SEED GERMINATION TIMING PATTERNS IN INTERMOUNTAIN PENSTEMON (SCROPHULARIACEAE)

Regulation of seed germination timing is an important component of population persistence, especially for short-lived plants in seasonal environments with unpredictable year-to-year weather variation. Seed germination patterns were examined in the laboratory for seeds from 135 populations belonging to 38 species and 13 sections of the genus Penstemon (Scrophulariaceae), from habitats ranging from warm desert to alpine tundra. Seeds of most species were dormant at dispersal and required a period of moist chilling to become germinable. Response to chilling was related to probable chilling duration at the collection site. Populations from habitats with severe winters produced seeds with long chilling requirements, while those from habitats with mild winters produced seeds with short chilling requirements. Populations from midelevation habitats produced seeds with intermediate chilling requirements but with a sizeable fraction whose dormancy was not broken by chilling of any length. Another pattern observed mostly in warm desert populations was little primary dormancy at autumn temperatures combined with induction of a fraction into secondary dormancy by short chilling. Species with a wide habitat range included populations with contrasting germination patterns. Parallel habitat-correlated patterns were observed in different sections. Most species showed germination patterns that combined predictive mechanisms with potential for carryover of a persistent seed bank. Results of common garden experiments suggested that germination differences had a strong genetic basis both among populations and among plants in a population. Adaptive radiation in the genus Penstemon has thus included the evolution of habitat-specific germination timing strategies in multiple lineages within the genus.     

Thermoregulation and microhabitat use in mountain butterflies of the genus Erebia: Importance of fine-scale habitat heterogeneity

Mountain butterflies have evolved efficient thermoregulation strategies enabling their survival in marginal conditions with short flight season and unstable weather. Understanding the importance of their behavioural thermoregulation by habitat use can provide novel information for predicting the fate of alpine Lepidoptera and other insects under ongoing climate change. We studied the link between microhabitat use and thermoregulation in adults of seven species of a butterfly genus Erebia co-occurring in the Austrian Alps. We captured individuals in the field and measured their body temperature in relation to microhabitat and air temperature. We asked whether closely related species regulate their body temperature differently, and if so, what is the effect of behaviour, species traits and individual traits on body to air and body to microhabitat temperature differences. Co-occurring species differed in mean body temperature. These differences were driven by active microhabitat selection by individuals and also by species–specific habitat preferences. Species inhabiting grasslands and rocks utilised warmer microclimates to maintain higher body temperature than woodland species. Under low air temperatures, species of rocky habitats heated up more effectively than species of grasslands and woodlands which allowed them to stay active in colder weather. Species morphology and individual traits play rather minor roles in the thermoregulatory differences; although large species and young individuals maintained higher body temperature. We conclude that diverse microhabitat conditions at small spatial scales probably contribute to sympatric occurrence of closely related species with different thermal demands and that preserving heterogeneous conditions in alpine landscapes might mitigate detrimental consequences of predicted climate change.     

Heat shock increases the reliability of a temperate calcareous grassland seed bank study

Reliable assessment of the density and species richness of the viable seeds in the soil is essential to estimate the probability of successful restoration of a particular plant community. Since temperate calcareous grasslands contain several thermophilous species typical of fire prone habitats, heat shock can be expected to break dormancy of target species for calcareous grassland restoration. We tested the effect of heat shock on the estimated density and species richness of seeds in soil samples. Heat shock treatments between 70 and 110°C resulted in both a higher estimated seed density and estimated species richness, in particular of specialist calcareous grassland species. Moreover, 25 species germinated exclusively from the heat-treated samples. This indicates that several species would have been missed by handling soil seed bank samples following the standard germination protocol, resulting in an underestimation of the restoration potential.     

盐生植物角果碱蓬种子二型性对环境的适应策略

角果碱蓬(Suaeda corniculata)是藜科一年生盐生植物, 在我国分布于北方盐碱滩涂和盐碱荒漠地区。角果碱蓬具有棕色和黑色两种异型体种子(简称棕色和黑色种子)。对采自内蒙古鄂托克前旗盐渍化生境的角果碱蓬二型种子的形态、休眠和萌发特性开展对比研究, 测定了二型种子休眠和萌发行为对温度、光照和盐分(NaCl)的响应, 以揭示盐生植物异型种子对温带盐漠生境的适应对策。结果表明: (1)二型性种子在大小、种皮特性和结实比例方面有显著差异。与黑色种子相比, 棕色种子个体较大, 种皮透水性强。黑色种子与棕色种子的结实比例约为5.6 : 1。(2)新成熟的棕色种子的萌发对各温度梯度和光照条件不敏感, 萌发率较高(84%-100%); 而新成熟的黑色种子萌发率较低(8%-78%), 萌发对光照敏感。(3)黑色种子具有浅度生理休眠, 种皮划破、赤霉素处理和低温层积均可有效地提高种子的萌发率。(4)二型种子萌发对土壤盐分的胁迫具有不同的响应。与黑色种子相比, 棕色种子对盐分胁迫不敏感, 在较高的盐分浓度下仍有较高的萌发率, 低温层积处理能够降低黑色种子对盐胁迫的敏感性, 有效地提高种子的初始萌发率、萌发恢复率和最终萌发率。角果碱蓬二型种子不同的形态、休眠和萌发特性, 提高了该物种在高度异质性生境中的适合度, 对种群成功地适应温带盐漠环境具有重要的意义。     

Genetic differentiation among populations of Sesleria albicans Kit. ex Schultes (Poaceae) from ecologically different habitats in central Europe

As observed for many other plant species, the populations of Sesleria albicans in Central Europe are located in habitats, which differ to a high degree from each other with regard to ecological factors such as nutrients, light and water as well as in type of land use. The species colonizes limestone cliffs, pavements, screes, grazed and mown grasslands, heaths, fens and open woodlands. In this study, we used random amplified polymorphic DNA (RAPD) analysis to investigate the genetic differentiation among 25 populations of S. albicans from six different types of habitat (beech forests, alpine and lowland rocky ridges, lowland screes, fens, calcareous grasslands). With RAPD analysis, 344 fragments could be amplified, of which 95.9% were polymorphic. The level of polymorphism ranged from 29.7 to 56.7% polymorphic bands per population and was correlated with population size. In an analysis of molecular variance (AMOVA), used to detect variation among individuals within populations, among populations from the same habitat and among different habitats, most of the genetic variation was found within populations (62.06%) and among populations from the same habitat (33.36%). In contrast, only a very low level of differentiation could be observed among different habitats (4.58%). The results of our study give only little evidence for an ecotypic differentiation of Sesleria albicans. This differentiation is principally conceivable, but obviously not related to the investigated RAPD loci.     

The impact of temperature regimes on development,dormancy breaking and germination of dwarf shrub seeds from arctic,alpine and boreal sites

It has been suggested that the infrequent sexual reproduction of arctic dwarf shrubs might be related to the harsh environmental conditions in which they live. If this is the case, then increases in temperature resulting from global climate change might drastically affect regeneration of arctic species. We examined whether recruitment of Empetrum nigrum ssp. hermaphroditum and Vaccinium uliginosum (hereafter E. nigrum and V. uliginosum) was affected by temperature during three reproductive stages: seed development, dormancy breakage and germination. Seeds were collected from an arctic, an alpine (only E. nigrum) and a boreal site with different climates; stored at different winter temperatures and incubated for germination at different temperatures. Seeds of V. uliginosum developed in the boreal region had a higher percentage germination than did seeds developed in the Arctic. In contrast, seeds of E. nigrum from the arctic site had a higher or similar percentage germination than did seeds from the alpine and boreal sites. Increased winter temperatures had no significant effect on resulting germination percentage of E. nigrum. However, V. uliginosum seeds from the arctic site suffered increased fungal attack (and thus decreased germination) when they were stratified under high winter temperatures. Seeds of both species increased germination with increased incubation temperatures. Our results suggest that both species would increase their germination in response to warmer summers. Longer summers might also favour the slow-germinating E. nigrum. However, increased winter temperatures might increase mortality due to fungal attack in V. uliginosum ecotypes that are not adapted to mild winters.     

Why some species cannot colonise restored habitats? The effects of seed and microsite availability

Restoration of calcareous grasslands was promoted as a conservation strategy to reduce the risks imposed by habitat loss and fragmentation. Restoration already provided promising results for several taxa, however some specialist species still fail at colonising restored habitats. We aimed at explaining this lack of colonisation success for three calcareous grasslands specialist species in southern Belgium: Pulsatilla vulgaris; Trifolium montanum; and, Veronica prostrata. We studied: (i) germination in control and outdoor conditions (cold, heat, smoke and litter effects); (ii) in situ seedling emergence patterns (effects of seed addition and germination microsites availability). The three species were able to germinate in Petri dishes in the absence of treatment. Cold enhanced the germination of V. prostrata. Fire-related treatments (heat shock and smoke exposure) did not enhance germination and were deleterious to V. prostrata. Litter cover improved P. vulgaris emergence in outdoor containers, but had a negative effect on V. prostrata. In the field, V. prostrata did not emerge. T. montanum seedlings were observed in the reference grasslands when seeds were added, but not in the restored grasslands. P. vulgaris emerged in the reference grasslands, and to a lower degree in the restored grasslands. The combination of seed addition and microsites availability for seed germination resulted in enhanced seedling emergence for P. vulgaris. Our results suggest that seed and microsite availability can be limiting factors for site colonisation, but the combination of both is likely much more limiting. Lower seedling emergence in restored than in reference grasslands suggests a lower habitat quality in restored grasslands.     

Diversity of germination strategies and seed dormancy in herbaceous species of campo rupestre grasslands

The effects of fire on the vegetation vary across continents. However, in Neotropical fire‐prone grasslands, the relationship between fire and seed germination is still poorly understood, while their regeneration, especially after strong anthropogenic disturbance, is challenging for their conservation. In the present study, we assessed diversity of germination strategies in 15 dominant herbaceous species from Neotropical altitudinal grasslands (locally known as campos rupestres). We exposed seeds to several fire‐related treatments. We also compared germination between regularly and post‐fire fruiting species. Finally, we investigated the diversity of dormancy classes aiming at better understanding the biogeography and phylogeny of seed dormancy. Germination strategies varied among families. Velloziaceae and Xyridaceae produced non‐dormant, fast‐germinating seeds. Cyperaceae and Poaceae showed an extremely low or null germination due to a high proportion of unviable or embryo‐less seeds. The seeds of campo rupestre grasslands are fire resistant, but there is no evidence that fire triggers germination in this fire‐prone ecosystem. Although heat and charred wood did not promote germination, smoke enhanced germination in one grass species and decreased the mean germination time and improved synchrony in Xyridaceae and Velloziaceae. Fire had a positive effect on post‐fire regeneration by stimulating fruit set in some Cyperaceae and Poaceae species. These species produced faster germinating seeds with higher germination percentage and synchrony compared to regularly fruiting Cyperaceae and Poaceae species. This strategy of dispersion and regeneration seems to be an alternative to the production of seeds with germination triggered by fire. Physiological dormancy is reported for the first time in several clades of Neotropical plants. Our data help advance the knowledge on the role of fire in the regeneration of Neotropical grasslands.     

Divergence in germination traits among arctic and alpine populations of <Emphasis Type="Italic">Koenigia islandica</Emphasis>: light requirements

Light is known to regulate conservative germination strategies and the formation of seed banks. Although these strategies are crucial to survival in tundra environments—especially for annuals—light requirements for germination in arctic–alpine species are seldom investigated. Furthermore, environmental differences between arctic and alpine regions are expected to lead to evolutionary divergence among conspecific populations in seed germination strategies. In this study, we report important differences in germination light requirements among six arctic and alpine populations of the annual Koenigia islandica. Light had little effect on germination of the seeds from Iqaluit (Nunavut, Canada), Yukon (Canada), and Jasper (Alberta, Canada), whereas the seeds from the most severe climates, Svalbard (Norway) and Colorado (USA), had strong light requirements. Stratification of the seeds had little influence on their germination light requirements, with the exception of the population from Dovre (Norway), in which it induced a strong light requirement. Possible adaptive explanations and some implications of these observed germination patterns are discussed.     

Species-rich Nardus stricta grasslands host a higher vascular plant diversity on calcareous than on siliceous bedrock

Background: Species-rich Nardus stricta grasslands are a priority habitat for conservation in Europe. They typically occur on siliceous substrates and less frequently are found on calcareous bedrock.

Aims: The present paper aimed to identify the environmental factors (i.e. bedrock type, topographic, and climatic factors) that are related with community diversity and to assess if differences in plant diversity between N. stricta communities on calcareous and siliceous bedrock occur. We hypothesised that Nardus grasslands on calcareous bedrock hosted a higher vascular plant diversity than those on siliceous bedrock.

Methods: Based on 579 vegetation surveys carried out in the south-western Alps, we assessed vascular plant diversity (species richness, Shannon diversity, and Pielou’s equitability index) of species-rich Nardus grasslands and compared it between N. stricta communities on calcareous and siliceous bedrock.

Results: Elevation was identified as the main factor related to species composition, while species diversity was mostly related to mean annual precipitation and bedrock type. Species richness, Shannon diversity, and Pielou’s equitability index were higher within the communities on calcareous rather than on siliceous bedrock and a total of 89 and 34 indicator species were detected, respectively.

Conclusions: Based on our results, we suggest to protect primarily, as a habitat of priority interest, N. stricta grasslands on calcareous substrates for the higher vascular plant diversity hosted.     

No positive correlation between species and genetic diversity in European alpine grasslands dominated by Carex curvula

The distributional range of alpine plants experienced dramatic changes during the Quaternary ice ages. These changes offer many opportunities for studying the impact of habitat contraction and fragmentation on both species and genetic diversity. Here, we examined the amplified fragment length polymorphism-based genetic diversity in the sedge Carex curvula All. in relation to the species diversity of siliceous European alpine grasslands in which the sedge is dominant. We found no relationship or even a negative relationship between genetic and species diversity indices. Local species richness was associated with the regional pool size of siliceous alpine species, which was itself dependent on the extant area of suitable habitats for these species. Genetic diversity of C. curvula was primarily shaped by the presumed location of glacial refugia and the routes of post-glacial colonization. We conclude that the two levels of diversity are not positively correlated because genotypes and species do not respond similarly to the spatial dynamics of suitable habitats induced by Quaternary temperature changes.     

Asymmetric carabid beetle spillover between calcareous grasslands and coniferous forests

Central European calcareous grasslands are considered biodiversity hotspots, but are severely threatened by the change in land-use and by habitat fragmentation. Coniferous forests are typical adjacent habitats to calcareous grasslands, as abandoned calcareous grasslands are often afforested or develop into coniferous forests by succession. To investigate spillover between calcareous grasslands and coniferous forests, a total of 144 pitfall traps for carabid beetles were placed at three different distances (1, 5, 20 m) from the edge in both habitats at eight locations from April to late August. We found that both habitats had a distinct species assemblage and a decrease in spillover with increasing distance from the habitat edge into the adjacent habitat. Calcareous grasslands were more affected by spillover from the adjacent coniferous forests than vice versa because more forest specialists penetrated into calcareous grasslands than grassland specialists penetrated into coniferous forests. We conclude that spillover into small and isolated habitats can severely change species assemblages, which has to be considered in conservation measures. The protection of large sites with small edge-interior ratios can reduce negative effects on species assemblages in endangered calcareous grasslands.     

Hybrid ecosystems can contribute to local biodiversity conservation

Calcareous grasslands have become severely threatened habitats in Europe. The aim of this study was to investigate the changes in plant species richness, and functional and phylogenetic diversity in northern Estonian calcareous (alvar) grasslands resampled after 90 years of land-use change. Functional traits characterizing species that have benefited most from decreased habitat area and altered environmental conditions, and additional species that can potentially inhabit the remaining grassland patches were identified. Also changes in the relative amount of habitat-specific species were studied to detect a possible decrease in habitat integrity. Although grasslands in the studied region had lost most of their original area (~90 %), species richness had substantially increased due to invasion by more competitive, nutrient-demanding native species. Functional diversity generally increased, whereas phylogenetic diversity showed no response to altered conditions. Overall, these grasslands have lost their integrity as calcareous grassland habitat type in the region, because the relative amount of habitat-specific characteristic species has declined significantly. However, although the grasslands have transformed to a ‘hybrid’ habitat type and restoration to their previous state is likely not reasonable, such degraded species-rich grassland fragments can still be recognized as important habitats to preserve high local biodiversity and several characteristic species of calcareous grasslands. As current landscapes consist of an increasing number of hybrid and novel communities, new tools to supplement traditional conservation or restoration practices are necessary to recognize and maintain regions and habitats of high local biodiversity.     

Seed bank dynamics of the desert cactus Opuntia rastrera in two habitats from the Chihuahuan Desert

In desert environments the main input to the seed bank of many succulents is the seed rain through zoochory while high levels of granivory by rodents, birds and ants are the main cause of subsequent losses. In the patchy environment of arid lands the characteristics of both processes may vary between habitats causing differences in the recruitment of new genets. To test this hypothesis we used populations of the desert cactus Opuntia rastrera which has different recruitment rates in the two adjacent habitats where it grows. In Opuntia-dominated scrublands (nopaleras, density ca. 4,000 plants/ha) 1 seedling out of 7,000,000 seeds establish, whereas in grasslands (density ca. 100 plants/ha) this ratio is 1:20,000. From 1996 until 1998 the seed rain, seed removal by granivores and seed abundance in the soil were monitored in both habitats. Results showed striking differences in the dynamics of the seed bank of both habitats. Seed rain was 8.5 times bigger in nopaleras than in grasslands. In nopaleras most seeds were removed by rodents while the quantities of seeds removed by rodents, birds and ants in grasslands were similar. One year after dispersal (the time necessary to break seed dormancy) only 6% of original nopalera seeds and 12% of grassland seeds remained. After germination trials only 1% (ca. 15,000 seeds/ha) and 2% (ca. 2,500 seeds/ha) respectively were viable. These differences in the effective seed bank (6 times bigger in nopaleras) can not explain the differences in genet recruitment (which is several orders of magnitude bigger in grasslands). Apparently the between habitat difference in nurse plant availability and in rodent density (which inflict a strong hervibory upon seedlings) can explain the differences in genet recruitment. It is speculated that this between habitat difference in genet recruitment suggests that the species evolved in less extreme environments (e.g. grasslands) than desert scrublands which, in turn, are colonised due to the singular ability of O. rastrera for vegetative propagation. This revised version was published online in August 2006 with corrections to the Cover Date.     

温带半干旱地区一年生植物种子的萌发特性

在实验室条件下研究了中国温带半干旱地区科尔沁沙地的23种1年生植物的种子萌发特性（新采集种子、冷藏和干藏种子）.大籽蒿、虎尾草、冠芒草、沙蓬和地锦的新种子萌发率达90%左右,11种植物新种子萌发率均低于70%,说明这些植物的新种子具有或多或少的休眠属性.经过150d的冷干藏后,大籽蒿、虎尾草萌发率保持在90%以上,说明这两种植物完全没有休眠机制;冠芒草、沙蓬和地锦的种子萌发率下降较多,可能是储藏的环境条件导致的2次休眠现象;冷藏和干藏处理均能使绿珠藜、毛马唐、细叶益母草、雾冰藜、金狗尾草、苋菜、马齿苋、碱地肤和水稗草的种子在生长季开始时完成生理后熟,萌发率达到80%以上;干藏有利于促进毛马唐、细叶益母草、马齿苋和鹤虱的种子成熟,冷藏有利于促进绿珠藜和金狗尾草的种子成熟;黄蒿、灰绿藜、画眉草和烛台虫实在不同处理下的萌发率都比较低,说明种子内在生理休眠作用较强,具有减少种子一次性萌发数量的风险分摊策略.大多数1年生植物均能在较短时间内达到最终萌发率的90%,表现出迅速萌发的特性;黄蒿、灰绿藜、碱地肤和沙蓬种子则在不同处理中表现出延长萌发时间的策略来适应半干旱地区不确定的环境条件.最后,探讨了几种主要1年生植物的种子萌发对策与其对环境适应机制之间的关系.     

Seed bank persistence of clonal weeds in contrasting habitats: implications for control

The ability of weeds to form a seed bank is important for their population dynamics and management because it provides a refuge enabling reinvasion after established target plants have died. However, knowledge of the differential seed behaviour of individual species over multiple years and varying environmental conditions is surprisingly rare but necessary for effective control of diverse weed populations. We established a seed burial experiment in alpine habitats differing in management regime (i.e., forest, hay meadow and pasture) to determine whether seeds of the unpalatable perennial weeds, Veratrum album (white hellebore) and Gentiana lutea (yellow gentian) were able to delay germination and remain viable over 3 years. Our study shows that both species formed a short-term persistent seed bank; in the third-year, the soil seed banks of both species were nearly depleted, having declined to <5% of their original size. Both species had strikingly different germination strategies: G. lutea seeds mainly germinated in their first-year, whilst the majority of V. album seeds germinated in their second-year. The fraction of dormant G. lutea seeds increased with seed age, indicating that seeds remained viable after forgoing germination in the previous year. Habitat-specific differences in seed germination increased with seed age, with germination fractions being lowest in moist hay meadows. This suggests that the negative effects of anoxic conditions became more pronounced as seeds aged in hay meadows. Conversely, seed dormancy was equal among habitats. The absence of a long-term persistent seed bank has important implications for the management of both nuisance and endangered-plant populations. In the case of V. album and G. lutea, re-colonization of habitats from the seed bank is unlikely after established plants have been removed.     

重要藏药川西獐牙菜种子萌发的研究

比较了温度、生长素(赤霉素GA3)和储藏条件对川西獐牙菜(SwertiamussotiiFranch)高海拔野生自然种群和低海拔栽培后种子发芽率的影响。未经任何处理的高海拔野生自然种群种子的发芽率明显高于低海拔栽培种群。无论是赤霉素处理还是低温处理对种子的发芽率都有显著提高。经过处理后,两种来源的种子最终发芽率没有明显的变化。结果表明:(i)野生的或栽培的川西獐牙菜种子都存在休眠现象;(ii)通过引种栽培不能打破川西獐牙菜种子的休眠,该机制可能是受遗传因素的控制;(iii)赤霉素处理和4℃低温冷藏对打破种子的休眠具有重要作用。     

VARIATION IN GERMINATION RESPONSE TO TEMPERATURE IN RUBBER RABBITBRUSH (CHRYSOTHAMNUS NAUSEOSUS: ASTERACEAE) AND ITS ECOLOGICAL IMPLICATIONS

Seed collections from 72 rubber rabbitbrush populations occupying a range of habitats in western North America were incubated at 3 C in the laboratory. Collections from warm desert habitats required less than 2 weeks to achieve 90% relative germination under these conditions, while collections from montane habitats showed delayed germination requiring up to 20 weeks. When 13 representative collections were incubated at constant temperatures from 5 to 30 C, all germinated completely at 30 C within 4 weeks. Collections from warm desert habitats germinated rapidly over the whole range of temperatures. Montane collections sometimes exhibited dormancy at intermediate temperatures (15 and 25 C) even though they were ultimately able to germinate at lower temperatures. Results suggest that dormancy is conditional and temperature-dependent in this species. Chilling the seeds extends the temperature range for germination downward to include the chill temperature itself. Germination response to temperature and its variation as a function of habitat are of apparent adaptive significance, serving to time germination so that the probability of seedling survival is maximized in each habitat. Within populations, response to temperature varied as a function of year of harvest and of within-year harvest date, indicating that germination patterns are probably not under rigid genetic control but represent an integration of genetic and environmental factors.     

Stock grazing of semi-natural habitats on National Trust land

Stock grazing is essential in many habitats owned by the National Trust if their nature conservation interests are to be maintained or enhanced. In this review, the benefits of grazing cattle, sheep and other livestock in different habitat types are described, together with constraints imposed by tenancy agreements and difficulties in obtaining the most appropriate animal. The majority of upland heaths, calcareous grasslands and lowland/coastal heaths on Trust land are being grazed satisfactorily. However, the nature conservation interest of parklands is currently being eroded in part because of inappropriate grazing regimes. It is argued that the best use of stock in the future is in extensive grazing regimes.     

Seed dormancy in alpine species

In alpine species the classification of the various mechanisms underlying seed dormancy has been rather questionable and controversial. Thus, we investigated 28 alpine species to evaluate the prevailing types of dormancy. Embryo type and water impermeability of seed coats gave an indication of the potential seed dormancy class. To ascertain the actual dormancy class and level, we performed germination experiments comparing the behavior of seeds without storage, after cold-dry storage, after cold-wet storage, and scarification. We also tested the light requirement for germination in some species. Germination behavior was characterized using the final germination percentage and the mean germination time. Considering the effects of the pretreatments, a refined classification of the prevailing dormancy types was constructed based on the results of our pretreatments. Only two out of the 28 species that we evaluated had predominantly non-dormant seeds. Physiological dormancy was prevalent in 20 species, with deep physiological dormancy being the most abundant, followed by non-deep and intermediate physiological dormancy. Seeds of four species with underdeveloped embryos were assigned to the morphophysiologial dormancy class. An impermeable seed coat was identified in two species, with no additional physiological germination block. We defined these species as having physical dormancy. Light promoted the germination of seeds without storage in all but one species with physiological dormancy. In species with physical dormancy, light responses were of minor importance. We discuss our new classification in the context of former germination studies and draw implications for the timing of germination in the field.