Seasonal cycle of seed dormancy controls the recruitment of Butia odorata (ARECACEAE) seedlings in savanna‐like palm tree formations in southern Brazil

Butia odorata (Barb. Rodr.) Noblick is a palm tree that grows in savanna‐like formations in subtropical regions of South America, and whose regeneration is threatened by agricultural management. Its diaspores are dormant after dispersal which takes place during the summer and early autumn. The aim of this study was to investigate seasonal and microhabitat effects on the germination and seedling recruitment of this palm species. Diaspores were sown in the field, in both open lands and forest patches. During 2 years, we measured seed germination, viability and moisture, seedling emergence and germination response to warm stratification of those seeds that failed to germinate in the field. Germination was concentrated during the summer, when soil temperatures were highest, whilst seedling emergence peaked in the autumn and early winter, when temperature and humidity conditions became less extreme. In open lands, there were two pulses of germination (first and second summer), whilst in forest patches, a single pulse (second summer) was detected. Although overall germination did not differ between microhabitats, the percentage of seedling emergence from seeds that remained buried until the end of the experiment was almost twice as large in the forest patches compared with open areas. The viability of seeds declined over time, particularly in open areas. Laboratory‐induced warm stratification was found to act on seed dormancy release in a cyclic way, being far more effective on seeds retrieved from the field in spring–summer months than in those retrieved in the winter. This cyclic pattern of dormancy in B. odorata seeds results in major seedling recruitment after the summer, under wetter and cooler conditions, thus reducing mortality risk. This process can be enhanced by the presence of surrounding vegetation, which both increases seedling emergence and/or prolongs seed viability.     

Ecophysiology of embryo development and seed germination of the European woodland herbaceous perennial Corydalis cava (L.) Schweigg. & Körte subsp. cava (Fumariaceae)

In this study we examined the germination ecology with special reference to the temperature requirements for embryo development and germination of Corydalis cava subsp. cava, under both outdoor and laboratory conditions. Corydalis cava is a spring flowering woodland tuberous geophyte widely distributed across Europe. Germination phenology, including embryo development and radicle and cotyledon emergence, was investigated in a population growing in northern Italy. Immediately after harvest, seeds of C. cava were sown both in the laboratory under simulated seasonal temperatures and naturally. Embryos, undifferentiated at the time of seed dispersal, grew during summer and autumn conditions, culminating in radicle emergence in winter, when temperatures fell to ca 5°C. Cotyledon emergence also occurred at ca 5°C, but first emergence was delayed until late winter and early spring. Laboratory experiments showed that high (summer) followed by medium (autumn) and low temperatures (winter) are needed for physiological dormancy loss, embryo development and germination respectively. Unlike seeds of C. cava that germinated in winter, in other Corydalis species radicle emergence occurred in autumn (C. flavula) or did not depend on a period of high summer temperature to break dormancy (C. solida). Our results suggest that subtle differences in dormancy and germination behavior between Corydalis species could be related to differences in their geographical distribution.     

Temperature controls seed germination and dormancy in the European woodland herbaceous perennial Erythronium dens-canis (Liliaceae)

We examined the germination ecology and the temperature requirements for germination of Erythronium dens-canis, under both outdoor and laboratory conditions. E. dens-canis is a spring flowering woodland geophyte widely distributed across Europe. Germination phenology, including embryo development and radicle and cotyledon emergence, were investigated in a natural population growing in Northern Italy. Immediately after harvest, seeds of E. dens-canis were either sown on agar in the laboratory under simulated seasonal temperatures or placed in nylon mesh sachets and buried in the wild. Embryos, undifferentiated at the time of seed dispersal, grew during summer and autumn conditions in the laboratory and in the wild, culminating in radicle emergence in winter when temperatures fell to ≈ 5 °C. Emergence of cotyledons did not occur immediately after radicle emergence, but was delayed until the end of winter. Laboratory experiments showed that temperature is the main factor controlling dormancy and germination, with seeds becoming non-dormant only when given warmth, followed by cold stratification. Unlike seeds of E. dens-canis that germinate in winter, in other Erythronium species radicle emergence occurs in autumn, while in some it is delayed until seeds are transferred from winter to spring conditions. Our results suggest that there is genetic and environmental control of the expression of seed dormancy amongst Erythronium species, which is related to local climate.     

ENVIRONMENTAL CONTROL OF SEED GERMINATION IN THLASPI ARVENSE (CRUCIFERAE)

The effect of environmental conditions during storage and imbibition on germination was investigated in field pennycress (Thlaspi arvense L.), a weed species that can behave as a winter or a summer annual. Freshly harvested seeds of an inbred line with a cold requirement for flowering exhibited primary dormancy that was rapidly lost following 1 month of afterripening in a dry state. Nondormant seeds were positively photoblastic. The light effect was mediated through phytochrome since germination was promoted by red light and inhibited by far red light. Seedling emergence was also inhibited by light filtered through a canopy of wheat leaves. Germination of field pennycress seeds was considerably more sensitive to moisture stress than two sympatric species, wild oat (Avena fatua L.) and wheat (Triticum aestivum L., cv. ERA). Seeds of the latter two species were chosen in order to compare the effect of water potential on germination in field pennycress with that in sympatric species. It was concluded that the major environmental factor limiting nondormant field pennycress seeds on the soil surface was water availability. Imbibition of fully afterripened seeds at low temperatures (6 C) induced a deep secondary dormancy. In contrast to primary dormancy, cold-induced dormancy was not alleviated by red light, alternating temperatures (21/5 C), or 2 months of dry storage at 6, 15, or 35 C. However, exogenous gibberellin A₃ or 24 weeks of dry storage resulted in germination in cold-induced dormant seeds. Secondary dormancy was not observed in fully afterripened seeds that were preincubated at 21 C for 1 or 2 days prior to the cold treatment. These results may explain the failure in field experiments to observe the cold-induced secondary dormancy that limits spring emergence in other winter annuals (J. Baskin, C. Baskin, Weed Res. 1979 19: 285–292).     

GERMINATION ECOLOGY OF SEDUM PULCHELLUM MICHX. (CRASSULACEAE)

Factors controlling the timing of seed germination were investigated in the small succulent winter annual Sedum pulchellum Michx. (Crassulaceae) in its natural habitat on unshaded limestone outcrops in northcentral Kentucky. At maturity in early July the dormant seeds are not dispersed but are retained in the fruits on the standing dead plants until September and October. Many, but not all, of the seeds afterripen in the fruits during summer, and at the time of dispersal some of them are dormant and some are nondormant. Germination and annual population establishment occur in September and October from seed reserves that have been in the soil for one or more years and from seeds produced in the current year. Germination of nondormant seeds may be prevented in autumn by lack of the appropriate combination of environmental factors including light, temperature and soil moisture in the seed's microsite. The effect of low winter temperatures on ungerminated seeds in the population is to induce nondormant seeds into secondary dormancy and to prevent afterripening of dormant seeds. Thus, in spring all the seeds in the population's seed reserve are dormant. During spring and summer some of these seeds afterripen, and they germinate in autumn when, and if, germination requirements are fulfilled.     

The dual role of temperature in the regulation of the seasonal changes in dormancy and germination of seeds of Polygonum persicaria L.

Summary The role of temperature in the regulation of seasonal changes in dormancy and germination was studied in seeds of Polygonum persicaria. Seeds were buried in the field and under controlled conditions. Portions of seeds were exhumed at regular intervals and germination was tested over a range of conditions. Seeds of P. persicaria exhibited a seasonal dormancy pattern that clearly showed the typical features of summer annuals, i.e. dormancy was relieved at low winter temperatures, the germination peak occurred in spring and dormancy was re-induced in summer. The expression of the dormancy pattern was influenced by the temperature at which germination was tested. At 30°C exhumed seeds germinated over a much longer period of the year than at 20° or 10°C. Nitrate added during the germination test occasionally stimulated germination. The seasonal changes in dormancy of buried seeds were regulated by the field temperature. Soil moisture and nitrate content did not influence the changes in dormancy. The fact that, on the one hand, field temperature determined the changes in dormancy and, on the other hand, germination itself was influenced by temperature, was used to describe the seasonal germination pattern of P. persicaria with a model. Germination of exhumed seeds in Petri dishes at field temperature was accurately described with this model. Germination in the field was restricted to the period where the range of temperatures over which germination could proceed (computed with the model) and field temperature overlapped.     

Light,temperature, dry after‐ripening and salt stress effects on seed germination of Phleum sardoum (Hackel) Hackel

Phleum sardoum is an endemic psammophilous species of Sardinia, growing exclusively on coastal sandy dunes. The effect of glumes on seed germination, germination requirements at constant (5–25°C) and alternating (25/10°C) temperatures, both in the light (12/12 h) and in the dark were evaluated, as well as the effect of a dry after‐ripening period (90 days at 25°C), the salt stress effect (0–600 mmol NaCl) and its recovery on seed germination. The presence of glumes reduced final germination percentages. For fresh naked seeds, high germination percentages were observed at 10°C. Dry after‐ripening increased germination rate at low temperatures, but did not affect final germination percentages. NaCl determined a secondary salt‐induced dormancy which recovery interrupted only partially. Our results highlighted that this species has its optimum of germination during autumn–winter when, under a Mediterranean climate, water availability is highest and soil salinity levels are minimal.     

The changing window of conditions that promotes germination of two fire ephemerals, Actinotus leucocephalus (Apiaceae) and Tersonia cyathiflora (Gyrostemonaceae)

BACKGROUND AND AIMS: Following a period of burial, more Actinotus leucocephalus (Apiaceae) and Tersonia cyathiflora (Gyrostemonaceae) seeds germinate in smoke water. The main aim of this study was to determine whether these fire-ephemeral seeds exhibit annual dormancy cycling during burial. This study also aimed to determine the effect of dormancy alleviation on the range of light and temperature conditions at which seeds germinate, and the possible factors driving changes in seed dormancy during burial. METHODS: Seeds were collected in summer, buried in soil in mesh bags in autumn and exhumed every 6 months for 24 months. Germination of exhumed and laboratory-stored (15 degrees C) seeds was assessed at 20 degrees C in water or smoke water. Germination response to light or dark conditions, incubation temperature (10, 15, 20, 25 and 30 degrees C), nitrate and gibberellic acid were also examined following burial or laboratory storage for 24 months. In the laboratory seeds were also stored at various temperatures (5, 15, 37 and 20/50 degrees C) for 1, 2 and 3 months followed by germination testing in water or smoke water. KEY RESULTS: The two species exhibited dormancy cycling during soil burial, producing low levels of germination in response to smoke water when exhumed in spring and high levels of germination in autumn. In autumn, seeds germinated in both light and dark and at a broader range of temperatures than did laboratory-stored seeds, and some Actinotus leucocephalus seeds also germinated in water alone. Dormancy release of Actinotus leucocephalus was slow during dry storage at 15 degrees C and more rapid at higher temperatures (37 and 20/50 degrees C); weekly wet/dry cycles further accelerated the rate of dormancy release. Cold stratification (5 degrees C) induced secondary dormancy. By contrast, no Tersonia cyathiflora seeds germinated following any of the laboratory storage treatments. CONCLUSIONS: Temperature and moisture influence dormancy cycling in Actinotus leucocephalus seeds. These factors alone did not simulate dormancy cycling of Tersonia cyathiflora seeds under the conditions tested.     

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.     

DORMANCY AND GERMINATION IN SEEDS OF COMMON RAGWEED WITH REFERENCE TO BEAL'S BURIED SEED EXPERIMENT

Common ragweed (Ambrosia artemisiifolia L.) was one of 19 herbaceous weedy species used by Beal in his buried viable seed experiment started in 1879. No seeds germinated during the first 35 years of the experiment when germination tests were performed in late spring, summer or early autumn. Germination did occur in seeds buried for 40 years when seeds were exhumed and tested for germination in early spring. Data obtained in more recent research provide the probable explanation for these results. Seeds of common ragweed that do not germinate in spring enter secondary dormancy by mid to late spring and will not germinate until dormancy is broken the following late autumn and winter. Thus, during the first 35 years of the experiment seeds were dormant when tested for germination, whereas seeds buried for 40 years were nondormant. Seeds buried 50 years or longer did not germinate when tested in spring, probably because they had lost viability and/or seeds germinated during burial and seedlings died.     

Temperature effects on dormancy levels and germination in temperate forest sedges (Carex)

The effects of stratification temperatures and burial in soil on dormancy levels of Carex pendula L. and C. remota L., two spring-germinating perennials occurring in moist forests, were investigated. Seeds buried for 34 months outdoors, and seeds stratified in the laboratory at temperatures between 3 and 18 °C for periods between 2 and 28 weeks, were tested over a range of temperatures. Seeds of the two species responded similarly to stratification treatments, except for an absolute light requirement in C. pendula. Primary dormancy was alleviated at all stratification temperatures, but low temperatures were more effective than higher ones . (≥ 12 °C). Dormancy induction in non-dormant seeds kept at 5 °C occurred when seeds were subsequently exposed to 18 °C. Dormancy was not induced by a transfer to lower temperatures. Buried seeds of both species exhibited seasonal dormancy cycles with high germination from autumn to spring and low germination during summer. Temperatures at which the processes of dormancy relief and of dormancy induction occurred, overlapped to a high degree. Whether, and when, dormancy changes occurred depended on test conditions. The lower temperature limit for germination (> 10%) was 9 °C in C. remota and 15 °C in C. pendula. Germination ceased abruptly above 36 °C. Germination requirements and dormancy patterns suggest regeneration from seed in late spring and summer at disturbed, open sites (forest gaps) and the capability to form long, persistent seed banks in both species.     

Seed germination of exotic and native winter annuals differentially responds to temperature and moisture,especially with climate change scenarios

Seeds of winter annuals require a summer after-ripening period for dormancy loss and low autumn temperatures for germination. With current and future changes in moisture and temperature, we tested the effects of warming along a relative humidity (RH) gradient on dormancy loss and effects of decreased diurnal temperature range (DTR) on germination. We further reasoned that the effects of changes in these variables would be disproportionate between the exotic and native winter annuals. Seeds of exotic species (Buglossoides arvensis, Lamium purpureum and Ranunculus parviflorus) and co-occurring native species (Galium aparine, Paysonia stonensis and Plantago virginica) were collected in middle Tennessee. After-ripening occurred over a 15–100% RH gradient at 25 and 30°C and germination was tested at 20/10 and 20/15°C. Niche breadth was calculated using Levins' B. Fresh Ranunculus seeds had high germination and those of other species did not. Germination for these species increased with after-ripening, mostly across the RH gradient irrespective of temperature. A decrease in DTR showed mixed results – the extreme being Ranunculus with no germination at 20/15°C. Most exotic species had wider germination niche breadths than native species. With climate change, we suggest that a decrease in DTR may have a larger effect on germination than increasing moisture or warming on dormancy break. Moreover, there is not a clear-cut winner with climate change when we compare exotic versus native species because the responses of our six species were species specific.     

GERMINATION ECOPHYSIOLOGY OF ARENARIA GLABRA,A WINTER ANNUAL OF SANDSTONE AND GRANITE OUTCROPS OF SOUTHEASTERN UNITED STATES

The germination ecophysiology of Arenaria glabra Michx., a characteristic winter annual plant species of granite and sandstone outcrops of southeastern United States, was investigated. Seeds germinate in early autumn, plants overwinter in the rosette stage and then flower, set seeds, and die in late spring; seeds are dispersed soon after maturity. Eighty-five to 90% of freshly-matured seeds were innately dormant, and the other 10–15% germinated only at temperatures lower than those that occur in the habitat at the time of seed dispersal in June. During the summer after-ripening period, seeds stored dry under ambient laboratory conditions exhibited progressive increases in rates and total percentages of germination, a widening of the temperature range for germination, and a loss of the light requirement. At a 14-hr daily photoperiod, seeds kept on continuously moist soil germinated to 83% at simulated July and August temperatures during July and August, and the remainder germinated at September temperatures in September. On the other hand, seeds subjected to alternate wetting and drying during July and August germinated to only 9% during those 2 months, and the remainder germinated after the soil was kept continuously moist, beginning on 1 September, at simulated habitat temperatures during September and October. Thus, the timing of germination of A. glabra in the field is controlled by an interplay of the seeds' physiological state with the dynamics of temperature and soil moisture conditions.     

Ecophysiology of seed dormancy and the control of germination in early spring‐flowering Galanthus nivalis and Narcissus pseudonarcissus (Amaryllidaceae)

Seed dormancy induction and alleviation in the winter‐flowering, moist temperate woodland species Galanthus nivalis and Narcissus pseudonarcissus are complex and poorly understood. Temperature, light and desiccation were investigated to elucidate their role in the germination ecophysiology of these species. The effect of different seasonal temperatures, seasonal durations, temperature fluctuations, the presence of light during different seasons and intermittent drying (during the summer period) over several ‘years’ on seed germination was investigated with outdoor and laboratory experiments. Warm summer‐like temperatures (20 °C) were necessary for germination at subsequent cooler autumn‐like temperatures (greatest at 15 °C in G. nivalis and 10 °C in N. pseudonarcissus). As the warm temperature duration increased, so did germination at subsequent cooler temperatures; further germination occurred in subsequent ‘years’ at cooler temperatures following a second, and also third, warm period. Germination was significantly greater in darkness, particularly in G. nivalis. Dormancy increased with seed maturation period in G. nivalis, because seeds extracted from green capsules germinated more readily than those from yellow capsules. Desiccation increased dormancy in an increasing proportion of N. pseudonarcissus seeds the later they were dried in ‘summer’. Seed viability was only slightly reduced by desiccation in N. pseudonarcissus, but was poor and variable in G. nivalis. Shoot formation occurred both at the temperature at which germination was greatest and also if 5 °C cooler. In summary, continuous hydration of seeds of both species during warm summer‐like temperatures results in the gradual release of seed dormancy; thereafter, darkness and cooler temperatures promote germination. Cold temperatures, increased seed maturity (G. nivalis) and desiccation (N. pseudonarcissus) increase dormancy, and light inhibits germination. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 246–262.     

THE GERMINATION STRATEGY OF OLDFIELD ASTER (ASTER PILOSUS)

At maturity in November, a high percentage of Aster pilosus Willd. seeds germinated in light at high temperatures (30/15, 35/20 and 40/25 C). Stratification during winter lowered the temperature requirement for germination, and high percentages of germination were obtained in light at 15/6 and 20/10 C., as well as at 30/15, 35/20 and 40/25 C. Stratification in darkness was completely ineffective, but stratification in light was partially effective in overcoming the light requirement for germination. Inability of seeds to germinate at low temperatures prevents germination after dispersal in late autumn and winter, when freezing temperatures could kill the seedlings. The lowering of the temperature requirement for germination during winter stratification allows the seeds to germinate and the resulting vegetative rosettes to become well established before the onset of the periodic summer droughts that occur in habitats occupied by A. pilosus.     

Soil seed bank,fire season,and temporal patterns of germination in a seeder-dominated Mediterranean shrubland

Soil seed banks play a major role in the post-fire regeneration of Mediterranean shrublands. They vary throughout the year in species composition, abundance, and readiness to germinate. After fire, germination occurs mainly during the following fall to spring. Time of germination can determine recruitment success. It is unclear what factors control post-fire germination and its timing. We tested the effects of season and fire on the readily germinable soil seed bank of a seeder-dominated shrubland. Plots were burned early and late in the summer season (ES, LS). Soil samples were collected before and after fire, and germinated in a chamber simulating successively autumn, winter, and spring conditions. Samples were kept moistened at all times. Fire intensity was similar between ES and LS. Several species of Cistus and herbs, mostly annuals, were dominant. Most germination occurred during the simulated-autumn period, with little subsequent germination during the following two periods. Germination speed (T ₅₀) during simulated-autumn was similar for shrubs and herbs, and independent of season or fire. Germination was lower for two shrubs (Rosmarinus officinalis, Cistus salvifolius) and higher for herbaceous dicots in LS than in ES soils. Fire reduced monocots and enhanced Cistus. Germination period significantly interacted with fire and season in some groups or species, altering the simulated-autumn germination peak. We demonstrate that the seed bank can germinate swiftly under simulated-autumn conditions. Hence, water availability is the main controlling factor of germination. Fire season differentially affected some species or groups, and could affect the post-fire regeneration.     

Evidence for physiological seed dormancy cycling in the woody shrub Asterolasia buxifolia and its ecological significance in fire‐prone systems

• Dormancy cycling is a key mechanism that contributes to the maintenance of long‐term persistent soil seed banks, but has not been recorded in long‐lived woody shrub species from fire‐prone environments. Such species rely on seed banks and dormancy break as important processes for post‐fire recruitment and recovery.
• We used germination experiments with smoke treatments on fresh seeds and those buried for 1 year (retrieved in spring) and 1.5 years (retrieved the following late autumn) to investigate whether Asterolasia buxifolia, a shrub from fire‐prone south‐eastern Australia with physiologically dormant seeds, exhibited dormancy cycling.
• All seeds had an obligation for winter seasonal temperatures and smoke to promote germination, even after ageing in the soil. A high proportion of germination was recorded from fresh seeds. but germination after the first retrieval was significantly lower, despite high seed viability. After the second retrieval, germination returned to the initial level. This indicates a pattern of annual dormancy cycling; one of the few observations, to our knowledge, for a perennial species. Additionally, A. buxifolia’s winter temperature and smoke requirements did not change over time, highlighting the potential for seeds to remain conditionally dormant (i.e. restricted to a narrow range of germination conditions) for long periods.
• For physiologically dormant species, such as A. buxifolia, we conclude that dormancy cycling is an important driver of successful regeneration, allowing seed bank persistence, sometimes for decades, during fire‐free periods unsuitable for successful recruitment, while ensuring that a large proportion of seeds are available for recruitment when a fire occurs.

     

Effects of temperature,light and gibberellic acid on the germination of seeds of 43 species native to Western Australia

Abstract. Species native to the southwest of Western Australia, representing a range of plant families, life-history strategies, fire-response syndromes, seed-store types and seed weights, were tested for viability using tetrazolium chloride and for germination under combinations of constant temperatures of 15 °C or 23 °C, constantly dark or 12 h diurnal whitelight conditions, and with, or without, addition of gibberellic acid (GA₃, 50 mg/l). Species previously known to require a heat-shock treatment to overcome dormancy due to an impervious testa were pre-treated prior to imposition of temperature, light and GA₃ conditions. The test environmental conditions related to differences between winter and autumn temperatures and surface and buried seed germination positions of post-fire habitats. The viability of the selection of native Western Australian species ranged from 0 to 100 %, averaging 71 %. For all taxa, no combination of temperature, light and gibberellic acid treatment induced all viable seeds to germinate. The greatest percentage germination achieved in any combination of treatments averaged 71 % of all viable seeds for all species. Larger seeds (> 10 mg seed weight) tended to have greater viability percentages, but no overall patterns of viability or germinability were attributable to seed storage syndrome, strategy of fire recovery response or life-form type. Germination of most of the dominant tree representatives (Eucalyptus calophylla, E. diversicolor, E. erythrocorys, E. gomphocephala, and E. patens) was indifferent to the trial conditions of temperature, light and GA₃. However, Eucalyptus marginata showed reduced germination in the light, which was overcome with GA₃. GA₃ also overcame the inhibition resulting from exposure to light in some understorey species (e.g. Allocasuarina campestris, Regelia ciliata, Xanthorrhoea gracilis and X. preissii). Germination of many hard-seeded, understorey shrub and herbaceous perennial species, especially those with small (< 10 mg) seeds (e.g. Bossiaea ornata, B. aquifolium and Acacia drummondii ssp. candolleana) was greater at the lower trial temperature and in the dark. Some large (> 10 mg) seeded, understorey species (e.g. Acacia extensa, Kennedia coccinea, K. prostrata, Hovea trisperma and Hardenbergia comptoniana) germinated in high percentages in both temperatures, but maximum germination percentages still tended to be at 15 °C. Large-seeded species were less sensitive to exposure to light compared to the smaller seeded species. The largest seeded species tested, Paraserianthes lophantha, germinated best in the warmer incubation temperature and in the light. The ecological significance of the tests would be that species which have seed dormancy mechanisms capable of delaying germination until the cool temperature, winter rainy period of this mediterranean-type climate would be more likely to survive than if germination followed summer rain showers or the first, intermittent rains of autumn. Burial of seeds becomes more important if germination occurs when rains first begin as this period has less available soil moisture and temperatures are high. Also survival of seedlings could be enhanced if germination of seed was restricted to the positions protected from high light, higher temperatures and lower soil moisture by the presence of a forest canopy. Therefore, seeds which have an ability to sense the presence of a previous fire in the habitat, conditions in light environment and appropriate temperature level have an adaptive advantage to time emergence to situations of time and space where survival is maximized. Variation in viability and germination percentages were apparent in some cases where more than one seed collection of available for testing, indicating that further aspects, such as seed age, maturity at collection, storage conditions and depth of seed dormancy, remain to be considered.     

Environmental factors affecting seed germination of gray birch (Betula populifolia) collected from abandoned anthracite coal mine spoils in northeast Pennsylvania

Germination of gray birch (Betula populifolia) seed collected from anthracite mine spoils in northeastern Pennsylvania was studied. Environmental conditions of the spoil banks are such that high mortality may occur at seed and germination stages because of low moisture availability and thermal stress. The mine spoil banks are harsh environments with respect to key seed germination factors: percent soil moisture as low as 1.8% and soil surface temperatures reaching 59°C. In the field, gray birch typically germinated in mid-April prior to severe environmental stress. Trends in germination success were inversely related to rising soil temperature and decreasing soil moisture availability. Although seeds were capable of survival and germination under laboratory conditions of constant temperatures in excess of 55°C, dramatic decline in germination was observed under fluctuating temperature regimes likely to be experienced in the field. No germinations occurred under fluctuating temperatures in excess of 30°C. Germinations in the field were seen to end after mid-June when substrate temperatures exceeded 30°C.     

The interaction of temperature, water availability and fire cues regulates seed germination in a fire-prone landscape

Ambient temperature and water availability regulate seasonal timing of germination. In fire-prone landscapes, the role of fire-related cues in affecting the range of temperatures and water potentials (ψs) across which germination can occur is poorly known, especially in non-Mediterranean landscapes. We examined interactive effects of temperature (15 or 25°C), ψ (0 to −0.9 MPa), and fire-related cues (heat and smoke) on germination for seeds of three shrub species from fire-prone southeastern Australia. Incubation temperature affected germination of untreated seeds of Kunzea ambigua and Kunzea capitata (Myrtaceae) (reduction at 25°C), but germination was uniformly low in Epacris obtusifolia (Ericaceae). Decreasing ψ reduced germination across both incubation temperatures. Fire cues increased germination at both incubation temperatures and across ψs, although in Kunzea the increase was smaller and occurred over a narrower range of ψs at 25°C. Hydrotime analysis suggested that fire cues reduced the amount of water necessary for germination of Kunzea seeds. Post-fire germination of the three study species may occur during the warm season, although it is reduced and confined to wet periods for the two Kunzea species. Warm season germination of the study species is consistent with a trade-off between the increased risk of failure of a cohort of seedlings, and benefits of early establishment of a cohort that may survive in an environment with aseasonal rainfall.