土壤种子库恒有种仅有6种;(2)土壤种子库种子密度及其中天山云杉种子密度存在巨大的年际变动,且不具有同步性;土壤种子库种子密度最大(2009年)值为(953.75±66.12)粒/m2,最小(2008年)值为(186.50±20.37)粒/m2,其中天山云杉种子密度最高(2006年)达到(584.50±53.58)粒/m2,最低(2005年)仅有(0.25±0.26)粒/m2;(3)天山云杉林土壤种子库年际间物种组成的相似性不高,Czekanowski相似系数均值仅为0.344,并随间隔年限的增加呈现减小—增大—减小的变化趋势.天山云杉林土壤种子库物种组成和种子密度稳定性差,年际间相差悬殊,物种组成的相似性不高,种子库中天山云杉种子密度主要受其种子库采样前一年天山云杉结实丰歉的影响,属间断型.土壤种子库年际变化特征可为天山森林的更新恢复和可持续经营提供科学依据.     

长久性紫茎泽兰土壤种子库

具有长久性土壤种子库的植物在适应多变的生境和不良的生长条件方面具有优越性。于紫茎泽兰（Eupatorium adenophorum）的主要萌发时段后（7月到次年的4月间）在云南的5个地点采集了共１９个不同植被覆盖下的土壤种子库样本,萌发实验结果表明,紫茎泽兰具有长久性的土壤种子库,其在云南的不同生境的土壤中广泛分布,所有19个样地中都有长久性的紫茎泽兰种子。 0～10 cm土层的种子密度变动于47～13 806 ind.·m-2,平均为2 202 ind.·m-2。种子密度与样地内地表的紫茎泽兰间没有直接的联系,但与植被的覆盖状况有关,种子库密度由滑坡堆积物（47 ind.·m-2）到草地（801 ind.·m-2）到灌丛（2 349 ind.·m-2）到森林（3 255 ind.·m-2）间逐渐增加。种子在各种类型土壤的采样点间出现的频度为60%～100%。在土壤的垂直方向上,0～2 cm土层分布有较多的种子,2～5 cm土层次之,5～10 cm土层最少,其各层占总数的比例的平均值分别为56.1%、25.2%和18.6%。但值得注意的是,虽然紫茎泽兰的种子在5～10 cm 深的土层内的存在量占总量的比例相对较少,但如果折合成密度值,其量仍高达270 ind.·m-2,仍有形成危害的潜在可能。广泛分布且数量巨大的具有长久性特性的紫茎泽兰土壤种子库对各种防治措施的制订意义重大,它要求我们长远地、大尺度地考虑防治措施。     

Germ banks affect the inference of past demographic events

Continuous progress in empirical population genetics based on the whole‐genome polymorphism data requires the theoretical analysis of refined models in order to interpret the evolutionary history of populations with adequate accuracy. Recent studies focus prevalently on the aspects of demography and adaptation, whereas age structure (for example, in plants via the maintenance of seed banks) has attracted less attention. Germ banking, that is, seed or egg dormancy, is a prevalent and important life‐history trait in plants and invertebrates, which buffers against environmental variability and modulates species extinction in fragmented habitats. Within this study, we investigate the combined effect of germ banking and time‐varying population size on the neutral coalescent and particularly derive the allele frequency spectrum under some simplifying assumptions. We then perform an ABC analysis using two simple demographic scenarios—a population expansion and an instantaneous decline. We demonstrate the appreciable influence of seed banks on the estimation of demographic parameters depending on the germination rate with biases scaled by the square of the germination rate. In the more complex case of a population bottleneck, which comprises an instantaneous decline and an expansion phase, ignoring information on the germination rate denies reliable estimates of the bottleneck parameters via the allelic spectrum. In particular, when seeds remain in the bank over several generations, recent expansions may remain invisible in the frequency spectrum, whereas ancient declines leave signatures much longer than in the absence of seed bank.     

Practical method of germination for a key jarrah forest species: Snottygobble (Persoonia longifolia)

Summary Snottygobble ( Persoonia longifolia ) is an ecologically and economically important species in the jarrah forest in Western Australia but is not well represented in jarrah forest restoration projects because it is difficult to germinate. In restored bauxite mines the establishment density of Snottygobble from the soil seed bank is variable and often inadequate. Alcoa considers it a priority to re-establish such key species at adequate densities in its restored mine areas The aims of the present research were: (i) to determine if the species could be cued to germinate; and (ii) to develop a practical method to re-establish the species in restored bauxite mines. We found that fresh Snottygobble seed has high viability (> 90%) but seed stored at 4°C rapidly loses viability over the first year after seed fall. We obtained up to 40% germination using fresh seed that had been treated with gibberellic acid (GA3) after having part of the endocarp chipped away, sown on the soil surface and watered twice daily in an ambient temperature glasshouse in winter/spring. We found that the key to successful germination was combining surface sowing, endocarp chipping and GA3 treatment. Germination involved the breakdown of mechanical and, probably, chemical dormancy. There also appears to be a cool temperature requirement for germination. Practical recommendations to germinate Snottygobble are made. This germination method will have application to land managers, restoration practitioners and the horticultural industry. Alcoa will continue work to translate this success into an adequate stocking of Snottygobble in restored bauxite mines.
Key words bauxite mine, dormancy, endocarp chipping, endogenous, germination, gibberellic acid , Persoonia longifolia , restoration, Snottygobble.     

Restoration of ditch bank plant species richness: The potential of the soil seed bank

Abstract. Small‐scale landscape elements, such as ditch banks, play an important role in preserving plant species richness in agricultural landscapes. In this study, we investigate whether the seed bank might be useful for restoring the above‐ground plant species richness. We studied the vegetation and seed bank composition at six species‐rich and six species‐poor ditch banks, where agri‐environment schemes are running to maintain and enhance ditch bank plant diversity. We show that the number of species in the seed bank was low, regardless of the number of species in the established vegetation. Moreover, the seed bank was always dissimilar to the established vegetation. Target species for nature conservation were occasionally present in the seed bank at both species‐poor and species‐rich sites, but rarely so if the species was absent from the established vegetation. We conclude that the potential use of the seed bank for restoration of ditch banks is minimal. At present, plant species richness seems to be largely controlled by germination opportunities; high biomass and competition appear to hamper germination at species‐poor sites. We recommend continued nutrient reduction at such sites. Soil disturbance measures and deliberate sowing should also be considered.     

青藏高原东缘封育和退化高寒草甸种子库差异

采用幼苗萌发法研究了青藏高原东部高寒草甸封育和退化地区土壤种子库的差异.结果表明:(1)两个样地总共有10161株幼苗萌发,分属55个物种,23个科.封育样地多年生植物比例高于退化样地,禾草和单子叶植物比例在两个样地间差异不大.(2)退化样地土壤种子库种子密度((6105±1530) m-2)显著高于封育样地((3883±798) m-2),而物种丰富度差异不显著.所研究区域拥有较为丰富的种子库资源,退化地区资源更丰富,说明恢复不存在种子限制问题,土壤种子库可以成为植被恢复的潜在资源.(3)种子密度和物种丰富度在垂直分层上差异显著,且随着深度的增加而显著减小.(4)用Srensen coefficient指数计算出土壤种子库和地上植被之间的相似性在整体上较低,封育样地(45.3%)略高于退化样地(40.4%),两个样地地上植被之间的相似性为50%,而种子库之间相似性高达84.6%,说明种子库起到一个"缓冲器"的作用.(5)Shannon-Wiener多样性指数分析显示,无论在地上植被或种子库中,物种多样性都是封育样地显著高于退化样地.过度放牧不仅导致地上植被的生物多样性丧失,而且使种子库中的物种多样性降低,而封育管理可以维持地上植被和种子库中的物种多样性.     

A stochastic population model for Lepidium papilliferum (Brassicaceae), a rare desert ephemeral with a persistent seed bank

Population viability analysis (PVA) is a valuable tool for rare plant conservation, but PVA for plants with persistent seed banks is difficult without reliable information on seed bank processes. We modeled the population dynamics of the Snake River Plains ephemeral Lepidium papilliferum using data from an 11-yr artificial seed bank experiment to estimate age-specific vital rates for viability loss and germination. We related variation in postgermination demographic parameters to annual variation in precipitation patterns and used these relationships to construct a stochastic population model using precipitation driver variables. This enabled us to incorporate realistic levels of environmental variability into the model. A model incorporating best estimates for parameter values resulted in a mean trajectory for seed bank size that remained essentially stable through time, although there was a measurable risk of extinction over a 100-yr period for the study population under this scenario. Doubling the annual seed viability loss rate resulted in near-certain extinction, as did increasing first-year germination to 100%, showing the importance of the persistent seed bank. Interestingly, increasing environmental variance substantially decreased the risk of extinction, presumably because this plant relies on extremely good years to restock the persistent seed bank, while extremely bad years have little impact. If every year were average in this desert environment, the species could not persist. Simulated effects of livestock trampling resulted in greatly increased extinction risk, even over time frames as short as 15 years.     

The coupling of recruitment and disturbance by fire in two resprouting Proteaceae species

Recruitment in plant populations is often tightly coupled to major disturbances such as fires. For species with persistent seed banks, fire-related cues may allow or enhance germination. The litter layer influences germination and may modify the impact of seed predators on seeds and seedlings. The litter layer is obviously affected by fire, providing one mechanism by which disturbance can determine recruitment. We tested the role of litter in the disturbance–recruitment coupling of two species with contrasting seed release timing after fire—Banksia serrata (canopy seed bank) and Telopea speciosissima (transient seed bank) by planting their seeds both early and late in the post-fire recruitment period (PRP) and manipulating litter density in orthogonal treatments. Vertebrate seed predators were excluded. Both species established more seedlings late in the PRP, although results were strongly influenced by very poor establishment at one site. Invertebrate seed predators consumed more T. speciosissima seeds in sites early (69.5%) than late in the PRP (51.2%), while consumption of B. serrata seeds was lower overall and comparable across sites (average 47.3%). Surprisingly, litter had very little effect on establishment and none on invertebrate seed predation, suggesting that other factors are more important. Recruitment was only loosely coupled to disturbance for the canopy seed bank species; for the transient seed bank species, the coupling was tighter but separated in time from the disturbance. Understanding both the strength and temporal aspects of the disturbance–recruitment coupling is necessary for appropriate management of plant functional diversity in fire-prone habitats.     

Importance of the soil seed bank of disturbed sites in Chilean matorral in early secondary succession

Abstract. The dynamics of the seed bank may provide clues to the process of recovery of the vegetation of disturbed sites. The role of the seed bank may be more important in areas with a seasonal climate than in areas where seedling recruitment is not limited to one season. We studied the seed bank and the seed rain in three sites of the Chilean mediterranean-climate region (33° 48'S) which differed in the degree of anthropic disturbance: a closed-canopy, second-growth forest; an open matorral; and an old-field. Additionally, we tested the germination of seeds from the soil and from the current-year seed crop. The seed bank varied considerably between the two years of study, although no change in the vegetation could be observed. Seed density and species richness were lower in 1989 than in 1988. The seed bank of the second-growth forest changed less between years, while the old-field showed the largest change. The highest seed rain occurred under shrub patches in the open matorral, while few seeds fell in the spaces between shrub clumps or in the old-field. In the forest, seed rain was low and correlated with species cover. Germination was low (0 - 15%) in tests using either soil samples or fresh seeds. These results indicate that matorral succession is a very slow process, limited mainly by low germination and low arrival of propagules to open areas. Most woody species have animal-disseminated fleshy propagules. The presence of established shrubs which may serve as perches or refuges for animals increases species richness in the seed rain and the seed bank.